Chemical Research

on Plant Growth
A translation of Théodore de Saussure’s
Recherches chimiques sur la Végétation

Translated, with an Introduction,

by Jane F. Hill
Chemical Research on Plant Growth
Portrait of Théodore de Saussure. On the table at his side is a “recip-
ient”, or receptacle, that encloses a plant; this was an experimental
set-up that he used for studying gas exchange between plants and
atmosphere of varying composition. (Image courtesy of Bibliothèque
de Genève, Centre d’iconographie genevoise)
Jane F. Hill
Translator

Chemical Research on Plant

Growth
A translation of Théodore de Saussure’s
Recherches chimiques sur la Végétation

2123
Translator
Jane F. Hill
Bethesda, Maryland
USA

ISBN 978-1-4614-4135-9 ISBN 978-1-4614-4136-6 (eBook)

DOI 10.1007/978-1-4614-4136-6
Springer New York Heidelberg Dordrecht London
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Contents

Foreword . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii

Preface to the Original de Saussure Book . . . . . . . . . . . . . . . . . . . . . . . . . . . . ix

Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi

Translator’s Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xiii

Plant Nutrition Concepts Before the Pioneering Research on
Photosynthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xiv
Contributions to Knowledge of Plant Nutrition and Physiology by the
Photosynthesis Pioneers who Preceded Théodore de Saussure . . . . . . . xv
De Saussure Provides Extensive Quantitative Data to Address the
Important Questions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xxi
De Saussure’s Work Enables Completion of the Overall Chemical
Equation for Photosynthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xxvii
Critical Reception of Recherches chimiques sur la Végétation, its
Legacy, and Historical Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xxvii
Some Errors and Misinterpretations in Recherches chimiques sur la
Végétation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xxxi
The Life and Scientific Background of Théodore de Saussure . . . . . . . . . . xxxii
References for the Translator’s Introduction . . . . . . . . . . . . . . . . . . . . . . . . . xxxix

A Note on the Translation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xlix

Contents of Original de Saussure Book . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lv

De Saussure’s Plate of Figures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lix

Appendices . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
Appendix 1: Conversions of Units of Measurement . . . . . . . . . . . . . . . . . . . 161
Appendix 2: Glossary of Terms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163

v
vi Contents

Appendix 3: Bibliography of References Cited by de Saussure

(as expanded by the translator) . . . . . . . . . . . . . . . . . . . . . . . . . 170
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183

Index of Authors de Saussure Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189

About the Author and the Translator . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 191

Foreword

Théodore de Saussure was the last of the early pioneers of photosynthesis research.
He filled in the gaps in the basic understanding of photosynthesis and presented the
first concise summary of the field in his 1804 book Recherches chimiques sur la
Végétation, which is translated into English here for the first time.
De Saussure’s work was built on the achievements of his predecessors in plant-
nutrition research. In the early 17th century, Jan van Helmont had shown that
plants obtain very little of their dry matter from the soil, contrary to the prevail-
ing view. Van Helmont concluded, instead, that plant dry matter is composed mostly
of “transmuted” water rather than of soil. In the early 18th century, Stephen Hales
demonstrated that “air” is fixed in plant matter, but, at the time he was working,
knowledge of the atmosphere was so limited that he was unable to distinguish one
kind of gas from another. In the early 1770s, Joseph Priestley discovered the funda-
mental fact that plants and animals exist in an interdependent relationship mediated
by gases: Plants “purify” air that has been “vitiated”, or corrupted, by the breathing
of animals or burning of candles, and that animals, in turn, thrive in the restored
air. In the last quarter of the 18th century, Antoine-Laurent Lavoisier and his col-
leagues made great strides in chemistry, which enabled rapid advances in knowledge
about photosynthesis and plant nutrition during this period. The discovery that the
atmosphere is not a homogeneous substance but instead is composed of a number of
gases was of great importance for this unfolding understanding of plant nutrition. At
about this same time, Jan Ingen-Housz established that light is necessary for plants
to thrive and to produce oxygen, and Jean Senebier provided the insight that carbon
dioxide is taken up in the production of oxygen by plants.
De Saussure showed that water is a component of plant dry matter (a fact that van
Helmont had inferred but not conclusively demonstrated), thus enabling completion
of the basic, overall equation of photosynthesis. De Saussure also showed that plant
carbon comes mainly from atmospheric carbon dioxide rather than organic carbon
in the soil. Further, he showed that plants obtain their minerals from the soil rather
than from “transmutations” of other substances by a vital force within the plant, and
that minerals are essential for plant growth.

vii
viii Foreword

Two German translations of de Saussure’s book appeared during the 19th century.
This first English translation now makes it readily accessible for English-speaking
students, researchers, and other readers who are not familiar with French or German.
Jane F. Hill, known to me for several years as an accomplished historian of photo-
synthesis research, has written a beautiful introduction for this book (see Translator’s
Introduction) that places de Saussure’s book in a historical context. This introduction,
as well as additional supplementary material in the book, incorporates editorial and
other suggestions that I made on draft versions of the manuscript. The introduction
provides an account of the gradual replacement of primitive ideas about plant nutri-
tion by the more accurate information supplied by the early pioneers, culminating in
the work of the last of these early researchers, de Saussure. The fate of the book is
also addressed, beginning with its positive initial reception, which was followed by
neglect of its fundamental conclusions until their revival by Jean-Baptiste Boussin-
gault and Justus von Liebig in the mid-19th century. The introduction describes how,
despite some errors due to the relatively primitive state of knowledge and experimen-
tal methodology in de Saussure’s day, the book is now widely recognized as a classic
and a basic document in the development of the understanding of photosynthesis and
plant nutrition. Finally, the life and scientific career of de Saussure are traced in detail.
In Chap. 10, Jane F. Hill presents succinct summaries of each of the nine chapters
of de Saussure’s book; in addition, she has provided at the end of the book three
very important appendices, which provide conversions of units of measurements, a
glossary of terms, and fleshed-out bibliography for most of the approximately 100
authors cited by de Saussure in his text.
I end this Foreword by making a reference to the Preface (pp. xxiii–xxvi), and
three editorials (pp. 5–35) in a 2005 book Discoveries in Photosynthesis, edited by
Govindjee, J. T. Beatty, H. Gest, and J. F. Allen, Springer, Dordrecht; they provide
a link to the many discoveries and discoverers in photosynthesis research since de
Saussure’s pioneering work.

Govindjee
Department of Plant Biology
University of Illinois at Urbana-Champaign
265 Morrill Hall, 505 South Goodwin Avenue
Urbana, IL 61801, USA
E-mail: [email protected]
Preface to the Original de Saussure Book

The object of the research that occupies me in this work is the influence of water,
air, and humus on plant growth. I do not pretend, however, to fathom all aspects
of this immense subject. I address the questions that can be decided by experiment
and forgo those that can give rise only to conjectures. In natural history, the facts
alone lead to the truth. In following this route, one is forced to acknowledge that
the discovery of the means used by nature for the development of plants and for the
composition of their substance must remain beyond our grasp for a long time. The
solution of these questions often requires information that we do not have. It requires
exact procedures for the analysis of plants and a full knowledge of their organization.
The functions of water and gases in the nutrition of plants, and the changes that
plants produce in their atmosphere, are the subjects that I have investigated most
deeply. The observations of Priestley, Senebier, and Ingen-Housz opened the road
that I have traveled, but they did not attain the goal that I set myself. If imagination
has sometimes filled the gaps that they left, it is by conjectures whose inadequacy has
always been shown by their unintelligibility and inconsistency. For my eudiometric
tests, I used either potassium sulfide [“sulfide of potash”] or phosphorus.1 These
procedures allowed me to achieve in my analyses an accuracy that the nitrous gas
eudiometer used by my predecessors could not attain.

1
In this work, when I have given readings of the phosphorus eudiometer, they have always been
cleared of the error that nitrogen gas can introduce through the expansion it undergoes in dissolving
the phosphorus. Berthollet determined precisely the correction that this expansion requires. There
is, however, a case in which this correction should be negligible: it is when the condensation of
oxygen gas is observed at the precise moment at which it has reached its maximum, or when the
phosphorus ceases to give off white vapors. In this operation, the nitrogen gas begins to expand only
when it contains no more oxygen gas. In the first moments of the disappearance of the oxygen gas,
the expansion of the nitrogen gas is undetectable. The expansion usually only reaches its extreme
limit 24 hours after the complete absorption of the oxygen gas.
For these experiments, I used rapid combustion and the bent tube indicated by Giobert (Analyse des
Eaux de Vaudier). I tilt the eudiometer when the phosphorus is melting, so that the phosphorus flows
and spreads over the full length of the tube. With this procedure, the analysis of air is completed in
less than half an hour, and at this time needs no correction.

ix
x Preface to the Original de Saussure Book

My research leads me to conclude that water and air contribute more to the for-
mation of the dry matter of plants growing in a fertile soil than does even the humus
matter that they absorb in aqueous solution through their roots.
I also focus on a subject that has given rise only to hypotheses: the origin of
the ash of plants. I investigate, through numerous experiments, the principles by
which these ashes vary, in both amount and composition, depending on the season
and the nature of the plants and their different parts. This work brought me several
new observations, which show that all the questions that I have just stated can be
resolved without attributing to vegetation creative forces and transmutations contrary
to known observations.
In experiments on plant growth, so many diverse and unforeseen factors tend
to influence the results that one should never fail to report all the accompanying
circumstances. The details I provide on this subject will serve to determine the degree
of confidence that can be placed in my research. They will prevent contradictions that
arise from differences in method. They will explain errors that I cannot flatter myself
to have avoided in a series of long, difficult experiments, whose results are perhaps
applicable only to the species of plants that I examined. The path that I set upon is
without doubt dry and fatiguing, but if one considers that the goal towards which it
is directed is the improvement of agriculture, one will bear with its difficulties and
excuse its shortcomings.
Acknowledgments

The translator gratefully acknowledges her deep indebtedness to a number of indi-

viduals who offered encouragement and support during the course of this project and
who consented to read critically, review, or proofread the entire manuscript or parts
of it. First, I want especially to thank Govindjee, Professor Emeritus, Department
of Plant Biology, University of Illinois at Urbana-Champaign, Urbana, IL, USA,
without whose encouragement and help this book would not have been possible, and
whose advice, editing, and insightful comments were invaluable.
I am also indebted to Mary Ellen Bowden and Robert Kenworthy, of the Chemical
Heritage Foundation, Philadelphia, PA, USA, for their generous assistance with
chemical terminology.
Others to whom I am deeply grateful include Marilyn Schiff for her unfailing
encouragement and for advice on French usage; and Anita Baker-Blocker, Dennis
G. Baker, and Barbara Wolanin for critical readings and comments on various parts
of the manuscript.
I could never have completed this project without the tremendous encouragement
and support of my husband, William A. Hill, whose tireless and careful proofreading
of drafts was of immeasurable value and is deeply appreciated.
The staffs of various libraries were of great assistance in helping track down ref-
erence works, especially older works, of limited accessibility, cited by Théodore de
Saussure in his book. I am particularly indebted to Eric Frazier, Reference Librar-
ian, Rare Book and Special Collections Division, Library of Congress, Washington,
D.C., USA, for generous help in finding many of these obscure references. Staffs
at a number of other libraries also provided valuable assistance: National Library
of Medicine, Bethesda, Maryland, USA; American University Library, Washington,
D.C., USA; United States Department of Agriculture, National Agricultural Library,
Beltsville, Maryland, USA; Public Library System of Montgomery County, Mary-
land, USA; Bibliothèque de Genève, Geneva, Switzerland; Bibliothèque nationale
de France, Paris, France; and Biblioteca Accademia delle Scienze, Turin, Italy.

xi
Translator’s Introduction

Summary: In his 1804 book, Recherches chimiques sur la Végétation, Théodore de

Saussure amalgamated his own research results with the findings of earlier investi-
gators of photosynthesis to produce the first concise summation of the basic facts
of plant nutrition. Important advances that he contributed to the overall knowledge
in the field were that water is incorporated into the dry matter of plants; that plant
carbon is derived from the carbon dioxide of the air, not from humus or soil; that the
minerals in plants are absorbed from the soil, not created by a vital force; and that
minerals are essential to plant growth.
In an introduction written by the translator, de Saussure’s book is placed in his-
torical context and details of his life are presented. In addition, the translator has
provided a glossary of terms and a list of de Saussure’s more than 100 citations, with
fuller reference details than were given in the original book.

Key words: Théodore de Saussure; photosynthesis; plant nutrition; history of sci-

ence; early pioneers of photosynthesis research; plant mineral nutrition; roles of car-
bon dioxide, water, and minerals in plant growth and development; chemical revolu-
tion; agricultural chemistry; 18th - and 19th -century agricultural science, agron-
omy, and soil science; “humus” theory of plant nutrition; historical importance of
Recherches chimiques sur la Végétation; reference details for literature cited by de
Saussure; chemical equation of photosynthesis

Between the early 17th and early 19th centuries, pioneering researchers unraveled
the fundamentals of photosynthesis and plant nutrition. The last of these pioneers,
plant physiologist Théodore de Saussure (1767–1845), of Geneva, Switzerland, con-
solidated his own findings with the contributions of earlier researchers to provide
the first concise, unified concept of photosynthesis and the physiology of plant nu-
trition in his book, Recherches chimiques sur la Végétation (de Saussure 1804a). De
Saussure’s most important findings were that plants incorporate water into their dry
substance; that they obtain their carbon from the atmosphere, not from humus in
the soil, as many others thought; and that their minerals come from the soil, not
from “transmutations” of other elements within the plants, as was widely believed
at the time. He also showed that minerals are essential to growth, and he performed
the most detailed analyses of the minerals in plant ash to that date. De Saussure’s

xiii
xiv Translator’s Introduction

demonstration that plants incorporate water enabled completion of the basic, overall
chemical equation of photosynthesis, which states that, from the simple, inorganic
substances water and carbon dioxide, plants, using the energy of sunlight, produce
fixed carbon, with the liberation of oxygen gas.
Recherches chimiques sur la Végétation met with initial acclaim and was soon
translated into German (de Saussure 1805), but the book’s fundamental conclusions
were then ignored for the next several decades. In the mid-19th century, the book’s
worth was re-established, ensuring de Saussure an enduring reputation.
Throughout the 19th century, even during the decades of eclipse, chemists and
plant scientists, including some of the most illustrious in the field, cited the book
extensively for various purposes (e.g., Thaer 1809–1812; Davy 1813; Keith 1816;
Liebig 1840; Schleiden 1842; Boussingault 1860–1891; von Sachs 1875; Pfeffer
1881; Aikman 1894). Late in that century, after the book’s conclusions had once again
been recognized, it was translated into German a second time (de Saussure 1890).
During the 20th and early 21st centuries, English-language writers on photosynthesis
and other aspects of plant science, agriculture, and the history of chemistry praised the
book highly, and many of them quoted passages that they had translated into English
(e.g., Harvey-Gibson 1919; Sherman 1933; Browne 1944; Reed 1949; Nash 1952;
Gabriel and Fogel 1955; Partington 1962; Aulie 1970; Morton 1981; Arnon 1991;
Magiels 2010). A facsimile edition of the original French text was published in 1957,
and a second facsimile edition appeared in 2010. Despite the book’s widely acknowl-
edged importance, however, the full text has not been available in English until now.

Plant Nutrition Concepts Before the Pioneering Research on

Photosynthesis1

Advances in the knowledge of both chemistry and plant nutrition were long hampered
by vague, untested assumptions, such as the idea that one substance could be “trans-
muted” into another. This concept had been invoked since the alchemical Middle
Ages, not only in connection with the attempt to change base metals into gold, but
also as an all-purpose explanation for a wide range of natural phenomena. Scientific
advances were also hindered by the doctrine, which had persisted from the time of
Aristotle (384–322 B.C.), that all matter is composed of just four elements: earth,
air, fire, and water. In addition, a vital force was thought to control processes within
organisms, operating differently from the principles that govern nonliving matter.
With knowledge so limited, plants were understandably thought to obtain all their
nutrients from the soil through their roots, which were seen as somewhat analogous to
animal mouths. This view persisted into the late 18th century in many quarters, even
as experimental evidence was accumulating showing that air and possibly water,
in addition to soil, contributed to plant nutrition. English agricultural writer and
farmer Jethro Tull (1674–1741), in his widely read book The New Horse-Houghing

1
The term “photosynthesis” did not originate until the late 19th century (see Gest 2002).
Translator’s Introduction xv

Husbandry (1731 and many subsequent editions), popularized this view, with his
advocacy of an agricultural system based on pulverizing the soil to such a fine powder
that the particles could easily be taken in by plant roots.
In a detailed discussion of early ideas about plant nutrition, Browne (1944, p. 256)
credits German physician and chemist Georg Ernst Stahl2 (1660–1734) with making
the first attempt at a scientific explanation of soil fertility, especially the benefits
of organic manures. Stahl attributed soil fertility to an oily or fatty principle (a
“phlogistic fatness”). Similarly, Külbel (1741) assumed that the main source of
plant fertility was a fatty magma (“magma unguinosum”) occurring in humus. Other
promulgators of this concept of soil fertility included Home (1757), Wallerius (1761,
per Browne 1944, p. 256), and Dundonald (1795).
The idea soon arose that humus had to be brought into solution by oxidation and
fermentation before it could be used by plants (Browne 1944, p. 256), and out of this
idea grew the “humus theory” of plant nutrition, which held that plants obtain their
carbon from water-soluble organic matter in the soil (Waksman 1936; Russell 1976;
Feller and Boulaine 1987; Boulaine 1994; van der Ploeg et al. 1999; Feller and Man-
lay 2001). Although some fertilizers, such as salts and lime, were acknowledged to
promote plant growth, they were thought only to aid the decomposition of humus and
the dissolution of organic matter in the soil water (van der Ploeg et al. 1999, p. 1057).
Often intertwined with the humus theory was the idea that a vital force in the plant
created the minerals that had long been detected in plant ash. These minerals were
thought to originate by transmutation of other constituents of plant matter (van der
Ploeg et al. 1999, p. 1057), not from the external environment.

Contributions to Knowledge of Plant Nutrition

and Physiology by the Photosynthesis Pioneers who Preceded
Théodore de Saussure3

One hundred years before Külbel (1741) posited a fatty magma as the main source
of plant fertility, Flemish physician Jan van Helmont4 (1579–1644) took the first

2
Stahl is best remembered for his advocacy of the phlogiston theory of chemistry, which held sway
during the 18th century before it was upended by the chemical revolution late in that century.
3
Publications focusing on the work of the early pioneers of research on photosynthesis and plant
nutrition include Rabinowitch (1945, Vol. 1, Chap. 1); Nash (1952); Rabinowitch and Govindjee
(1969, see Introduction); Hill (1970); Rabinowitch (1971); Egerton (2008); Magiels (2010); and
Hill (2012). The early pioneers are treated within the larger context of the history of chemistry by
Partington (1957, 1962) and Ihde (1964); within the context of agricultural chemistry by Browne
(1944); and within the context of soil science by Russell (1976). Overviews of photosynthesis
research that go beyond the era of the early pioneers, up to various later dates, include von Sachs
(1875, 1890 English translation); Spoehr (1919); Sherman (1933); Morton (1981); Höxtermann
(1992); Huzisige and Ke (1993); Govindjee and Gest (2002); Govindjee et al. (2003); Govindjee
et al. (2004); Govindjee and Krogmann (2004); Morton (2008); and Nickelsen (2010).
4
For more on van Helmont, see Redgrove and Redgrove (1922), Harvey (1929), Pagel (1972), and
Newman and Principe (2002), among many other sources.
xvi Translator’s Introduction

major step towards a broader, more accurate view of plant nutrition. He was the first
to demonstrate convincingly that soil could not be the only source of plant nutrients.
He planted a 5-pound willow tree in an earthen pot in which he had placed 200 pounds
of oven-dried soil, and covered this vessel with a perforated lead shield to exclude
foreign matter. He added only water to the pot for 5 years. After removing the tree
from the soil and re-weighing it (without the leaves that had been shed during the
5-year period), and also weighing the re-dried soil, he found that the tree had gained
about 164 pounds and the soil had lost only 2 ounces. From these results, van Helmont
concluded that the solid substance of plants is formed not of soil, but instead entirely
of water (one of the four Aristotelian elements) that has been transmuted into solid
matter, or “earth” (another of the Aristotelian elements). Van Helmont’s experiment
was briefly reported in a posthumously published collection of his writings (van
Helmont 1648). Although he was correct in his conclusion that water is an important
component of plant dry matter, he did not prove a role for water (an achievement
belonging to Théodore de Saussure, 150 years later), and he mistakenly ruled out
soil and the atmosphere as other sources of plant nutrients. He did establish, though,
how little of their solid matter plants derive from the soil (Hill 1970).
The first experimental evidence showing that soil does play a role in plant nutrition
was provided at the end of the 17th century, by English naturalist and geologist
John Woodward (1665–1728). Woodward (1699) showed that plants grown in water
containing impurities or soil gained more weight than did plants grown in distilled
water. He concluded that the impurities or soil were the essential nutrient and that
water was merely a vehicle carrying them into the plant.
Also in the late 17th century came evidence that air might be involved in plant
nutrition, when the first-ever comprehensive studies of plant anatomy were under-
taken, by Malpighi (1675–1679) and Grew (1682). They found that leaves contain
minute pores (stomata) in their surfaces and have internal structural features that
appeared capable of functioning as air ducts. Then, in the early 18th century, the first
functional evidence was found for the involvement of gases in plant life: Stephen
Hales5 (1677–1761), an English physiologist, chemist, and cleric, reported in his
classic book Vegetable Staticks (1727) that “air” becomes “fixt” (solidified) in plant
dry matter and thereby loses its “elastick” (gaseous) state. Because Hales accepted
the prevailing doctrine that all matter is composed of the four Aristotelian elements,
however, he did not distinguish one kind of gas from another.
During the late 18th century, the understanding of photosynthesis progressed
rapidly, as the chemical revolution yielded both new chemical knowledge and im-
proved experimental techniques that could be applied to the study of plant nutrition
and physiology. In turn, research in plant nutrition contributed to knowledge of
chemistry (see Partington 1962; Ihde 1964). Of the many European natural philoso-
phers who contributed to the dramatic advances in chemistry during that period, the

5
For biographies of Hales, see Clark-Kennedy (1929) and Allan and Schofield (1980).
Translator’s Introduction xvii

French researchers and theorists, led by Antoine-Laurent Lavoisier6 (1743–1794),

stand out. Although Lavoisier did not advance the understanding of photosynthesis
directly, his work helped make rapid progress in that field possible. For example,
in 1770, Lavoisier took an early step towards overthrowing the doctrine of trans-
mutation when he showed, through an experiment with water in a vessel, that the
water was not transmuted into earth, or dry substance. He thereby cast doubt on van
Helmont’s conclusion that plant dry matter consists of transmuted water. Lavoisier
also played a key role in establishing that there is a broad array of chemical ele-
ments, which are definable much more precisely than the four, all-encompassing,
Aristotelian elements. To qualify as an element, a substance had to be shown to persist
through complex chemical reactions and not be decomposed into more fundamental
substances.
Another chemical advance important for photosynthesis research was the isolation
of individual gases and the determination of their composition. This work was made
possible by improved methods of catching gases, transferring them from vessel to
vessel, and studying their properties (Rabinowitch 1971). Much of this work in
“pneumatic” chemistry was done by British researchers during the third quarter
of the 18th century (see Ihde 1964, Chap. 1). The work was greatly aided by the
development of a piece of equipment, the pneumatic trough, which consisted of a
receiver, or vessel, inverted in a bath of water or mercury to trap gases.7
The first gas to be isolated was carbon dioxide. It had originally been discovered
by van Helmont (1648), who did not appreciate its significance. The gas was re-
discovered in 1754 by Scottish physician Joseph Black (1728–1799), who called it
“fixed air” (1756). Hydrogen was discovered by Henry Cavendish (1731–1810) in
1766, and nitrogen by three separate researchers in the early 1770s (see discussion
by Partington 1962, p. 222, pp. 263–265).
English chemist and theologian Joseph Priestley8 (1733–1804) broadened aware-
ness that the atmosphere has a functional role in living organisms, with his startling
discovery that plants “restore” air that has been “injured” by the burning of a candle
or the breathing of an animal (1772, p. 168). Priestley, however, could not explain
the relationship.
Priestley also developed a chemical technique, which he called the “nitrous air
test” (but which actually used nitric oxide), to measure the degree of “purity” of a
gas sample (1772, pp. 210–216).9 The nitrous air test came to be widely used by

6
Biographies and other treatments of Lavoisier include McKie (1952); Guerlac (1961, 1975);
Holmes (1985, 1998); Donovan (1993); Poirier (1996); and Bell (2005).
7
For a history and description of the pneumatic trough, see Badash (1964) and Parascandola and
Ihde (1969).
8
For a comprehensive treatment of Priestley, see the two-volume biography by Schofield (1997,
2004).
9
The nitrous air test was based on the reaction between oxygen and nitric oxide (NO), a colorless,
insoluble gas. The solubility of the reaction product, nitrogen dioxide (NO2 ), in water caused a
decrease in the volume of the gas phase and thus provided a measure of the amount of oxygen in
the original sample (see discussions by Conant 1950; Nash 1952; Partington 1962; Ihde 1964).
De Saussure used the nitrous air test in some of his work with gases and plants, although by
xviii Translator’s Introduction

18th-century chemists. Although Priestley did not realize it when he developed it, this
was a test for oxygen, a gas that he himself was to discover only two years later, in a
nonbiological context (see Priestley 1775). He named his new gas “dephlogisticated
air” (as distinguished from “phlogisticated,” or “bad,” air). Lavoisier (1778) renamed
the gas “oxygine”—i.e., oxygen.
The discovery of oxygen was the central event that enabled Lavoisier to over-
throw the older, phlogiston theory of chemistry, which had dominated through most
of the century (Conant 1950, p. 49), in favor of a new, more scientifically based, oxy-
gen chemistry. The phlogiston theory, promulgated by Stahl (see Partington 1957,
pp. 85–89), held that combustible materials contain an “inflammable principle,”
called “phlogiston,” which is transferred to the atmosphere upon burning or, in the
case of metals, upon calcination. Once the atmosphere had become “phlogisticated,”
or saturated with phlogiston, it could support no additional burning or calcination,
nor could it support animal life, because respiration was presumed to saturate the air
with phlogiston given off by slow burning within the animal body. In combustion and
respiration, then, the role of phlogiston was essentially the converse of the role that
oxygen was shown to play.10 Armed with the new knowledge of oxygen, Lavoisier
(1783) launched a vigorous attack on the phlogiston theory. He and his colleagues
(de Morveau et al. 1787) revised and modernized chemical terminology, removing
the phlogiston-associated language. Soon thereafter, Lavoisier set forth his new ap-
proach to chemistry in his pioneering book Traité élémentaire de Chimie (1789).
Most of the chemical elements recognized in this book are still accepted today.
Late-18th-century chemical discoveries that were especially important for the
understanding of plant and animal physiology included: the finding that fixed air
(carbon dioxide) is composed of carbon and oxygen (Lavoisier 1781); that water,
rather than being an element, is a compound of hydrogen and oxygen (see discus-
sion by Partington 1962, p. 344 ff); and that fixed air is different from “mephitic”
air (nitrogen) (see Partington 1962, p. 263). In physiology, animal respiration was
demonstrated to be a process of oxidation that provides heat for the body (Lavoisier
1777; Lavoisier and de la Place 1780).11
Lavoisier developed the technique of analyzing organic substances by combustion
(Holmes 1985, Chap. 9), and used it to analyze spirit of wine (alcohol), olive oil,
and wax (1784). He noted (1789, 1790 English translation, p. 123) that the “true”
constituent elements of plants are hydrogen, oxygen, and carbon.

the mid-1780s, this technique of “eudiometry,” or measurement of oxygen-gas content, had been
largely abandoned as imprecise and unreliable (Golinski 1992, p. 93). In his research on plants, de
Saussure preferred to use phosphorus or potassium sulfide in his eudiometric tests. For the history
of the development of eudiometric methods, see Benedict (1912), Golinski (1992, pp. 117–128),
and Magiels (2010, Chap. 4).
10
See Conant (1950) and Nash (1952) for detailed discussion of the phlogiston theory and its
upending by oxygen chemistry.
11
For details about Lavoisier’s findings in physiology, see Holmes (1985).
Translator’s Introduction xix

A major advance in the understanding of photosynthesis itself came in 1779,

when, during a single summer of research, Dutch physician Jan Ingen-Housz (1730–
1799) made the critical discovery that sunlight is essential for plants to thrive. Based
on this pivotal finding, Ingen-Housz has been credited by some (e.g., Gest 1997;
Magiels 2010) with the discovery of photosynthesis.12 Ingen-Housz (1779) reported
his results during the autumn immediately following his summer of work.
Ingen-Housz achieved his important insight about light partly because he used an
innovative experimental set-up. Priestley’s experiments were done on whole plants.
Ingen-Housz, however, made use of a 1754 observation by naturalist Charles Bonnet
(1720–1793) that bubbles form on leaves that are submerged in water. Bonnet had
not recognized their significance but, with the benefit of the increase in knowledge of
gases over the intervening years, Ingen-Housz was able to demonstrate that the bub-
bles consist of dephlogisticated air and that only illuminated leaves produce this gas.
Ingen-Housz had no idea of the identity of the starting material for the production of
dephlogisticated air, however, instead invoking a vague “transmutation,” excited by
light, within submerged leaves, and a different process—a withdrawal of phlogiston
from the air—by plants in aerial environments.
Ingen-Housz also showed that the non-green parts of plants, and the green parts
when not illuminated, produce fixed air. Thus he discovered respiration in plants, two
years after Lavoisier (1777) had reported it in animals. Priestley, in contrast, with his
use of whole plants, had difficulty obtaining clear results in his later experiments be-
cause his experimental material contained both non-green, carbon dioxide-producing
tissues and green, oxygen-producing ones and because he did not understand the
requirement for light in photosynthesis. Ingen-Housz noted that production of de-
phlogisticated air by plants during the day exceeded their production of fixed air
by night, so that overall the improvement of the atmosphere during the day far
outweighed its vitiation, or corruption, at night (1779, p. 47).
Swiss clergyman and naturalist Jean Senebier13 (1742–1809) contributed the next
major insight on photosynthesis. He repeated many of Ingen-Housz’s experiments,
with mostly similar results, and he did not always give Ingen-Housz due credit
(Magiels 2010), but Senebier (1782) was the first to establish that fixed air is con-
sumed in a plant’s production of dephlogisticated air. Senebier (1782, 1783) also
found that the amount of dephlogisticated air produced is roughly proportional to the
amount of fixed air available to the plant. (This was a line of research that de Saus-
sure was to expand upon and refine.) Further, Senebier (1782) pinpointed the green,
fleshy parts of leaves (the parenchyma) as the sites where fixed air is transformed
into dephlogisticated air.

12
Rabinowitch and Govindjee (1969, p. 7) conclude that each of the early pioneers made an invalu-
able contribution to the understanding of photosynthesis and that “there is fame enough to share
among them”.
13
Kottler (1973) provides a comprehensive historical description of the scientific contributions of
Senebier.
xx Translator’s Introduction

In contrast to Ingen-Housz, Senebier considered the production of fixed air by

plants to be an indication of stress or disease caused by the unnatural growing con-
ditions in experimental situations, rather than a natural process carried out by all
plants. This subject became a major area of contention between the two men.
During the final two decades of the 18th century, Ingen-Housz (1787, 1789, 1796)
and Senebier (1783, 1788, 1791, 1792, 1800) revised their views, speculated, and
quarreled about various aspects of plant metabolism.14 The two men agreed that
carbon has a nutritional role in plants: Senebier (1791, p. 164; and 1800, Vol. 3,
p. 151) stated that carbon combines with plant constituents, and Ingen-Housz (1796,
p. 4) concluded that plants derive some of their most essential substances—such as
their acids, oils, and mucilages—from the two elements that compose carbon dioxide
(or carbonic acid, as it was then called). They disagreed about the source of plant
carbon, however, right to the end: Ingen-Housz (1796) attacked the humus theory,
conjecturing that plants growing in the open air obtain almost all of their carbon from
the atmosphere, whereas Senebier (1800, Vol. 3, p. 148 ff) clung to the idea that they
obtain it through their roots, from carbon dioxide dissolved in the soil solution. The
quantitative data necessary to settle the matter would be supplied by de Saussure.
Senebier, in his writings on fixed air, initiated an error of interpretation that was
to be perpetuated by Ingen-Housz (1796), de Saussure, and many others, for some
150 years. This was the idea that the oxygen liberated in photosynthesis comes from
carbon dioxide rather than from water. This view seemed logical based on the close
equivalence between the volumes of carbon dioxide absorbed and oxygen released.
The idea is often attributed to Ingen-Housz (1796)—by, for example, Arnon (1971,
1991); and Raven, Evert and Eichhorn (1999)—but, although Ingen-Housz briefly
stated this view in 1796, he confusingly also mentioned water as a source of oxygen,
and, more importantly, it was not Ingen-Housz, but Senebier, who earlier had done
many, repetitious studies and analyses that suggested carbon dioxide as the precursor
of the oxygen released. Senebier’s very verbosity may have eclipsed his findings,
and Ingen-Housz, with his concise summary of the field in 1796, was more easily
read and comprehended (Hill 2012, p. 792). It is now known that water, not carbon
dioxide, is the source of the oxygen gas liberated in green-plant photosynthesis.
At century’s end, unresolved questions also remained about the source of plant
nitrogen and hydrogen. Lavoisier (1784) had found that plant matter contained abun-
dant hydrogen, and Berthollet (1785) had found nitrogen in both plants and animals,
but from what source did plants obtain these two important nutrients?15 Because
the most logical source of plant hydrogen was water, there was conjecture (by, e.g.,
Berthollet 1786; Senebier 1792) that plants obtain it from that liquid. Senebier’s
opinion (1792, p. 208) was based on his demonstration that plants transpire much

14
See Nash (1952) for a detailed, comparative account of the ideas of Ingen-Housz and Senebier
during this period.
15
Adequate concentrations of the four major nutrients in dry plant tissue are now considered to be,
by weight, 45 % carbon, 45 % oxygen, 6 % hydrogen, and 1.5 % nitrogen (Stout 1961, as cited by
Raven et al. 1999, p. 728).
Translator’s Introduction xxi

less water than they take in through their roots. The contribution of water to plant
dry matter was to be established by de Saussure.
For plant nitrogen, the most logical source seemed to be the large reservoir in
the atmosphere, which is approximately four-fifths nitrogen gas, by volume. Ingen-
Housz (1796) believed in an atmospheric source, but Senebier (1800, Vol. 3, p. 159)
thought that it came from the soil, along with the carbonic acid that he also believed
was absorbed from there. Senebier failed to establish that soil was the source, how-
ever, and de Saussure, although speculating correctly against gaseous nitrogen as the
source, did not settle the matter.
In addition, neither Ingen-Housz nor Senebier clarified the source of plant min-
erals. The idea that a vital force within plants formed these substances was still
prevalent when these two researchers completed their work. Senebier (1791, p. 252)
pondered the question of vital force versus environment, but did not provide an
answer.
Although both Ingen-Housz and Senebier clung to the phlogiston concept until
late in life, both of them eventually re-interpreted their findings, to a large extent,
in terms of Lavoisier’s chemical system. Senebier probably adopted the new system
about 1789 (Smeaton 1978); Priestley never did.16
Of the later publications by Senebier and Ingen-Housz, it was Ingen-Housz (1796)
that was the most succinct and that, according to Nash (1952, p. 105), provided “a
fair approximation to the conceptual scheme that we now hold to be correct. . . .
[A] view representing a closer approach to our present conceptual scheme was not
proposed until 1804” (in de Saussure’s book). Ingen-Housz (1796), in an appendix
to an obscure British government publication, presents the first formulation of plant
nutrition and the photosynthetic process in accordance with the new chemical theory
(Rabinowitch 1945). Although not entirely consistent in his terminology, Ingen-
Housz mostly used “carbonic acid” rather than “fixed air,” and “oxygen” or “vital
air” rather than “dephlogisticated air.” He used “carbon,” but also sometimes “coal,”
for the element. His scheme, however, included only brief mention of the idea of the
decomposition of water, and he paid scant attention to a possible nutritional role for
minerals or to the question of how plants obtained minerals.
Thus, at the dawn of the 19th century, with the contributions of Priestley, Ingen-
Housz, and Senebier completed, quantitative data on photosynthesis and plant
nutrition were still incomplete and critical questions remained.

De Saussure Provides Extensive Quantitative Data

to Address the Important Questions

The position of the Swiss plant physiologist Théodore de Saussure in the history of
the early research on photosynthesis has been compared to that of Jan van Helmont,
as both were transitional figures: Van Helmont was an alchemist, but he addressed

16
See Holmes (2000), however, for the view that Priestley’s concept of phlogiston was only loosely
allied with older phlogiston theories.
xxii Translator’s Introduction

subjects that became major fields of study in the burst of chemical and plant physi-
ological research of the late 18th century. De Saussure, in turn, completed the basic
experimental work and provided a unified theoretical interpretation of the field; he
also opened the way for further experiment and advances in knowledge (Nash 1952,
p. 106).
Younger than his immediate predecessors, Théodore de Saussure did not have to
labor under the weight of the phlogiston theory, because, by the time of his research,
that theory had largely been replaced by Lavoisier’s new chemistry. In accepting the
new chemistry as a young man, Théodore was probably following the lead of his
father, the noted scientist Horace-Bénédict de Saussure (1740–1799), who as early
as 1788 adopted the new system and was probably the first scientist in Geneva to do
so (Smeaton 1978).
With his more modern outlook, Théodore de Saussure recognized from the start
that a facile recourse to the “transmutation” of one kind of substance into another as
an explanation of natural phenomena was no longer tenable. Instead, he approached
problem-solving analytically, designing experiments that were carefully targeted to
answer important physiological questions of the day. He followed Lavoisier’s lead
and used a balance-sheet approach, carefully recording his quantitative results. He
was thus able to build quantitatively on his predecessors’ qualitative observations.
Naturally a cautious man, not given to extravagant speculation as were his immediate
predecessors in the field, he stuck to facts as revealed by experiments.
In many of his experiments, de Saussure used whole plants, which he was careful
to maintain in a normal, healthy state. Unlike his predecessors, he did not simply
observe whether the plants thrived or wilted, but measured their weight gain, and
he did not merely determine whether the air “improved,” but measured the amounts
of its gaseous constituents. Also unlike his predecessors, he made liberal use of
controls, comparable samples for determining quantities of substances in the initial
plant material, and repetition of experiments with different plant species or plant
organs. He made errors, some due to the primitive experimental techniques of his
time, and others—of interpretation—due to the still-rudimentary state of knowledge.
Nevertheless, he was able, through his thorough and wide-ranging investigations, to
come close to identifying the sources of most of the major elements found in mature
plants, and to determine the paths by which plants obtained them.
Some of the important questions de Saussure addressed—and the conclusions that
he reached based on his experiments—were:
1. What is the quantitative relationship between the carbon dioxide taken in and the
oxygen released by green plants in the light?
De Saussure found that, under many conditions, there was a close equivalence
between these quantities, but that in general somewhat more carbon dioxide was
taken in than oxygen released.17 This led him to conclude that plants retain, in
their substance, a small amount of residual oxygen from the carbon dioxide. He

17
The finding that the carbon dioxide taken up exceeded the oxygen released may be attributable to
a slow reaction of oxygen with the chemicals in the particular eudiometric tests de Saussure used
(Morton 1981, p. 360, note 74).
Translator’s Introduction xxiii

often found, erroneously, that plants released a small amount of nitrogen gas to
the atmosphere.
De Saussure’s findings supported Ingen-Housz’s view that, in ordinary air and
with exposure to light, the uptake of oxygen and emission of carbon dioxide are,
in a healthy plant as a whole, invariably exceeded by the uptake of carbon dioxide
and emission of oxygen. These two processes (photosynthesis and respiration),
although directly opposed, seemed to be inextricably linked in plant metabolism.
2. Do plants obtain their carbon mainly from the soil or the atmosphere?
Using experimental plants grown with their roots in distilled water and their
shoots in the open air, de Saussure provided the first convincing demonstration
that plants assimilate carbon mainly from atmospheric sources. This was a major
blow to the humus theory. He dealt that theory a further blow by showing that
carbon is proportionally more abundant in the humus than in the plants whose
decomposition gave rise to the humus, due to the loss of hydrogen and oxygen,
in the form of water, from the humus. If plants were taking a large amount of
carbon from the soil, he reasoned, there should be a decrease, not an increase, in
the carbon in the humus. Further, he found that, in humus extracts, which are the
soluble part of the humus and the only part available to plants, there is too little
solid substance to account for the dry weight gained by plants that are growing
and developing in the humus. Further, he found that, in atmospheres to which
he had added a limited amount of extra carbon dioxide, plants in the sun gained
more in dry weight than they did in ordinary air.
3. Does some of the oxygen released by green plants in the light come from the
decomposition of water?
De Saussure evaluated other authors’ claims that some of the oxygen released
by plants comes from water rather than carbon dioxide. He generally found no
evidence for this. In the case of a plant growing in a normal day/night cycle, in an
oxygen-free atmosphere, the plant may release several times its own volume of
oxygen gas, but de Saussure did not consider this an indication of the breakdown
of water, with release of the water’s oxygen. Instead, he reasoned that the plant,
having had no contact with oxygen gas during its early stages of growth, formed
carbon dioxide entirely from the carbon and oxygen of its own substance and then
decomposed it. In contrast, this same plant, at least if it was not fleshy and was
growing in an atmosphere that already contained oxygen, did not form carbon
dioxide entirely from its own substance but instead combined its carbon with
the oxygen in the atmosphere. The plant’s decomposition of this carbon dioxide
simply returned to the atmosphere the free oxygen that had been there at the start.
Because he found no net increase in atmospheric oxygen content under these
conditions, he concluded that plants did not add oxygen to the atmosphere by a
direct decomposition of water.
4. Does atmospheric oxygen combine directly with plant constituents other than
carbon?
De Saussure found that atmospheric oxygen generally reacts only with the carbon
of the plant, forming carbon dioxide gas. These findings countered some previous
xxiv Translator’s Introduction

researchers’ assertions, such as the claim that oxygen combines directly with
substances in a germinating seed to produce sugar. He based this conclusion on
his finding of a near-equivalence between the volumes of oxygen taken up and
carbon dioxide released, both in germinating seeds and in more-developed plants.
Leaves exposed to an alternating day/night regime, in a receptacle filled with
ordinary air, consumed oxygen at night but restored it by an approximately equal
volume of carbon dioxide during the day. He also found that atmospheric oxygen
gas is not fixed in the tissues of dead or fermenting plants or plant substances
(such as wine that is fermenting to vinegar), during the early stages, nor does it
combine with plant hydrogen to form water. He concluded that the sole role of
oxygen in these stages is to combine with plant carbon, yielding carbon dioxide
gas.
5. Is the presence of oxygen gas necessary for plants to break down carbon dioxide
and to grow?
Plants, de Saussure found, need oxygen gas in their atmospheres in order to be
able to break down carbon dioxide (i.e., to photosynthesize) and to grow. Plants
can survive in environments lacking oxygen gas only if they are able to produce
free oxygen through the decomposition of carbon dioxide that they form entirely
from their own tissues. Even plants that can survive in an atmosphere of pure
nitrogen gas, however, die there if carbon dioxide is added to that atmosphere in
concentrations that would have allowed plant growth in an ordinary atmosphere.
(He also determined that plants do not absorb nitrogen, hydrogen, or carbon
monoxide gases.)
Like Ingen-Housz, de Saussure concluded that the union of atmospheric oxygen
with plant carbon (i.e., respiration) is a normal plant process and not, as Senebier
had thought, an indication of unnatural growing conditions or of plant stress
or disease. He considered respiration to perform the same function in plants as
in animals, that is, to bring about a release of heat, through the combination
of atmospheric oxygen with the carbon of the organism. He noted that plants’
release of heat often went undetected. De Saussure was unable to develop as
clear an interpretation of the metabolic role of oxygen in plants as he was of
carbon dioxide, as noted by Nash (1952, p. 110).
6. Do plants fix the elemental components of water in their dry substance?
In regard to a possible role for water in plant nutrition, de Saussure reasoned—
quite plausibly, but, as it turned out, incorrectly—that, because the amount of
oxygen the plant loses to the atmosphere is approximately equal to the amount
of oxygen in the carbon dioxide that the plant takes in, the most likely source for
most of the oxygen assimilated by plants is water. He also reasoned—correctly,
in this case—that water is probably the source of the hydrogen in plant dry mat-
ter. He showed experimentally that growing plants do, indeed, assimilate water
(although not as much as he thought, since they assimilate only the hydrogen,
the oxygen coming from the carbon dioxide). He thus confirmed van Helmont’s
guess that water is an important component of plant dry matter. Many consider
de Saussure’s recognition of the role of water to be his single greatest contribu-
tion to the understanding of plant nutrition (Nash 1952, p. 113). He provided the
Translator’s Introduction xxv

additional, important insight that the weights of hydrogen and oxygen in a plant
cannot be increased substantially unless the weight of carbon is simultaneously
increased. By linking the incorporation of carbon and water, he foreshadowed
the discovery of plant carbohydrates, but in his day, of course, there was no
knowledge of organic carbon compounds (Naef 1987, p. 336).
7. Do plants form and release water?
De Saussure found that water was formed and released during some processes
within living plants, such as in seed germination, and also in some processes
occurring in nonliving plants and plant products. For example, when atmospheric
oxygen combined with the carbon of humus or of wood extracts and formed
carbon dioxide, water was also produced. This water, however, was generally
composed from hydrogen and oxygen already contained within the dry substance
of the plant or plant product, not from the combination of atmospheric oxygen
gas directly with the hydrogen of the substance. He believed that the later stages
of fermentation presented a possible exception to this pattern, in that atmospheric
oxygen combined directly with plant hydrogen.
8. Are the minerals that are found in plant ash created by a vital force within the
plant or do they come from the soil?
De Saussure demonstrated that mineral substances disappeared from test solu-
tions in which plants were immersed by their roots, and that these substances
reappeared, in roughly the same proportions as they were absorbed, in the incin-
eration products (ash) of the plants. He thereby showed that the minerals in plants
do not owe their presence to a transmutation of carbon or other elements by a
vital force. Further, he demonstrated that roots do not absorb all solutes in the
same proportions, and that they absorb more of the water of solution than they
do any of its solutes.
De Saussure’s analyses of plant ash were the most accurate and extensive per-
formed up to that time, and his tabulations of the percentages of the various
minerals in the ash were the first of their kind (Browne 1944, p. 200). His anal-
yses revealed all of the minerals now known to be plant macronutrients, except
sulfur. He found traces of other minerals, in amounts too small for him to analyze
quantitatively.18
Another of de Saussure’s important findings was that the composition of plant ash
reflects the mineral composition of the soil in which the plants have grown. He
thereby showed that humus does, after all, play a role in plant life—not as a carbon
source, but as a mineral source. In addition, he found that mineral concentrations
in plants vary among different species, among different tissues, and with plant

18
Nine essential plant macronutrients are now known: carbon, oxygen, hydrogen, nitrogen, potas-
sium, calcium, magnesium, phosphorus, and sulfur. Adequate concentrations of the last four of
these nutrients in plant dry tissue are 1 % or less. In addition, there are eight known essential plant
micronutrients: chlorine, iron, boron, manganese, zinc, copper, nickel, and molybdenum (Stout
1961, as cited in Raven et al. 1999, p. 728).
xxvi Translator’s Introduction

age. For example, although all plants contain phosphorus,19 young tissue contains
more than old, and seeds are especially rich in it. Some minerals are lost from the
plant due to leaching by rainwater.
9. Are minerals essential for plant growth?
Although plants contain minerals in only very small amounts, these substances
are essential for their growth, de Saussure concluded. Some of the elements
absorbed may not be essential, he thought, but this had not been demonstrated
for the elements that plants always contain.
De Saussure did not trace the source of plant nitrogen convincingly, but he cor-
rectly concluded that the source of this nutrient is not atmospheric nitrogen gas,
as Ingen-Housz (1796) and many others had believed. De Saussure thought, in-
stead, that plant nitrogen came from extracts of plants and animals in the soil, or
ammonia that these extracts released to the soil and atmosphere.
In his studies of gases, de Saussure generally considered the volume of his exper-
imental plant to be a standard, or threshold, for determining whether oxygen or
carbon dioxide gas was being taken in or released significantly. He considered that
the “water of vegetation” in a plant’s parenchyma could hold a volume of carbon
dioxide approximately equal to, or slightly greater than, the volume of the plant,
and thus could store it and sequester it, at least temporarily, from decomposition
into carbon and oxygen by the plant’s photosynthetic apparatus. This stored carbon
dioxide could arise as a result of direct uptake of carbon dioxide by the plant from
the atmosphere or as a result of the combination of oxygen from the atmosphere with
the plant’s own carbon—or even (in oxygen-free atmospheres) from the combination
of carbon and oxygen that were both drawn from the plant’s own substance. Thus,
in order for plant uptake or release of a gas to be considered significant, the volume
of this gas had to exceed considerably the volume of the plant. If it did not, the gas
could be assumed to be in storage, as carbon dioxide, in the parenchyma.
The significance of one of Théodore de Saussure’s discoveries regarding gas
exchange was not understood by him and, in fact, came to be elucidated only in the
20th century. He found that the physiology of gas exchange in succulents, such as
cactus (Opuntia), differed from that in thin-leaved plants. Although not relevant to
the major physiological questions of his time, this was the first documentation (per
Moyse 1990) of the photosynthetic pathway known as CrassulaceanAcid Metabolism
(CAM), an important variant of the usual photosynthetic pathway (see Black and
Osmond 2005).

19
Analyzing phosphate content proved challenging for de Saussure. Although he showed that phos-
phates were important to plants, he had great difficulty in isolating them. Agricultural chemists
who came after him also had difficulty in developing better tests for phosphates, which were often
mistaken for alumina (Rossiter 1975, p. 25).
Translator’s Introduction xxvii

De Saussure’s Work Enables Completion of the Overall Chemical

Equation for Photosynthesis

The work of the early pioneers of photosynthesis research could, after Théodore de
Saussure’s contribution, be expressed in the basic, overall, chemical reaction20 of
photosynthesis that still stands today:
Light
Carbon dioxide + water → oxygen gas + carbon fixed in plant
Green plant
De Saussure was responsible for showing the participation of water in this reaction.
This equation does not incorporate the concept of energy, which was not fully
developed until the mid-19th century, when the law of the conservation of energy
was propounded: that energy can be neither created nor destroyed. Initially this
law was applied only to physical systems, because a vital force was believed to
create the energy that powered biological systems. German physician and physicist
Julius Robert von Mayer21 (1814–1878) contended in 1845, however, that energy
conservation applied to biological systems, as well. He concluded, correctly, that
photosynthesizing plants store, in the form of chemical energy, part of the light
energy reaching them from the sun. The stored chemical energy is later broken down
by the plants, or by organisms nutritionally dependent on the plants, and used as
tissue building blocks and for doing cellular work.
Before von Mayer’s insight, green plants could be understood only as chemical
manufacturers of organic matter, but after von Mayer, plants were also seen as
energy providers for life (Rabinowitch and Govindjee 1969, p. 9). The equation for
photosynthesis could then be re-written to show both the material and the energy
balance:
Light
Carbon dioxide + water → oxygen gas + carbon fixed in plant
Green plant
+ chemical energy

Critical Reception of Recherches chimiques sur la

Végétation, its Legacy, and Historical Perspectives

Recherches chimiques sur la Végétation was favorably received upon publication.

French chemist Claude-Louis Berthollet analyzed it and read extracts at the Institute
of France (l’Institut de France22 ), and his review of the book was published in the

20
This reaction is not written in chemical formula notation here because that notation was not in
use at the time de Saussure’s book was published. It was not developed until 1808, by Swedish
chemist Jöns Jacob Berzelius.
21
For a biography of von Mayer, see Caneva (1993).
22
Established in 1795, the Institute of France incorporated many of the former royal academies, in-
cluding the Royal Academy of Sciences (l’Académie royale des sciences), which had been abolished
2 years previously during the French Revolution.
xxviii Translator’s Introduction

Annales de Chimie (1804). An English translation of the review soon appeared in the
Philosophical Magazine (1805). Extracts of the book were published in the Journal
de Physique (de Saussure 1804b), and a German translation of these extracts appeared
in the Annalen der Physik (de Saussure 1804c). The year after the book appeared,
biologist Friedrich Siegmund Voigt (1781–1850) published a German translation of
the entire work (de Saussure 1805).
Despite its successful debut, however, the book’s important conclusions on the
source of plant carbon and mineral nutrients were largely ignored for about four
decades, although various authors cited the book liberally. No revised edition of the
book ever appeared.
Eminent German plant physiologist Julius von Sachs (1832–1897), although
praising the book highly, identified a problem: the writing style. He commented
(1875, 1890 English translation, p. 498):
It is not every one who can follow a work like this, which is no connected didactic exposition
of the theory of nutrition, but a series of experimental results which group themselves round
the great questions of the subject, while the theoretical connection is indicated in short
introductions and recapitulations, and it is left to the reader to form his own convictions
by careful study of all the details. . . . [T]he writer seems to confine himself too anxiously
within the limits of what is given in experience, and there is no doubt that many errors in
later times might have been avoided if the inductive proof of de Saussure’s doctrines had
been accompanied with a deductive exposition of them of a more didactic character.

Von Sachs continued (pp. 501–502), “Unfortunately de Saussure neglected to state

[the results on minerals] with due emphasis and to point out their fundamental im-
portance, and consequently doubts were entertained even till after 1830 respecting
the necessity of the constituents of the ash to vegetation”. Grandeau (1879, p. 46)
expressed similar regrets. Waksman (1936, p. 23) considered de Saussure’s ideas re-
garding the origin and chemical nature of soil organic matter or humus too advanced
for his times.
Another problem for the book was that some of the new research undertaken after
it was published yielded unclear or inconsistent results. Further, closely following
on the heels of the book were two highly influential works on agricultural chemistry
that continued to promulgate the humus theory, despite referring to de Saussure’s
findings. These books were by German agronomist Albrecht Daniel Thaer (1752–
1828), in his work of 1809–1812; and British chemist Humphrey Davy (1778–1829),
in his work of 1813. Davy accepted (1813, p. 273) de Saussure’s conclusions that
plants cannot form any of their ash constituents by synthesis or transmutation, but
Thaer did not (see Browne 1944, p. 180).
Other influential writers who clung to the humus theory during this period included
Swedish chemist Jöns Jacob Berzelius (1779–1848), in a treatise he co-authored on
animal chemistry (1815), and Swiss botanist Augustin Pyrame de Candolle (1778–
1841), in a treatise on plant physiology (1832). De Candolle was more modern in
his view on plant minerals, however, contending that they are derived entirely from
the soil (1832, Vol. 1, p. 390) (See Browne 1944, Chaps. 5 and 6, for more detail on
research and opinions during this period).
Translator’s Introduction xxix

Another factor in the delay in general acceptance of de Saussure’s conclusions

was an error of interpretation that cost him his scientific reputation for a time. He
mistakenly thought (his p. 56) that the pigment that gives a certain variety of Atriplex
hortensis its red or purple color was associated with photosynthetic oxygen produc-
tion, leading him to conclude, erroneously, that the green pigment was not essential.
“This oversight on the part of a man so highly regarded engendered doubts which
unfortunately persisted for a long time. . . .” (Morton 1981, p. 341).
De Saussure’s conclusions on plant carbon and the soil minerals did find support
in some new research studies, however, especially among German researchers. One
of these was agricultural chemist Carl S. Sprengel (1787–1859), who, in a work on
plant humus (1826), denied that plants could generate their own lime or other non-
combustible constituents but obtained them instead from the environment. Sprengel
also rejected Thaer’s notion that plants assimilate humus (Browne 1944, pp. 231–
239). Investigations by German researchers Wiegmann and Polstorff (1842), who
used a balance-sheet approach, also supported de Saussure’s findings on plant mineral
matter.
De Saussure’s ideas owe their real revival, however, chiefly to the influential
German chemist Justus von Liebig23 (1803–1873). Liebig (1840, 1847 English trans-
lation) stressed both the atmospheric source of plant carbon and the implications of
de Saussure’s findings on plant mineral nutrition for increasing crop yields through
fertilization.24 Liebig criticized the plant physiologists of his day for clinging, despite
abundant evidence to the contrary, to the idea that plants obtain their carbon from
the soil rather than the atmosphere.25 He argued that, since the humus itself consists
of the decayed matter of preceding plants, it was circular to contend that it was the
original source of plant carbon (Aikman 1894). Liebig also pointed out that there is
not enough soil humus to supply carbon to all of the plants growing in it. Based on
de Saussure’s ideas, manures could be seen in a different way: Animal dung provides
silicates of potash and small amounts of nitrogen, ashes provide potash, and bone
dust provides phosphates (Rossiter 1975, p. 25).
The renewed appreciation for de Saussure’s contributions was also aided by the
field research of French agricultural chemist Jean-Baptiste Boussingault26 (1802–
1887). Like Wiegmann and Polstorff (1842), Boussingault used a quantitative,
balance-sheet approach, and, in extensive field trials on crop plants, which he re-
ported in 1843–1844 and 1860–1891, he weighed and analyzed the materials he
applied to the soil and the crops he obtained under the various regimes. His balance
sheets showed the extent to which the fertilizers had met crop needs, and the extent
to which air, rain, and soil had contributed. He demonstrated that plants can grow
normally with nutrient salts alone, without any humus whatsoever (Waksman 1936).

23
For a biography of Liebig, see Brock (1997).
24
Liebig later advocated and developed artificial fertilizers.
25
See Werner and Holmes (2002) for the view that plant physiologists as a group did not deserve
such singling out.
26
For a biography of Boussingault, see McCosh (1984).
xxx Translator’s Introduction

It was not until the 1850s that the source of plant nitrogen was traced to the soil.
This accomplishment was also the work of Boussingault (1855, 1856, and 1857),
as discussed in detail by Aulie (1970). Another three decades would pass, however,
before Winogradsky (1890) isolated the soil bacteria that fix nitrogen.
Even after publication of Liebig’s book and Boussingault’s early research, how-
ever, the humus theory of plant carbon clung to life for a time. In 1845 plant
physiologist Jean-François Macaire-Prinsep, of Geneva, noted that the theory was
still controversial among physiologists. Evidence that de Saussure’s ideas were tak-
ing hold, however, include the publication in 1842 (English translation from the
German in 1849) of what Morton (1981, p. 377) called the first textbook of botany
that is modern in form and spirit, by Matthias Jakob Schleiden (1804–1881), co-
founder of the cell theory. Emphasizing plant physiology and its relationship to plant
structure and development, Schleiden cited de Saussure numerous times and rejected
both the humus theory and the concept of a vital force.
The renewed respect for Recherches chimiques sur la Végétation as the 19th cen-
tury wore on is reflected in the appearance of the second German translation of the
work, in 1890, by botanist Arwed Wieler27 (b. 1858). This translation appeared in a
two-volume set in the series Ostwald’s Klassiker der Exakten Wissenschaften. Wieler
stated (Vol. 1, p. 96) that he undertook the translation at the request of plant physi-
ologist Wilhelm Pfeffer because of the rarity and high price of the original. Wieler’s
German is more modern than that of the 1805 translation by Voigt, and thus more
accessible to a modern audience. (In 2009, the Wieler translation was reprinted in
two paperback, “optical character recognition,” volumes. In this reprint, Théodore
de Saussure’s name was, regrettably, rendered as “Théodore Laussure”).
The continuing appreciation of de Saussure’s book is also evident from the recog-
nition and praise bestowed on it, from Liebig’s time through the present, by numerous
authors from a wide range of disciplines—chemistry, agriculture, agronomy, soil sci-
ence, plant physiology, and plant nutrition.28 Despite this renewed respect, however,
French agronomist Paul Robin (2007, p. 47) laments, “Although it is constantly re-
ferred to by Liebig, de Saussure and the date 1804 are largely forgotten by today’s
agronomists”.

27
Wieler subsequently published one of the first studies (1905) of the long-term effects of industrial
air pollution on plants.
28
These include, from the mid-19th through late-20th centuries: de Candolle (1846, p. IX); von
Sachs (1875, 1890 translation, pp. 497–498); Pfeffer (1881, 1900 English translation, pp. 308–
309); Aikman (1894, pp. 14–15); Brown (1899, p. 476); Harvey-Gibson (1919, pp. 79–82); Spoehr
(1919, p. 36); Waksman (1936, p. 12); Browne (1944, pp. 192–193, p. 220); Rabinowitch (1945,
Vol. 1, p. 23); Gabriel and Fogel (1955, p. 162); Leicester (1956, p. 232); Solovine (1957, p. VIII);
Pilet (1975, p. 124); Russell (1976, p. 7); Morton (1981, p. 338); Buchs (1987, p. 172); Naef (1987,
p. 334); and Boulaine (1994, p. 21). In the 21st century, de Saussure’s life and work continue to
garner attention, especially in the French-language literature on agriculture and agronomy: See
Robin and Blondel-Mégrelis (2001), Candaux (2007), and Robin (2007). Magiels (2010), writing
in English, addresses de Saussure in the context of Ingen-Housz’s work.
Translator’s Introduction xxxi

Some Errors and Misinterpretations in Recherches chimiques sur

la Végétation

Despite the achievement represented by de Saussure’s book, there were, inevitably,

flaws. As already noted, the writing style is, in places, opaque. The text is plagued
with minor errors, including mistakes in conversion between Parisian and metric mea-
surement systems, some mis-statements, and mis-citings of his own tables. There
are also some more significant, conceptual errors, attributable to limitations in ex-
perimental techniques and the general state of knowledge of plant physiology and
nutrition in de Saussure’s day.
I have already pointed out de Saussure’s erroneous conclusion that the green
pigment in plants is not necessary for photosynthesis. In addition, he believed that,
in the light, plants decompose carbon dioxide into carbon and oxygen, and that the
carbon is then hydrated by a reaction with water (see p. 77 of his text), with most
of the oxygen of the carbon dioxide released to the atmosphere. It is now known
that, in green plants, water is the hydrogen donor in the photoreduction of carbon
dioxide, yielding an organic building block (CH2 O), with the release of gaseous
oxygen derived from the water. As already mentioned, the idea that the source of the
liberated oxygen is the absorbed carbon dioxide rather than water seemed logical
based on the near-equivalence between the volumes of carbon dioxide absorbed and
oxygen released. In fact, this view of the source of the oxygen prevailed until the
20th century.
The possibility that the oxygen comes instead from water had been suggested by
Lavoisier’s chemical colleagues Antoine-Francois Fourcroy (per Moyse 1990) and
Claude-Louis Berthollet (1786), but without experimental evidence. It was not until
the work of van Niel (1932) and Ruben et al. (1941)29 that water was shown to be the
source of the released oxygen. Because excess water is almost always present in plant
cells, however, and because oxygen is liberated from water only in proportion to the
carbon dioxide absorbed by the plant, the net effect is as envisioned by de Saussure
(Nash 1952, p. 119). The view that carbon dioxide is the source of the liberated
oxygen persisted, however, among some workers30 for more than a quarter-century
after the work of Ruben et al. (1941).
De Saussure’s erroneous conclusion that plants release nitrogen gas arose from
limitations in experimental methodology (Gabriel and Fogel 1955, p. 164, fn. 2). In
the early 19th century, gas analysis was still difficult (Partington 1962, p. 758) and
the techniques were not noted for their accuracy (Ihde 1964, p. 291).
Methodological limitations also hampered de Saussure’s chemical analyses of
the minerals in plant ash, as indicated by the large “deficits” he obtained (i.e., the
difference between the sum of the separate constituents he measured and the amount

29
Van Niel (1932) proposed, based on his research with bacteria, that water was the source of the
oxygen liberated by green plants, and Ruben et al. (1941), using heavy oxygen (O18 ), confirmed
the hypothesis.
30
For example, Warburg et al. (1969). (See Govindjee (1999) and Nickelsen and Govindjee (2011)
for a challenge to the conclusions of this paper.)
xxxii Translator’s Introduction

of ash he analyzed). Nevertheless, his mineral analyses represent a major advance

over previous work in this area. Liebig, who criticized de Saussure’s techniques, also
suggested (1840, 1847 English translation, p. 256) that, because all the analyses were
done by similar methods, the results were of interest because they were comparable
among themselves. (De Saussure’s lengthy tables of incinerations and analyses are
impressive, but, unfortunately, he made many mistakes when referring to the entries
in them.)
De Saussure did not always identify elements clearly. Given the state of chem-
ical knowledge in his day, this is understandable. It must be remembered that, in
1804, when Recherches chimiques sur la Végétation was published, elements and
compounds had not been clearly differentiated. Although Antoine Lavoisier had
propounded in 1789 that elements were substances that chemical analysis failed to
break down into simpler substances, and very early in the 1800s English chemist
and physicist John Dalton (1766–1844) had formulated an atomic explanation for
the apparent indestructibility of Lavoisier’s elements, Dalton’s theory was not well
publicized and accepted until 1808–1827.
Magiels (2010) criticized de Saussure’s book on the grounds of historical inac-
curacy, asserting that de Saussure failed to study the previous literature in the field
thoroughly and thus mistakenly attributed to Senebier—who was a close friend of
the de Saussure family—many contributions that were in fact made by Ingen-Housz.
Magiels noted, for example, that de Saussure credited Senebier with the discovery
that the nongreen parts of plants do not expel oxygen, when in fact it was Ingen-
Housz (1779) who first made this observation. Magiels also criticized de Saussure
for dissecting the humus theory without mentioning Ingen-Housz, who had attacked
the theory 8 years previously. Ingen-Housz, however, stated his view briefly and
without experimental data to support it, whereas de Saussure based his conclusion
on detailed experimentation. It is true that, given the discord between Senebier and
Ingen-Housz, Senebier may have tended to ignore Ingen-Housz’s work and that de
Saussure to some extent followed suit. Still, de Saussure’s book, which is of only
modest length, contains more than 100 citations, including six of Ingen-Housz, in-
dicating that de Saussure examined a considerable amount of the previous literature
(see Appendix 3: Bibliography of References Cited by de Saussure (as expanded by
the translator)).
I have not thoroughly annotated de Saussure’s text, but I have noted some clear
errors in brackets within the translation.

The Life and Scientific Background of Théodore de Saussure

When Recherches chimiques sur la Végétation was published, Théodore de Saussure

was 36 years old and well settled into a quiet, somewhat reclusive life as a plant
researcher in his own private laboratory. In contrast to his staid middle and later
years, however, de Saussure as a young man (see Fig. 1) had known adventure and
scientific accomplishment with his father under arduous conditions in the high Alps,
Translator’s Introduction xxxiii

Fig. 1 Portrait of Théodore

de Saussure as a young man

and he had suffered when the intense political turmoil of the French Revolution had
reached his home city of Geneva.31
Théodore de Saussure was born in Geneva on October 14, 1767. His parents,
Horace-Bénédict de Saussure (1740–1799) and Albertine-Amélie Boissier de Saus-
sure (1745–1817), were wealthy, and Théodore grew up in a luxurious family
townhouse (Fig. 2). His full name was Nicolas-Théodore, but he was usually called
Théodore to distinguish him from his grandfather Nicolas de Saussure (1709–1790),
a wealthy landowner and agriculturalist who had published a number of papers on
agricultural subjects (Zumkeller 2001). Théodore was the second of three children.
The first child was Albertine (1766–1841) and the third Alphonse (1770–1853).
As his children were growing up, Horace-Bénédict, who was learned in geology,
physics, meteorology, chemistry, and botany, was developing an illustrious career
as a geologist, Alpine explorer, and professor of natural philosophy at the Academy
of Geneva (l’Académie de Genève), the predecessor of the University of Geneva,32
where he taught from 1762 until 1786 (Carozzi 1981).

31
The political upheaval in Geneva began in the early 1790s. In 1798, the French Republic annexed
all Swiss territory. This territory became part of the Napoleonic Empire—which began in 1804, the
year de Saussure’s book was published—and remained a part of the empire until the end of 1813.
32
For detailed treatments of Horace-Bénédict de Saussure’s life and work, see Freshfield (1920),
Sigrist (2001), Bungener (2002), and Carozzi (1981, 2005).
xxxiv Translator’s Introduction

Fig. 2 Geneva townhouse where Théodore de Saussure grew up

Horace-Bénédict de Saussure and his son Théodore were among the natural
philosophers responsible for a burst of scientific achievement in Geneva in the late
18th century (Sigrist 1990). The tradition of natural history in Geneva has been
characterized as relying strongly on observation and experiment, in contrast to a
more speculative and theory-based tradition that was prevalent in some other parts
of Europe (Roger 1987). Photosynthesis pioneer Jean Senebier and plant researchers
Charles Bonnet and Augustin Pyrame de Candolle were members of this scientific
circle in Geneva.
Horace-Bénédict disapproved of the public education in Geneva and therefore
directed the early schooling of his children at home. Macaire (1845) blamed
Théodore’s shy disposition on this private mode of instruction, because it was so
isolating.
From 1782–1786, Théodore attended the Academy of Geneva, where he took
courses in mathematics, science, and history (Robin and Blondel-Mégrelis 2001;
Candaux 2007). He studied physics under the noted scientist Marc-Auguste Pictet
(1752–1825) (Candaux 2007). At the outset, Théodore shared his father’s interest
in the physical sciences, especially geology and meteorology, and he began his
scientific career as his father’s companion and field assistant on Alpine expeditions.
Horace-Bénédict devised hygrometers, eudiometers, and other precision scientific
instruments and trained Théodore in their use (Browne 1944), eventually giving the
young man responsibility for recording various physical data on the expeditions.
Théodore first became involved, in a limited way, in 1787, when his father made
Translator’s Introduction xxxv

the second-ever conquest of the summit of Mont Blanc (elevation 4,810 m), the
highest peak of the Alps (Freshfield 1920; Rowlinson 1998). Judging that his 20-
year-old son was not strong enough to make this climb, Horace-Bénédict left him
in the village of Chamonix, at the foot of the mountain. There, Théodore recorded
meteorological data for comparison with his father’s readings on the summit (Macaire
1845).
In 1788, the year following the Mont Blanc ascent, Théodore de Saussure became
a full partner with his father on an arduous expedition to the snowfields of the Col
du Géant, a high glacial pass. On that expedition, which was the most ambitious and
challenging of Horace-Bénédict’s career (Freshfield 1920), father and son spent 17
days and nights under harsh, storm-wracked conditions at an elevation of 3,436 m.
There, Théodore assisted in various physical, chemical, mineralogical, and meteoro-
logical experiments and analyses. He made numerous observations on the density and
composition of the atmosphere (Briquet 1940, p. 425). To measure atmospheric den-
sity, he observed the oscillation of a pendulum—a method he found disappointingly
inexact. He measured the oxygen content of the air using nitric oxide (Priestley’s
“nitrous-air” method), and tested for the presence of atmospheric carbon dioxide
using limewater and caustic potash (Robin and Blondel-Mégrelis 2001). He also
conducted experiments on the dissolution of metals in sulfuric acid (Macaire 1845).
The following year, Théodore published his first scientific paper, a physico-
chemical analysis of the mineral “sappare” (aluminum silicate) (de Saussure “fils”
1789).33 Also that year, he and his father made an expedition to Monte Rosa,
where Théodore again measured the density of the atmosphere, this time using a
large, empty, tightly closed glass flask, which he weighed at 70 different elevations.
Théodore found that the differences in weight were exactly proportional to the differ-
ences in barometric pressure readings (Macaire 1845). Théodore thereby confirmed
Boyle’s Law (known as Mariotte’s Law in France) by a new method (Hoefer 1867,
p. 374; Briquet 1940, p. 425). Théodore’s publication on this subject (de Saussure
“le fils” 1790) brought him to the attention of the scientific community (Robin and
Blondel-Mégrelis 2001). In each of the subsequent three years, he published a paper
on mineralogy (de Saussure “fils” or “le fils,” 1791, 1792, and 1793). These early
chemical, mineralogical, and atmospheric experiments and analyses, although con-
fined to the physical sciences, presage the studies that Théodore was later to conduct
on the roles of gases and minerals in plant chemistry and physiology.
The French Revolution, which had begun in Paris the month of the de Saussures’
Monte Rosa climb, soon diverted Horace-Bénédict’s attention. He made only one
further Alpine expedition—a visit, with Théodore, in 1792, to the St. Théodule
glacial pass, to measure the height of the Matterhorn (Freshfield 1920).
During the early years of the revolution, while the de Saussure family fortune
remained intact, Théodore was able to travel abroad to meet with foreign researchers

33
Following the custom of the day, the papers Théodore published during his father’s lifetime were
under the authorship of “de Saussure fils” or “de Saussure le fils,” indicating that he was the son of
a scientist of the same surname.
xxxvi Translator’s Introduction

(Hart 1930). He made his first trip to London in the summer of 1791. In 1793–
1794, once again in Britain, he met with Joseph Priestley, Joseph Black, and Henry
Cavendish, all of them experts in chemistry and eudiometric methods (Robin and
Blondel-Mégrelis 2001; Candaux 2007). Much of Horace-Bénédict’s fortune was
invested in France, however, and was lost during the revolution (Freshfield 1920),
while his wife’s income was drastically reduced (Carozzi 1981). In late December
of 1793, Théodore’s mother wrote to him in Britain of the financial difficulties, and
advised him to seek a traveling tutorship (Hart 1930). Unable to find such work,
Théodore returned to Geneva in the summer of 1794 (Candaux 2007).
The revolutionary turmoil in Geneva soon forced Théodore and his brother to flee
to Rolle, about 30 km away, and, that same summer, his father suffered the first of a
series of strokes. According to Freshfield (1920), Théodore spent a great deal of his
time in Rolle during 1794 and 1795 correcting proofs of the last two volumes of his
father’s magnum opus, Voyages dans les Alpes (1779–1796). Théodore remained in
Rolle at least until the summer of 1795 (Candaux 2007). In 1796, upon returning to
Geneva, he married Renée Fabri (1767–1847) and began research in plant chemistry
(Freshfield 1920; Hart 1930; Robin and Blondel-Mégrelis 2001; Candaux 2007).
When Théodore began his plant research, knowledge of biological chemistry
lagged behind that of mineral chemistry, providing an opportunity for significant
contributions (Macaire 1845). Other factors may also have propelled Théodore to-
wards plant research. Eighteenth-century Geneva was an important center for botany,
especially plant physiology (Naef 1987; Sigrist and Bungener 2008), and some of
Théodore’s family and their friends were deeply involved in botanical pursuits. In-
deed, botany had been his father’s first love (Freshfield 1920; Bungener 2001).34
Further, Horace-Bénédict and his own father, Nicolas, maintained a botanical garden
at the family estate at Frontenex (Fig. 3), on the outskirts of Geneva. There, Horace-
Bénédict grew plants that he had collected, in order to observe them at different
developmental stages for purposes of identifying and describing them (Bungener
2001; Sigrist and Bungener 2008). Thus, from early in life, Théodore would have
been familiar with the scientific study of plant growth. In his experimental work,
Théodore used a wide variety of plant material (Moyse 1990), including pea, beans,
wheat, periwinkle, pellitory, loosestrife, mint, cactus, rhododendron, bilberry, pine,
juniper, and fir. Some of these plants probably came from the family garden, although
some came from the Alps.
Théodore de Saussure was profoundly influenced by pioneering photosynthesis
researcher Jean Senebier, who was a close friend of Horace-Bénédict’s and instru-
mental in Théodore’s education. A major line of research pursued by Théodore was
one that had intensely occupied Senebier—the absorption of carbon dioxide and
liberation of oxygen by plant leaves. In addition, Théodore was undoubtedly influ-
enced by his great-uncle Charles Bonnet, whose 1754 book on plant physiology had

34
Horace-Bénédict had launched his scientific career in 1760 by collecting plants near Geneva and
from high Alpine areas for Albrecht von Haller (1708–1777), a Swiss physiologist and anatomist
who was also a noted botanist and author of a major work (1768) on the Swiss flora. Horace-Bénédict
had also published (1762) on plant anatomy.
Translator’s Introduction xxxvii

Fig. 3 De Saussure family estate at Frontenex (on the outskirts of Geneva), where Théodore de
Saussure’s father and grandfather maintained a botanical garden

inspired Ingen-Housz’s experimental methodology and who was a neighbor and close
friend of the de Saussure family as Théodore was growing up (Freshfield 1920). Fur-
ther, a number of the subjects Théodore chose to investigate—soils, humus, and plant
mineral nutrition—indicate an agricultural impetus that probably reflected the influ-
ence of his agriculturist grandfather Nicolas. In his preface to Recherches chimiques
sur la Végétation, Théodore acknowledged an agricultural motivation.
Théodore’s early papers on plant nutrition provided the groundwork for
Recherches chimiques sur la Végétation. The first was a major study of carbon diox-
ide metabolism, published in the Annales de Chimie (de Saussure “le fils” 1797) and
reprinted in the Journal de Physique (de Saussure “le fils” 1798). This paper demon-
strated that plants, like animals, continually form carbon dioxide from atmospheric
oxygen and their own carbon, and that, if carbon dioxide production is not detectable
in a plant, it is because the plant is decomposing this gas as it is formed. Further, he
showed that, in sunlight, plants must decompose carbon dioxide in order to grow,
and that, although growth in sunlight is promoted to some extent by elevated concen-
trations of this gas, such concentrations are harmful to plants in shade. This paper
formed the nucleus for those chapters of Recherches chimiques sur la Végétation
dealing with gas exchange and the metabolism of carbon and oxygen. De Saussure’s
second paper on plant physiology (de Saussure “fils” 1799), a study of the effects of
oxygen on seed germination, laid the foundation for Chap. 1 of his book.
In the late 1790s, Théodore again left Geneva, this time with his wife. The couple
remained abroad for several years. In the spring of 1800, on his first trip to Paris,
Théodore took courses from the noted French pharmacist and chemist Louis-Nicolas
xxxviii Translator’s Introduction

Vauquelin (1763–1829) and kept a journal (de Saussure 1800a) that provides infor-
mation on his links to the world of Lavoisier (Robin and Blondel-Mégrelis 2001;
Candaux 2007). In Paris, Théodore also presented a paper (de Saussure “le fils”
1800b) to the Philomatic Society (la Société Philomathique),35 on his comparative
analyses of the minerals contained in the ash of plant samples he had collected,
starting in the summer of 1798, from calcareous and granitic Alpine mountains. His
results showed that the mineral composition of plants varies according to the nature of
the bedrock underlying the soils in which the plants are rooted. This paper contained
the first data for those chapters of his book that address plant mineral nutrition. Robin
and Blondel-Mégrelis (2001) express regret that, although this paper was the first
application, on the ground, of the methods of the “new chemistry” and was therefore
a significant contribution to knowledge of the role of soil in plant development and
thus to agronomy, it has largely been forgotten.
In 1799, Théodore’s father died. Soon thereafter, Théodore’s papers began ap-
pearing under his own name, Théodore de Saussure. His early publications under this
name were in the physical rather than the plant sciences. The first (de Saussure 1801)
concerned alumina (aluminum oxide), and the second (de Saussure 1802) focused
on carbon dioxide and the electric spark.
Théodore and his wife remained in Britain and France until 1802 (Hart 1930).
Upon returning to Geneva, he expected to occupy a promised chair in natural history
and chemistry at the Academy of Geneva, but instead was named honorary professor
of mineralogy and geology. Acutely disappointed at not receiving an appointment
in plant chemistry, he immediately requested and obtained an 18-month leave of
absence in order to complete his book Recherches chimiques sur la Végétation and
to prepare to teach a course in a field he had not chosen. Upon finishing the book, he
taught a single course in mineralogy and geology, in 1805 (Candaux 2007), but then
no longer taught, despite his presence on the faculty roster for 33 years (Borgeaud
1909). Macaire (1845) attributed his failure to teach to his shy nature.
Recherches chimiques sur la Végétation is Théodore de Saussure’s sole book. The
initial acclaim for the work brought him recognition as a major contributor to the
understanding of plant physiology. He was elected a corresponding member of the
Institute of France in 1805 (Haag and Haag 1859; Pilet 1975; Naef 1987).
After publication of the book, Théodore lived another 41 years—more than half his
lifetime. During this period, he continued his research on plant chemistry and physi-
ology, publishing numerous papers. He also conducted studies in organic chemistry.
For example, he analyzed alcohol and ether by three different methods between 1807
and 1814 (Ihde 1964). De Saussure (1821) showed that, with regard to the air, green
fruits behave as do leaves. His other significant contributions included the demon-
stration (de Saussure 1825) that seeds whose germination has been interrupted by
drying can resume normal development if they are rehydrated. In 1841, in a notable
presentation on plant nutrition to a scientific congress in France, he showed that

35
The year he made this presentation, Théodore was elected a member of the Philomatic Society,
which had been founded in 1788 (Moyse 1990).
Translator’s Introduction xxxix

nitrogen must be absorbed by the roots, although he did not disprove the absorption
of atmospheric nitrogen.
Théodore de Saussure was elected to the Royal Society of London as a foreign
member in 1820. He served a number of times as a member of the representative
council of Geneva. He died in Geneva on April 18, 1845, leaving no descendants.
De Saussure’s archives are housed at the Library of Geneva (Bibliothèque de
Genève; formerly called Bibliothèque publique et universitaire de Genève). Candaux
(2007) lists the contents of these archives, which include a number of drafts of
Recherches chimiques sur la Végétation. Further information about de Saussure’s
life and work can be found in Macaire (1845); Haag and Haag (1859, pp. 191–192);
Hoefer (1867, pp. 374–375); Borgeaud (1909, pp. 55, 60, 76, 83, 112, 171, 177);
Freshfield (1920); Hart (1930); Briquet (1940, pp. 425–428); Cannon (1961–1962,
1963); Pilet (1975); Robin and Blondel-Mégrelis (2001); and Candaux (2007).

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A Note on the Translation

In the mid-1790s, when Théodore de Saussure began the research that laid the ground
work for Recherches chimiques sur la Végétation, the turmoil of the French Revolu-
tion had spread to his home city, Geneva. The Swiss Republic was annexed by the
French Republic in 1798 and became part of the Napoleonic Empire in 1804, the
year this book was published. In the upheaval of this period, many of the conventions
of the ancien régime were replaced. The innovations included the metric system and
a drastic redesign of the calendar system. In addition, the forms of address, or hon-
orifics, changed. The text of de Saussure’s book reflects the transitional nature of
this period and adheres neither to the old nor to the new conventions completely. He
compiled some of his book from papers he had written, or research he had under-
taken, starting in the mid- to late-1790s, before the new systems had taken hold, and
he did not complete the work until 1804, after these newer conventions had become
more accepted.
De Saussure mostly, but not always, reported his measurements (linear, volume,
weight) in units of both the old Parisian system and the newer metric system. The
Parisian values that he reported are commonly more rounded than their metric equiv-
alents, strongly suggesting that his laboratory equipment was calibrated in Parisian
units. Although some of the Parisian units sound familiar to English speakers, such
as inch (pouce), pound (livre), and grain (grain), these units are slightly different
from their English counterparts. For the convenience of the reader, I have provided,
in Appendix 1, conversions among the Parisian units, the English units (which de
Saussure himself does not use), and the metric ones. De Saussure often rounds off
his figures roughly, and his conversions between units of the Parisian and metric
systems are not always exact.
In some places in his text, de Saussure used the familiar, pre-Revolutionary,
January-December, calendar system, which was re-adopted in 1805 by Napoleonic
France. At other places he used the French Revolutionary system (monthsVendemaire
through Fructidor, and years that began with An I (year 1, or 1793) and continued
through An XIV (year 14, or 1805)). Some of the literature he cited was in French-
language publications that appeared during the Revolutionary era and thus used this
system. In Appendix 3, in which I list the authors de Saussure cited and as much
information about the exact sources as I could find, I use only conventional dates.

xlix
l A Note on the Translation

Although Europe was roiled by the French Revolution during the time de Saussure
was working on his book, the chemical revolution, which had begun some years
before the French Revolution, was well advanced, and by the time de Saussure’s
book was published, the new chemical concepts and nomenclature were largely in
place. Many of the chemical terms de Saussure used are the same as, or similar to,
modern terms.
Chemical terminology has evolved since de Saussure’s day, however, and some of
his terms are now obsolete. This situation led to some complexity in the way I dealt
with chemical nomenclature in the translation. For some of the older chemical terms
for which there are unambiguous modern equivalents, the modern terms have been
substituted throughout the translated text. For example, I used “carbon dioxide” in
place of the now-archaic term “carbonic acid gas”, and “chloride” in place of the
now-obsolete “muriate”. Where I have made such a substitution, I have provided de
Saussure’s term in square brackets at the first occurrence of the term in the text, and
sometimes again later to remind the reader.
For some terms, however, the substitution of modern for older terminology was
problematic, as in the case of “potash” and “soda”—two terms that de Saussure
employed frequently. At the time he wrote his book, the elements potassium and
sodium had not yet been isolated from potash and soda, respectively, and the terms
“potash” and “soda” might refer to more than one chemical that contained potassium
or sodium. In this translation, I have substituted the terms “potassium” and “sodium”
for “potash” and “soda” only in cases where de Saussure used these terms as part of
the name of a compound that is still valid today, and only where the general context
did not make such substitution confusing for the reader. In cases where it could be
confusing, I retained the older terminology, even in the names of still-valid chem-
ical compounds. For example, in Chap. 9, which contains detailed descriptions of
complex chemical manipulations and is heavily laden with chemical terminology,
de Saussure frequently used the terms “potash” and “soda”, both by themselves and
to refer to components of compounds (for example, “phosphate of potash”, which,
in modern terminology, is potassium phosphate). The substitution of “potassium”
for “potash” in the names of chemical compounds but not for the term “potash”
when used by itself would have resulted in a disconnected narrative that adhered
neither to the new nor to the old terminology, and could have been confusing for the
reader. Therefore, in that chapter, I have generally retained the older names for all
terms (including lime and magnesia, because the elements calcium and magnesium,
respectively, had not yet been isolated from these compounds). I have provided
a glossary of terms (Appendix 2) that gives the older terms and their modern
equivalents.
In addition to the changes in chemical terminology, I have made some changes
where a literal translation of the original sounded awkward in English. Further, for
passages in which the flow of ideas seemed difficult to follow, or where unusual
terms were used, I have inserted, in italicized type in square brackets, explanations
or clarifications that I thought would be helpful to the reader. (This translation is
not a thoroughly annotated version of the text, however). I have also inserted, in
bold type in square brackets, the page numbers of the original French text, for ease
A Note on the Translation li

in consulting that text. All footnotes are de Saussure’s, as is text that is set off in
ordinary parentheses.
De Saussure made a number of clear errors, mostly minor, in his text. These
include mistakes in conversion between the old, Parisian measurement units and
their metric equivalents; misinterpretation of another author’s writing; some mis-
statements; and mis-citings of his tables. I have generally corrected, or pointed
out, the errors that I have spotted, giving, where appropriate, de Saussure’s original
wording or numerical figures, or an explanation for my change, in italicized type
in square brackets after the corrected text. (Errors were a particular problem in de
Saussure’s references in Chap. 9 to his extensive tables of incinerations and ash
analyses.) I have also corrected the four errata that de Saussure listed at the end of
his text.
Although de Saussure’s book is of only modest length, it includes more than
100 citations, drawn from diverse fields. As was common in his day, however, he
supplied no bibliography and provided only sketchy details for many of his sources—
sometimes no more than an author’s name. I have traced these references as far as I
could, obtaining nearly all authors’ full names and, for most of them, the full titles
of books, journals, and papers. For many of them, I found relevant page numbers.
The fleshed-out list is provided in Appendix 3. Some of the entries in this list are,
necessarily, only plausible guesses on my part about his sources (and I note these as
such), but the list should provide the reader an entrée to the broad range of literature
de Saussure consulted in his research.
In the text itself, I have left the references largely as de Saussure cited them,
except for a few changes related mainly to forms of address and name renderings.
For authors in French-language publications in the older, pre-Revolutionary era, the
honorifics used were “M.” and “MM.” (for Monsieur and Messieurs), whereas in the
Revolutionary-era publications, “C.” and “CC.” (for Citoyen (Citizen) and Citoyens
(Citizens)) were commonly used instead. For the sake of simplicity, I have deleted all
honorifics in the translation itself and in the fleshed-out citations listed in Appendix 3,
as well as in the reference list that I compiled for my introduction. In the text of the
translation, I have italicized foreign words and the titles of all books and journals
cited by de Saussure.
In addition, in the translation and in Appendix 3, I have made the authors’ sur-
names conform to generally accepted spellings or forms. The name changes I made
are as follows:
My rendering of the name De Saussure’s original
DeCandolle Decandolle
Haüy Hauy
Ingen-Housz Ingenhoutz
Lefebure Le Febvre
Marggraf Margraff
Perthuis Pertuis
Rückert Ruckert
Rumford Rumfordt
van Helmont vanhelmont
lii A Note on the Translation

I have also standardized the scientific (Latin) names of the plant species mentioned
in the text. De Saussure was not consistent in this regard. In some places he used a
lower-case initial letter for genera, other times a capital letter. Sometimes he italicized
the Latin names, other times not. For the sake of consistency and to conform to
modern nomenclature standards, I have italicized all Latin names and capitalized
the first letter of all Latin generic names. Otherwise, I have preserved the scientific
names as he presented them, even though many of these have been changed since
his time.
De Saussure’s sentences tended to be long, with a much more liberal use of
commas, semicolons, and colons than is usual today. I have taken the liberty of
removing many of his commas, and of substituting periods for many of his semi-
colons and colons, because periods seem more natural by modern standards.

Hill, J. F.
Title page of Recherches chimiques sur la Végétation
Contents of Original de Saussure Book

1 Influence of Oxygen Gas on Germination . . . . . . . . . . . . . . . . . . . . . . . . . 1

1.1 Of Germination . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.2 Use of Oxygen Gas in Germination . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.3 Of the Changes that a Seed Undergoes Through Germination . . . . . . 5
1.4 Of the Influence of Light on Germination . . . . . . . . . . . . . . . . . . . . . . . 8

2 Influence of Carbon Dioxide Gas on Plant Growth . . . . . . . . . . . . . . . . . 11

2.1 Influence of Carbon Dioxide Gas on Germination . . . . . . . . . . . . . . . . 11
2.2 Influence of Carbon Dioxide Gas on Developed Plants . . . . . . . . . . . . 11
2.2.1 Results in Sunlight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
2.2.2 Results in the Shade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
2.3 Elaboration of Carbon Dioxide Gas by Leaves is Necessary
to Their Growth in the Sun . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
2.4 Of the Decomposition of Carbon Dioxide Gas by the Green Parts
of Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
2.4.1 First Experiment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
2.4.2 Second Experiment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
2.4.3 Third Experiment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
2.4.4 Fourth Experiment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
2.4.5 Fifth Experiment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
2.5 Plants Grown with Pure Water in the Open Air Take Carbon from
the Small Amount of Carbon Dioxide Gas that Occurs Naturally
in Our Atmosphere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
2.5.1 Further Remarks on the Decomposition of Carbon
Dioxide Gas by Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
2.6 Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23

3 Influence of Oxygen Gas on Developed Plants . . . . . . . . . . . . . . . . . . . . . 25

3.1 Of the Changes that Oxygen Gas Undergoes Through its
Contact with Leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
3.2 Experiments on Inspiration [this section title supplied by translator] 26
3.2.1 Inspiration by Cactus opuntia . . . . . . . . . . . . . . . . . . . . . . . . . . 26

lv
lvi Contents of Original de Saussure Book

3.2.2 Leaves Saturated with Oxygen Gas Form Carbon

Dioxide Gas in Darkness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
3.2.3 A Cactus that has Inspired Oxygen Gas Cannot Expire it
Through the Effect of the Vacuum of an Air Pump . . . . . . . . . 28
3.2.4 Leaves do not Inspire Appreciably in Gaseous Environments
that Lack Free Oxygen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
3.2.5 A Cactus Inspires Carbon Dioxide Gas in the Same
Proportion as Oxygen Gas, When the Carbon Dioxide
is Mixed with the Oxygen in a Small Amount . . . . . . . . . . . . . 29
3.2.6 A Cactus Never Appears Saturated with Oxygen Gas After
Staying for Several Days in Darkness, or an Indefinite Time
in the Open Atmosphere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30
3.2.7 A Cactus Whose Structure has been Destroyed Makes no
Appreciable Inspiration in Atmospheric Air . . . . . . . . . . . . . . . 30
3.2.8 Inferences from the Preceding Observations: The Oxygen
Gas Inspired by Leaves in the Dark is Converted into
Carbon Dioxide Gas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
3.3 Experiments on Expiration [this section title supplied by translator] 33
3.3.1 Expiration by Cactus in Atmospheric Air . . . . . . . . . . . . . . . . . 33
3.3.2 Expiration by Cactus Under Distilled Water and in
Nitrogen Gas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
3.3.3 Expiration is Proportional to Inspiration . . . . . . . . . . . . . . . . . 34
3.3.4 Effect of Quicklime or Potash on Expiration by Cactus . . . . . 35
3.3.5 The Results Obtained on Expiration by Cactus are
Applicable to the Leaves of Other Plants . . . . . . . . . . . . . . . . . 36
3.4 Plant Growth in Pure Oxygen Gas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
3.5 Relative Amounts of Oxygen Gas Consumed in Darkness
by Different Leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
3.6 Influence of Atmospheric Oxygen Gas on Plant Roots . . . . . . . . . . . . 41
3.7 Of the Changes that Oxygen Gas Undergoes Through its
Contact with Roots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
3.8 Of the Changes that Oxygen Gas Undergoes Through
Woody Stems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
3.9 Of the Changes that Oxygen Gas Undergoes Through its
Contact with Flowers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
3.10 Influence of Oxygen Gas on Fruits . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
3.11 Utility of Oxygen Gas in the Nutrition of Plants . . . . . . . . . . . . . . . . . 50
3.12 Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

4 Influence of Oxygen Gas on Some Plant-Derived Substances

[“proximate constituents of plants”] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
4.1 Use of Oxygen Gas in the Precipitation of Extracts . . . . . . . . . . . . . . . 53
4.2 Use of Oxygen Gas in the Conversion of Wine to Vinegar . . . . . . . . . 55
4.3 Influence of Oxygen Gas on Dead Wood . . . . . . . . . . . . . . . . . . . . . . . 57
Contents of Original de Saussure Book lvii

4.4 Condensation of Oxygen Gas by Oils . . . . . . . . . . . . . . . . . . . . . . . . . . 59

4.5 Condensation of Oxygen Gas by Putrefying Plant Substances . . . . . . 59
4.6 Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61

5 Of Plant Humus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
5.1 Research on the Composition of Humus . . . . . . . . . . . . . . . . . . . . . . . . 63
5.2 Extractive Substances of Humus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
5.3 Of the Salts Contained in Humus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
5.4 Of Changes that Oxygen Gas Undergoes by its Contact with Humus 68
5.5 Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
5.6 Note on the Carbonization of Various Plant Substances . . . . . . . . . . . 70

6 Of Plant Growth in Oxygen-Free Environments . . . . . . . . . . . . . . . . . . . 75

6.1 Of Plants that Cannot Maintain their Growth in Nitrogen Gas . . . . . . 75
6.2 Of Plants that can Grow in Nitrogen Gas . . . . . . . . . . . . . . . . . . . . . . . 76
6.3 Of Plant Growth in Carbon Monoxide [“oxide of carbon”] Gas
(Berthollet’s oxycarburetted hydrogen) . . . . . . . . . . . . . . . . . . . . . . . . . 80
6.4 Of Plant Growth in Hydrogen Gas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
6.5 Of Plant Growth in a Vacuum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
6.6 Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82

7 Of the Fixation and Decomposition of Water by Plants . . . . . . . . . . . . . 83

7.1 Investigations on the Fixation of Water by Plants Growing in
Atmospheric Air Freed of Carbon Dioxide Gas . . . . . . . . . . . . . . . . . . 83
7.2 Fixation of Water by Plants Growing in a Mixture of Ordinary
Air and Carbon Dioxide Gas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
7.3 Of the Decomposition of Water by Plants . . . . . . . . . . . . . . . . . . . . . . . 86
7.4 Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89

8 Of the Absorption of Solutions by Plant Roots . . . . . . . . . . . . . . . . . . . . . 91

8.1 Water and Gas are Insufficient Nourishment to Bring about
the Full Development of Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
8.2 Do Plants Absorb the Substances that are Dissolved in Water
in the Same Proportion as they Absorb the Water? . . . . . . . . . . . . . . . 93
8.3 Do Plants Absorb Certain Substances in Preference to Others,
from a Liquid Containing Several Substances in Solution? . . . . . . . . 96
8.4 Considerations on the Saline or Mineral Substances that Enter
into the Composition of Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
8.5 Application of the Foregoing Observations to Research on the
Amount of Nutrients that the Humus Substance Alone Provides
to Plant Roots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100
8.6 Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
lviii Contents of Original de Saussure Book

9 Observations on the Ash of Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103

9.1 Observations Made by Various Authors on the Amounts of Ash
Produced by Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
9.2 Principle by Which Ash Varies in Amount in Woody and
Herbaceous Plants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
9.3 General Composition of Ash; Influence of Soil . . . . . . . . . . . . . . . . . . 105
9.4 Of the Alkaline Salts in Ash . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107
9.5 Of Earthy Phosphates in Ash . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110
9.6 Of Free or Carbonated Lime in Ash . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
9.7 Of Silica in Ash . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112
9.8 Of Metallic Oxides in Ash . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
9.9 Influence of the Atmosphere on the Ash of Plants . . . . . . . . . . . . . . . . 113
9.10 Details on the Procedures used for Incineration . . . . . . . . . . . . . . . . . . 115
9.11 Details on the Procedures used for Analyzing Ash . . . . . . . . . . . . . . . 116
9.11.1 Note on the Combination of Phosphate of Potash
with Lime . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119

10 Chapter Summaries . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155

 De Saussure’s Plate of Figures

Plate from de Saussure’s 1804 book, Recherches chimiques sur la Végétation, showing Figures I–VIII.

lix
[p. 1] Chapter 1
Influence of Oxygen Gas on Germination

1.1 Of Germination

Oxygen gas and water are two agents that nature uses simultaneously to stimulate the
development in seeds known as germination. Water alone, by entering their cellular
tissue and filling it, increases their volume but does not cause them to germinate.
The [appearance of the] radicle is, for many but not all seeds, an [p. 2] indication
of germination. Some seeds reveal this organ by a swelling due to the imbibition
of water and not due to an act of plant growth. We cannot doubt it, knowing that
certain seeds that are dead or have lost, through decrepitude, the ability to germinate
nonetheless sprout their radicles as if they were capable of new development.
Coffee, for example,1 which can sprout only if sown 5 or 6 weeks after being
harvested, produces the phenomenon in question after an indefinite number of years.
It produces the effect not only in cold water, without contact with air, but also in
liquids certainly incapable of eliciting or sustaining growth, such as boiling water,
vinegar, and some saturated saline solutions. This very small increase, the end of
which can be anticipated before the swelling that produces it, does not change [p. 3]
the taste, odor, or other properties of the seed. The imbibition of water without
contact with air has no other effect than to dispose the seed to rapid decay.
Some seeds, such as peas, lentils, and aquatic seeds, can germinate under water,
but only in water that is impure or contains oxygen gas foreign to its composition.
I passed boiling water under a receptacle [de Saussure used, almost exclusively, the
term “recipient” for this vessel; see Glossary (Appendix 2)] filled with mercury,
and, after this water had cooled, peas, lentils, and seeds of Alisma plantago and
Polygonum amphibium. They gave no sign of germination when the amount of boiled
water was only seven to eight times the weight of the seeds, but when this amount
was 100 or 200 times greater, they germinated, and the elongation of their radicle
was proportional to the amount of water surrounding them. It is perhaps unnecessary
to explain the reason for these differences, or to recall that boiling does not deprive

1
Valmont de Bomare, Dictionnaire d’Histoire naturelle (article Café), made this phenomenon
known, but it has been ascribed, baselessly, to a true germination. Lentils that have lost their
germinative ability have the same property in a less striking manner.

J. F. Hill, Chemical Research on Plant Growth, 1

DOI 10.1007/978-1-4614-4136-6_1, © Springer Science+Business Media New York 2013
2 1 Influence of Oxygen Gas on Germination

water of all the oxygen gas that is contained in it in a dispersed or dissolved state,
and that in increasing the amount of fluid, I added enough air to elicit a very feeble
development in the seed. It has been claimed that peas that were swollen under water
[p. 4] germinated when they were then submerged in oil. I repeated this experiment
several times but was unable to obtain the reported result.
Humboldt found that seeds germinated in oxygenated muriatic acid [chlorine; see
Glossary] that was very diluted with water. This experiment, repeated in darkness
with strong oxygenated muriatic acid dissolved in boiled water and without contact
with external air, gave me the same results. It shows that it is not really necessary
that oxygen gas be free or uncombined to elicit germination. [A misinterpretation
by chemists of that day; chlorine, or “oxygenated muriatic acid,” does not contain
oxygen; see Glossary] But this acid is, up to now, at least according to my experi-
ments, the only substance that can produce this effect. Dilute nitric acid, to which
the same influence had been attributed, does not exercise it without contact with
external air. I likewise tried, unsuccessfully, sulfuric acid and several other acids
taken from the class of acids in which oxygen appears to be the least tightly bound.
I was no more successful in subjecting to the same tests, and without contact with
external air, a mixture of boiled water and different metallic oxides, such as black
oxide of manganese, red oxide of mercury, and red oxide of lead. [p. 5] I carried out
these experiments in glass flasks that I filled with liquid heated to the boiling point.
When the liquid had cooled to 30◦ Réaumur, I introduced the seeds. [In the Réau-
mur temperature scale, the freezing point of water is 0◦ , the boiling point 80◦ ; see
Conversions of Units of Measurement, Appendix 1] The flasks were then carefully
closed [see “closed”, in Glossary, for explanation] and plunged under mercury. The
amount of water used in each experiment was no more than seven or eight times the
weight of the seeds. The seeds tested were lentils, haricots, fava beans, peas, Alisma
plantago, and Polygonum amphibium.
A seed of fava, pea, or haricot, swollen by water without contact with oxygen gas,
shows a marvelous structure [“organization”] when its integuments are removed.
The microscope reveals, in the thinnest lamina that can be detached from the surface
of the cotyledons, a regular tissue formed of angular cells. This tissue is speckled,
on the back of the seed and on the edges of the cotyledons, with the openings or
glandular pores [presumably stomata] that produce such a beautiful effect on the
surface of mature leaves.2 They are [p. 6] very abundant on the opposing surfaces
of the cotyledons of pea seeds.
When seeds begin to germinate through contact with oxygen gas, the elongation
of their radicle beyond the length it could have attained by a purely aqueous swelling
is the only indication of their growth. But, other than the lack of a growth spurt, no
essential difference could be seen between the structure of a seed swollen by pure
water and the structure of a newly germinated seed. It is only by more prolonged
growth that the changes undergone by the seed become detectable to us. The seed

2
This observation, which I am merely confirming, was made by Sprengel (Anleitung zur Kenntniss
der gewachse, Vol. 1. p. 396). It shows, along with other, similar observations, that these openings
do not form through contact with air.
1.2 Use of Oxygen Gas in Germination 3

does not decay as it would if it had been deprived of contact with oxygen gas, but its
taste changes. It becomes bitter or sweet, according to the nature of the plant. The
cotyledons turn green through exposure to light and empty into the developing plant
by means of branched ducts that can be revealed by injected dye.

1.2 Use of Oxygen Gas in Germination [p. 7]

Our means are too limited to discover the causes of plant development, to see the
ultimate basis for their marvelous organization, and to explain how their growth
protects them from the decay or destruction to which they would seem to be destined
by the respective affinities of their elements. We have seen that oxygen gas was
necessary for germination to take place. We will limit ourselves to investigating
whether oxygen gas changes the seed by combining with it or rather by removing
some element from it. Oxygen gas could also act as a stimulant, as an irritant, but I will
not concern myself with this vague and hidden effect, which cannot be conclusively
tested here.
When a seed is placed in oxygen gas to germinate, the oxygen gas disappears
and is at the same time replaced by carbon dioxide [de Saussure used the older term,
“carbonic acid gas”, throughout the book]. Rollo,3 who observed these effects [p. 8]
in barley seeds, believed that the oxygen gas consumed was (1) for the most part
absorbed by the seed; [to de Saussure, the consumption of a gas meant its uptake,
whereas absorption meant assimilation into the plant’s dry matter] and (2) formed,
with the carbon of the seed, carbon dioxide. This opinion is only conjecture, not
based on any precise analysis.
It seemed to me that we could clarify what happened in this regard during germi-
nation only by comparing the amount of oxygen gas consumed with the amount of
carbon dioxide gas produced at the same time. If the amount of oxygen gas consumed
exceeds the amount entering into the carbon dioxide formed in the process, we could
conclude that the seed, in fact, absorbed oxygen gas, but if these two quantities are
always equal, we must infer that the oxygen gas was not absorbed by the seed but
was used only to remove carbon from the plant, by forming, with this carbon, carbon
dioxide gas.
I gave details of my research on this subject, in the Journal de physique, year
7 [of the French Republican calendar, i.e., 1799; see Note on the Translation]. I
carried out my research in bell-jars filled with ordinary air and closed by mercury. I
introduced only a very small amount of water, just enough [p. 9] to elicit germination,
in order that this fluid would not cause an appreciable loss [via solution in water]
of the carbon dioxide produced in germination. I performed these experiments on
seeds of pea, fava, haricot, barley, wheat [“blé”], lettuce, purslane, and garden cress.
They all provided me the result that I have communicated [presumably, in his paper
of 1799].

3
Annales de Chimie, Vol. 25, p. 37.
4 1 Influence of Oxygen Gas on Germination

Carbon, in combining with oxygen gas through combustion, should not apprecia-
bly change the volume of the oxygen gas, for we find, by calculating according to the
data of Lavoisier, that 100 cubic inches of carbon dioxide gas contain 49.67 grains of
oxygen, which are 98 cubic inches of oxygen gas.4 [Note: In the Paris measurement
system, which preceded the metric system and which de Saussure uses along with the
metric system in this text, the units of measurement, such as inch, pound, grain, differ
from the comparable English units; see Appendix 1.] Germinating seeds, as well as
burning carbon, do not appreciably change the volume of the oxygen gas that they
transform into carbon dioxide. If, for example, we germinate seeds [p. 10] in 100
cubic inches of ordinary air containing 21 cubic inches of oxygen gas and 79 cubic
inches of nitrogen gas [de Saussure used the term “azote”, which is still used for
nitrogen in French], we find that if germination produced 14 cubic inches of carbon
dioxide, only 7 cubic inches of free oxygen gas remain in their atmosphere; or if
7 cubic inches of carbon dioxide gas are produced, 14 cubic inches of oxygen gas
remain in the receptacle. The volume of oxygen gas consumed is thus equal to the
volume of carbon dioxide gas that is produced at the same time. This result, which
so far has admitted no exception, is an observation that should clarify for us the use
of oxygen gas in germination, whatever opinion one adopts on the proportions of the
constituent elements of carbon dioxide.
To observe the effects that I have communicated, the seeds must be in direct
contact with oxygen gas. If they are deprived of it by a layer of water or by being too
heaped up, they decay, and then yield hydrogen and carbon dioxide gases, whereas
in germinating, they lose only carbon.
Rollo, who saw that moistened seeds in a medium free of oxygen gas [p. 11]
yielded carbon dioxide, concluded that seeds develop this gas, or its two constituents,
when they are in the presence of oxygen gas, but there is no comparison, no analogy
to be drawn between a seed that germinates under the influence of oxygen gas and a
seed that decomposes in a place totally devoid of it.
The effect of oxygen gas on cotyledons, even when they remain underground, as
do those of favas and haricots, is useful to the growth of the plant after it germinates,
for, when these seeds are germinated in water, the plant that arises can thrive in air
only insofar as the cotyledons are above the surface of the liquid.
Seeds that germinate with the aid of water, under a receptacle full of pure oxygen
gas, produce, under equivalent conditions, much more carbon dioxide gas than under
a receptacle full of atmospheric air. Carbon dioxide, in all proportions, appears to be
detrimental to the initiation of germination. If moistened quicklime is placed under
the air-filled receptacle in which seeds are being germinated, in such a way that the
seeds are not in contact with this earth, the growth [p. 12] of the radicles is a little
accelerated. Carbon dioxide, however small the amount, added to the ordinary air
in which seeds are being germinated retards germination more than does a similar
amount of hydrogen gas or nitrogen gas.

4
Lavoisier found, in his experiment on the combustion of charcoal in oxygen gas, that the oxygen
gas underwent a small decrease, equivalent to around 5/100 of its volume. This difference arises
from the fact that charcoal cannot be entirely freed of hydrogen; therefore, the purer and drier the
charcoal that was used, the less the difference (Mémoires de l’Académie des Sciences, 1781).
1.3 Of the Changes that a Seed Undergoes Through Germination 5

I could see no difference in the time of germination of seeds placed at the same time
in ordinary air and in pure oxygen gas. Nor have I found that pea, wheat [“blé”],
or cress seeds placed in the open air on two sponges, one of which was wetted
with water strongly oxygenated by a compression pump and the other with boiled
water, germinated at different times. When I prolonged the experiment, keeping
the radicles immersed in the boiled and oxygenated waters, they always elongated
less in the oxygenated water. This effect may be attributed to two different causes:
one, that pure oxygen gas removes too much carbon from the seeds; the other, that
carbon dioxide, if more abundant, is harmful to their early development. In general,
carbon dioxide seems useful to plants only insofar as they can decompose it. Seeds,
before the [p. 13] development of their plumule, do not seem able to bring about this
decomposition.
It may be superfluous to note that seeds, depending on their species, require
different amounts of oxygen gas to begin germinating. The seeds of haricot, fava,
and lettuce consume, at equal weights, more of it than do peas, and peas more than
wheat [“blé”], barley, and purslane. It is very difficult to determine these amounts
accurately, by numbers, because there is always some uncertainty about the precise
time of germination. The amount of oxygen gas consumed to elicit the beginning
of germination appeared to me to be the same for haricot, fava, and lettuce, around
a hundredth part of their weight. The oxygen gas consumed by wheat [“froment”,
i.e., Triticum sativum], barley, and purslane seeds is equal to a thousandth or two
thousandth part of their weight. The carbon that these seeds lose at the same time is
only about a third of these amounts.
The amount of oxygen gas used by the same [kinds of ] seeds for germination
is, other things being equal, proportional to their weight and not to their number.
I chose four large favas whose weight was equal to that of 23 other, smaller ones.
I germinated [p. 14] the large and small favas at the same time, but separately, in
two receptacles closed by mercury. In this operation, the four large favas consumed
as much oxygen gas as the 23 other, smaller ones. A large seed thus requires more
oxygen gas to develop than does a small seed of the same species. The small seed can
germinate more deeply in the ground than the former. If experiment demonstrates
that large seeds develop at a depth where small ones perish, it is not because the
small ones do not germinate there [i.e., it is not because oxygen levels deep in the
ground are too low], it is because their plumules, which are weaker, cannot lift the
soil that covers them.

1.3 Of the Changes that a Seed Undergoes Through

Germination

In germination, we see no convincing evidence of the decomposition of water, for

the seeds emit neither hydrogen gas nor oxygen gas. Oxygen makes up part of the
carbon dioxide gas that the seeds produce, but the oxygen belonged to [p. 15] the
atmosphere of the seeds before germination.
6 1 Influence of Oxygen Gas on Germination

Seeds submerged in water or placed with this fluid in pure nitrogen gas emit
carbon dioxide, methane [de Saussure’s term for methane is “carburetted hydrogen
gas”], and nitrogen gas, but these emissions are of constituents that have separated
from the substance of the seed itself when it ferments. They are observed only when
the seed begins to putrefy, never during its growth or development, or even during
fermentation when that process occurs in direct contact with oxygen gas.5
[p. 16] Some seeds, when germinating in atmospheric air, pass from a mucilagi-
nous to a sweet state. Rollo noted that the seeds do not undergo this change of taste
in pure water or in oxygen-free environments, and analysis showed long ago that
sugar contains more oxygen than does mucilage. It was therefore concluded that
sugar does not form in a germinated seed unless the seed combines with the oxygen
gas in the atmosphere or decomposes the surrounding water.
These inferences or explanations are unnecessary, for the mucilage of seeds is
formed of hydrogen, oxygen, and carbon, and the proportion of oxygen in this
compound can as well be increased by the removal of another element (the carbon)
as by the addition of foreign oxygen gas.
When, with the aid of pure water in a closed vessel, I germinated any seed that had
been fully dried before the experiment, [p. 17] and then dried this same seed after its
germination [the second drying kills the seed], I always found that this germinated,
dried seed weighed less than the same seed before germination, although I added
back to it, by calculation, the carbon that the seed had lost in carbon dioxide either
during growth or during drying, and also the small amount of mucilage or extract
that it had left in the water in which it had developed.
This singular result can, I believe, be explained only by accepting that the seed has
lost water that was previously fixed in its own substance. It remains to be determined
whether this water is lost at the time of germination or after the death of the seed
during drying. It seemed to me that it was only during this last operation, because
when I doubled or tripled the growing time, the weight loss always remained the same,
whereas when I prolonged the drying time, the loss was always greater. Seventy-three
pea seeds, harvested 5 years earlier and placed for several weeks in a drying-oven
heated continuously to 20◦ Réaumur, [p. 18] had a combined weight of 200 grains.
They were placed, with five times their weight of distilled water, in a large flask filled
with atmospheric air, closed by a glass stopper, and inverted in mercury. I withdrew
these seeds after 2 days. They had almost all germinated, and during this development
they had formed 4-1/2 cubic inches of carbon dioxide, which contained, following

5
I say in direct contact because the contact of atmospheric air with the water in which dead seeds
are submerged singularly augments the development of hydrogen gas. Seeds placed under a little
water covered by a layer of oil, or without oil under a receptacle filled with a small amount of water
and sealed by mercury, can emit very little gas there, or at most three or four times their volume,
and the plant does not deteriorate noticeably. But if the mercury or oil is removed, the emission of
hydrogen gas is limited only by the almost complete disappearance of the seed. The mechanical
pressure of the oil or mercury could stop this release, but in my experiments it was not great enough
to produce this effect, because a column of water equal in weight to that of these external fluids did
not stop the emission of the gas. I believe that the carbon dioxide that the water retains in greater
quantity when the water is not in contact with the atmosphere serves as an antiseptic to the seed.
1.3 Of the Changes that a Seed Undergoes Through Germination 7

Lavoisier, 0.85 grains of carbon. The water remaining in the flask, evaporated at a
gentle heat, left a dry residue of 0.75 grains of mucilage or extract.
I dried these seeds on a flat vessel in the same drying-oven in which they had
germinated, and I examined, by comparative experiments made on other germinated
seeds, the changes that they caused in their atmosphere during drying. I found that
they produced no hydrogen gas and that they formed, as in germination, carbon
dioxide gas with the surrounding oxygen gas, without producing carbon dioxide
entirely from their own substance. The amount of carbon lost by the dead seeds
appeared to me less, during the same time and at the same temperature, than during
growth. They took 2 days to dry, or the same length of time as to germinate. [p. 19]
For brevity, I will suppose that the amount of carbon they lost in these two operations
was the same. The seeds, in drying, gave no indication of putrefaction or deteriora-
tion, either by odor or other products, or by external appearance. Their volume alone
diminished appreciably. These seeds, based solely on the results that I have reported,
should have had a dry weight of 200 − 0.85 × 2 − 0.75 = 197.5 grains. Yet I found
that they weighed only 189 grains. They could have lost only water, other than the
substances I have mentioned, and this lost water amounted to about 8 grains.
I repeated this experiment, at the same temperature, on identical seeds, leaving
them to grow or germinate in the flask for 4 days instead of 2. They lost very nearly
twice as much carbon, but only 8 grains of water, as in the preceding experiment.
The drying time remained the same. The amount of lost water increased when I dried
them for a longer time, exposing them to a lower temperature and in a more humid
place.
These experiments were repeated on haricots and favas, with similar results.
[p. 20] The dry, germinated seed weighed less than the dry, ungerminated seed,
and this difference was much greater than should result from the loss of pure carbon
and of extract.
As the seed, in drying and germinating, does not change the volume of its atmo-
sphere, and as the oxygen gas that the seed causes to disappear from the atmosphere
is recovered in an exactly equal amount in the carbon dioxide gas produced, we must
accept that atmospheric oxygen gas has no direct part in the formation of this water,
or, in other words, that the oxygen does not combine with the hydrogen of the plant
to form the water. The water arises entirely from the substance of the seed itself.
The seed thus loses, in the form of water, a part of its oxygen and hydrogen, and
this loss should increase the proportion of its other substances, particularly carbon.
Indeed, I found that 100 parts by weight of dried seeds, which had germinated in
a closed vessel in pure oxygen gas, contain more carbon than 100 parts of dry,
ungerminated seeds. Taking the mean of several observations, 100 pounds of pea
seeds yielded, by carbonization, 17-3/4 pounds of charcoal, [p. 21] whereas 100
pounds of the same seeds, dry, that had germinated in a closed vessel in oxygen gas,
produced 18 pounds of charcoal.
This loss of water and the resulting increase in the proportion of carbon occurs
only after the death of the plant. A seed growing in a closed vessel in an atmosphere
of pure oxygen gas does not lose water that was previously fixed; neither does it fix
8 1 Influence of Oxygen Gas on Germination

it. It only loses carbon. The direct effect of oxygen gas on the dead seed6 and on
the germinating seed is essentially the same. In both cases, it is limited strictly to
the removal of carbon. But the results are different: The dead seed loses fixed water,
namely, oxygen and hydrogen, and the germinating seed retains them.

1.4 Of the Influence of Light on Germination

Senebier is the first natural philosopher who has been led to admit that light is [p. 22]
harmful to germination. This opinion was confirmed by the comparative experiments
carried out by Ingen-Housz in the shade and in sunlight. Moreover, it has perhaps
also been confirmed by the practice of gardeners, who have found it advantageous
to protect their seedbeds from the direct effects of solar rays. [It is now known that
some seeds, especially small ones, require light to germinate].
It remains to be determined if the harmful effect of this star is attributable to its
heat, which could destroy the plant, or to the light alone, considered separately.
It has been believed that the influence of light was seen here, because the com-
parative experiments, in shade and sun, were performed at the same temperature
according to the thermometer. But it must be noted that this instrument, placed un-
der a receptacle in the atmosphere of the seeds, does not indicate the real heat that
the seeds experience on their surface due to the effect of solar rays. This heat is
so promptly dispersed by the surrounding bodies that it eludes our instruments. It
may be brought to the level of incandescence, as Rumford observed.7 A small plant
should be especially affected by it, [p. 23] as all its organs are conjoined in a smaller
space, it transpires less, and it decomposes less carbon dioxide.8 Now, experiment
shows that young plants transpire less, as they are less developed, and that young
leaves decompose, at the same volume, less carbon dioxide than do adult leaves.
I tried to germinate precisely weighed seeds, simultaneously, under two equivalent
receptacles, one opaque and the other completely transparent, but the seeds under the
transparent receptacle received only diffuse sunlight, such as reaches us through a
thick layer of clouds. The temperature was exactly the same in the two experiments,
according to the reading of a very sensitive thermometer. I could see no difference in
the time of germination of the seeds placed under these two vessels. When the trial
was prolonged on the germinated seeds, the seeds under the transparent receptacle
gained more weight and their growth was more vigorous and more advanced. The
plumules of the seeds [p. 24] in darkness were more slender and elongated due to
the effect of etiolation. I believe I can conclude from these experiments that there
is no evidence that light has, apart from the heat that accompanies it, a detrimental
influence on the germination and growth of young plants.

6
I am not speaking here of effects that occur in the final stages of putrefaction. I will return to those
later.
7
Essais politiques, économiques, et philosophiques, Vol. 2, p. 273.
8
The decomposition of carbon dioxide should produce cold, since composing it yields heat.
1.4 Of the Influence of Light on Germination 9

Lefebure9 notes that it is not by drying the seeds that the sun is detrimental to
their germination, because he observed that germination occurs more readily under
water in the shade than in sunlight. Although the reasons that I have given for the
difficulty of estimating the very intense heat that the surface of seeds placed in air in
the sun may acquire could be applied here, they are not the only cause of the results
obtained by that author. Growth under water may also be retarded in the sun because
the air that is disseminated in the water, and with whose aid germination could have
occurred in the shade, is partly disengaged through the influence of the solar rays.

9
Expériences sur la Germination des Plantes, by Lefebure, p. 136.
[p. 25] Chapter 2
Influence of Carbon Dioxide Gas on Plant
Growth

I would perhaps have followed a more methodical order if I had addressed the in-
fluence of oxygen gas on mature plants before proceeding to an examination of the
effect of carbon dioxide gas, which is only secondary and which is exerted only in
conjunction with oxygen gas, but a knowledge of the effects of oxygen gas assumes
preliminary data that force me to invert the natural order.

2.1 Influence of Carbon Dioxide Gas on Germination

Seeds do not germinate in pure carbon dioxide gas. A small amount of this gas (such as
one-twelfth), which, when mixed with atmospheric air, promotes the [p. 26] growth
of developed plants in the sun, is detrimental to germination and retards it, both in the
light and in the shade, more than does the same amount of hydrogen gas or nitrogen
gas. If we introduce, under a receptacle in which seeds have been germinated in
pure water and atmospheric air, some potash or a [i.e., another] substance capable of
absorbing the carbon dioxide gas that the seeds form with the surrounding oxygen gas,
their development is accelerated somewhat. It always seemed to me that germination
occurs sooner in moist sand or between two wet sponges than in humus, and humus
produces carbon dioxide gas. In general, this gas seems useful to plants only insofar
as they can break it down, and seeds, at the earliest stage of their development, do
not seem to carry out this decomposition appreciably. We note, however, that, as
germinating seeds produce too much carbon dioxide gas for us to deprive them of
it entirely, it is impossible to decide if its complete absence is harmful or helpful to
them.

2.2 Influence of Carbon Dioxide Gas on Developed Plants [p. 27]

When recently sprouted seeds are nourished with water lightly impregnated with
carbon dioxide gas, the water appears to be less advantageous to them than it is at
later stages of their growth. In two jars, one of which was filled with distilled water

J. F. Hill, Chemical Research on Plant Growth, 11

DOI 10.1007/978-1-4614-4136-6_2, © Springer Science+Business Media New York 2013
12 2 Influence of Carbon Dioxide Gas on Plant Growth

and the other with acidulated water,1 I floated two plates pierced with 24 holes, which
were intended to support the same number of pea seeds germinated in distilled water.
When the experiment began, their radicles were 6 millimeters (2 1/2 lines) long.
After 10 days, the roots in contact with the distilled water had elongated
1.3 decimeters (5 inches) more than had those [p. 28] in the gassed water. The stems
and leaves had developed in the same proportion. But when, after a month, the plants
nourished by the acidulated water were more developed, they no longer differed from
those that had grown with the aid of pure water and that had attained their maxi-
mum growth several days previously. The latter, in turn, were even surpassed by the
former, for, after 6 weeks, the pea plants in the gassed water had gained 46.4 grams
(12 gros + 10 grains), [“gros” is an obsolete French unit of measurement, equal to
1/8 French ounce; see Appendix 1] whereas those grown in pure water had gained
45.5 grams (11 gros + 66 grains).2 [Note: De Saussure made an error in decimal
placement here: He wrote 4.55 grams instead of 45.5. Also, he referred to footnote 1,
when he clearly meant footnote 2]. It is important to note that Senebier found that
young leaves decompose, at equal volume and in the same time, less carbon dioxide
than do adult leaves.
[p. 29] In the experiments just communicated, the stems of the plants grew in the
open air and received almost entirely through the roots the extra carbon dioxide that
was provided to them dissolved in water. It remains for me now to consider whether
this gas is useful to plants when it serves as their atmosphere.
Percival (Memoirs of the Manchester Society, Vol. 2) observed that a mint plant
nourished by water and exposed to a stream of atmospheric air mixed with carbon
dioxide fared better than did a similar plant exposed to a stream of pure atmospheric
air.
I tried to verify this initial observation and to determine the amount of carbon
dioxide that, when mixed with atmospheric air, may promote plant growth.
I germinated peas with the aid of water, until each plant had reached about
1 decimeter (4 inches) in height and weighed 1 gram (20 grains). I then placed three
plants for each experiment in a stemmed glass filled with water, such that the roots
alone were immersed in this liquid, and I presented them with various mixtures of or-
dinary air and carbon dioxide in receptacles closed by water that was [p. 30] covered
on the interior by a layer of oil if the receptacles contained more than half their vol-
ume of carbon dioxide. The three pea plants in each experiment had an atmosphere
of 990 cubic centimeters (50 cubic inches) and they did not displace 1/400 part of it.
Each day, for 5–6 hours, they received direct sunlight, which was moderated when

1
At first this water contained about a quarter of its volume of carbon dioxide, but as it was exposed
to the open air in the sun, it retained only a much smaller amount during the general course of the
experiment, which lasted 6 weeks. During this period, the gassed water was renewed four times.
2
Rückert (Annales de Crell) found that favas and violets planted in pots full of garden humus fared
better when they were sprinkled with water containing a third of its volume of carbon dioxide than
when they were sprinkled with distilled water. I did not find a detectable difference, in performing
the same experiment on wheat [blé]. It is possible that my humus, which was richer in manure,
provided the plants excess carbon dioxide.
2.2 Influence of Carbon Dioxide Gas on Developed Plants 13

too intense. At the same time and in the same place, I set up similar apparati exposed
to a weak and diffuse light. I call the latter manner of placement exposure to shade.

2.2.1 Results in Sunlight

The mean weight gain of the plants exposed to sunlight for 10 days in pure
atmospheric air was 425 milligrams (8 grains) per pea.3
[p. 31] These plants, at the same exposure, wilted as soon as they were in contact
with pure carbon dioxide.
They met the same fate in the atmospheres containing 3/4 and 2/3 of their volume
of carbon dioxide.
They grew for 7 days in the vessel containing 1/2 its volume of carbon dioxide.
After this period, they stopped growing.
The plants whose atmosphere contained 1/4 of its volume of carbon dioxide sur-
vived for the 10 days devoted to the experiment but they did not thrive. Each pea
gained only 265 milligrams (5 grains).
With 1/8 carbon dioxide, the mean gain was 371 milligrams (7 grains).
Finally, the mean gain of each plant in an atmosphere of ordinary air in which
carbon dioxide occupied 1/12 part was 583 milligrams (11 grains). I repeated this
experiment several times, and [p. 32] the plants consistently fared better than in
pure atmospheric air. The plants that grew in pure atmospheric air did not change it
perceptibly, either in purity or in volume, but those that grew in the artificial mixture
changed nearly all the carbon dioxide gas into oxygen gas.
I performed another experiment, which confirms this one and shows directly that
humus is useful to plants, not only through the nutrients that the plants may draw
from it through their roots, but also through the influence that humus has on the
atmosphere (influence that, as we know, consists largely of forming carbon dioxide).
I suspended, at the upper part of a receptacle containing about 3 liters (150 cubic
inches) of atmospheric air, 61 grams (2 ounces) of moistened humus. With this
receptacle, closed by water, I covered some partially developed pea plants, whose
roots were immersed in pure water during the experiment. After 10 days in the
sunlight, these plants, which did not touch the humus at all, had grown one-third
more than other, similar plants placed at the same time, without humus, under a
receptacle similar to the preceding one. [p. 33] But I must note that I renewed the air
of the receptacles twice in 24 hours, for without this precaution the plants growing
with the humus would have fared less well, either because the humus releases too
much carbon dioxide or because it produces vapors or miasmas that, in an unrenewed
air and through an unknown cause, are highly detrimental to plant growth.

3
This weight gain was due largely, and maybe even entirely, to the introduction of liquid water, that
is, water of vegetation, into the leaves that developed during the experiment and that drew their solid
substance from the cotyledons, which were still large and attached to the plant. These cotyledons
contained three or four times less water of vegetation than the leaves to whose development they
contributed. I will return elsewhere to this subject, which is unrelated to what occupies me here.
14 2 Influence of Carbon Dioxide Gas on Plant Growth

2.2.2 Results in the Shade

In the apparatus exposed to shade, the smallest amount of carbon dioxide added to
ordinary air was detrimental to plant growth. In the atmosphere containing 1/4 of its
volume of carbon dioxide, the plants were dead after the sixth day. They survived, at
the same exposure, for 10 days in an atmosphere in which carbon dioxide occupied
1/12 part, but their gain there was only 159 milligrams (3 grains), whereas it was
265 milligrams (5 grains) in pure atmospheric air.
We have just seen that carbon dioxide added artificially in very small proportions
to the atmosphere of plants [p. 34] is useful to their growth in the sun, but it exerts
this beneficial effect only insofar as this atmosphere contains free oxygen gas. Thus,
plants that can maintain their growth in nitrogen gas die in it even in the sun when
it is combined with the proportion of carbon dioxide gas that would have promoted
their development in atmospheric air.

2.3 Elaboration of Carbon Dioxide Gas by Leaves is Necessary

to Their Growth in the Sun

The experiments that I have detailed on the growth of peas in pure atmospheric air
yielded the same results when I washed this air in limewater and thus removed the
very small amount of carbon dioxide that this air contains naturally. But the results
were very different when I introduced into the atmosphere of the plants a substance
that absorbs the carbon dioxide that the plants contribute to forming. I suspended, at
the top of the receptacles covering the peas, 7 or 8 grams (2 or 3 gros) of lime slaked
with water [p. 35] and then dried quickly by the heat of boiling water. I rested the
openings of these receptacles on saucers filled with limewater.
From the second day, the atmosphere of the plants exposed to the sun in this
apparatus decreased in volume. The third day, the lower leaves began to turn yellow,
and between the fifth and sixth days, the stems died or were completely defoliated.
The atmosphere of the plants, examined at this time, was found to be corrupted
[“vitiated”]. It contained no more than 16/100 oxygen gas. [Ordinary air is approxi-
mately 21 % oxygen, 79 % nitrogen, by volume]. Peas that had grown at the same time
without lime, under receptacles filled with ordinary air, had changed the air neither
in purity nor in volume, and all their parts were healthy and vigorous. We see by
the experiment with lime that there was absorption [by the lime], and consequently
formation of carbon dioxide gas, for the absorbing substance acted only on this gas.
We see, further, that the presence, or rather, the elaboration, of carbon dioxide is
necessary to plant growth in the sun.4 Finally, we [p. 36] find that when we do not

4
One might think that the withdrawal of the part of the atmospheric oxygen gas retained by the
carbon dioxide in the lime was the reason why growth ceased, but developed pea plants can survive
in pure nitrogen gas. Lime and potash exert their full deleterious influence on marsh plants, which
thrive as well in pure nitrogen gas as in atmospheric air.
2.3 Elaboration of Carbon Dioxide Gas by Leaves is Necessary . . . 15

see carbon dioxide production by plants growing without lime in ordinary air, it is
because they decompose it as they form it with the surrounding oxygen.
In the shade I obtained a different result. Not only did the plants not die in the
receptacle containing the lime and limewater, but they fared better there than in a
similar receptacle lacking these substances.
The mean weight gain of each plant growing with the lime was 371 milligrams
(7 grains) in the period of 10 days. The air of the receptacle contained 3/100 carbon
dioxide after the experiment. But in the ordinary air without [p. 37] lime, each pea
acquired [on average] only 212 milligrams (5 grains). Limewater showed 11/100
carbon dioxide in this atmosphere.
It is evident from these results that one cannot judge the effect of the complete
deprivation of carbon dioxide on plant growth in the shade because too much of this
gas is produced in this case for the lime to absorb it all as it is formed, but that the
effect of a partial deprivation is to promote growth.
For the same reason, lime does not cause plants that are growing in the sun
in an atmosphere of pure oxygen gas to lose their leaves. This atmosphere has a
superabundance of carbon dioxide gas that the alkaline earth did not have time to
absorb as quickly as it was formed.
The above observations were made only on plants growing in pure water, and
it was important to make sure that the same results would be obtained with plants
rooted in plant mold. But this earth could not be placed under a receptacle because the
earth produces too much carbon dioxide for the lime, which acts only at a distance,
to be able to remove it before its elaboration by the plant.
I put 31 grams (1 ounce) of the same lime that I used in the preceding experiments
into a glass globe. [p. 38] I moistened the lime slightly to remove any doubt as
to its dehydrating properties. Into the globe, which had a capacity of about 4 liters
(200 cubic inches), I then placed a woody branch,5 covered with leaves, that was
exposed to the sun and whose roots were in plant mold. I took care that the leaves
touched neither the lime nor the walls of the globe, whose neck was carefully luted
[cemented] to the branch. I fitted a similar apparatus, but without lime in the globe,
to a branch located next to the preceding one. This branch retained its freshness for
more than 2 months, but it was not so for the branch growing with the lime. Its leaves
remained green for 12 days, then began to dry and, after 3 weeks, they had all fallen.
The branch was not dead. A month later, it sprouted new leaves in the globe, which
[p. 39] had not been unsealed. But at this time, the lime no longer had an effect on
the surrounding air. Its surface was saturated with carbon dioxide. I withdrew it and
found that it effervesced with acids.

I must note that quicklime and potash have no very noticeable effect on the growth of succulent
plants, because, with their very thick parenchyma and their epidermis that is less porous than that
of other plants, they retain carbon dioxide more tenaciously. For the same reason, the stems of all
plants are much less affected than are the leaves in these experiments.
5
The plants on which I performed these experiments were honeysuckle (Lonicera caprifolium),
plum (Prunus domestica), privet (Ligustrum vulgare), and peach (Amygdalus persica).
16 2 Influence of Carbon Dioxide Gas on Plant Growth

This new growth shows that the leaf shedding was due not to deprivation of oxygen
gas, which could be retained in the lime by the absorption of carbon dioxide gas, but
only to the absence of carbon dioxide. If the effects occurred more slowly in these
experiments than in the previous ones, it is partly because the plants rooted in soil
were not deprived of the effects of the carbon dioxide that they received from the
plant mold through the roots, but only of the external effects of this gas on the leaves.
[Although de Saussure says here that plants obtain some carbon via carbon dioxide
in the soil, he generally considers the main source of plant carbon to be atmospheric
carbon dioxide.]

2.4 Of the Decomposition of Carbon Dioxide Gas by the Green

Parts of Plants

Priestley was the first to recognize that leaves have the property of improving air
corrupted by combustion or respiration, but he did not trace the cause of this phe-
nomenon. Senebier discovered that leaves [p. 40] decompose carbon dioxide by
appropriating its carbon and eliminating its oxygen. He noted that fresh leaves ex-
posed to the sun, in spring water or water lightly impregnated with carbon dioxide
gas, produced oxygen gas as long as some carbon dioxide remained in the water.
He saw that when this gas was used up, and when leaves in distilled water were
exposed [to the sun], they did not produce more air than could be interposed in their
own volume. But no one has yet analyzed the effects of the decomposition of carbon
dioxide, or seen if the amount of oxygen gas eliminated is greater than, less than,
or equal to the amount in the carbon dioxide gas. The following experiments were
intended to answer this question. I will go into this subject in long and minute detail,
but without it the results would be almost meaningless.

2.4.1 First Experiment

2.4.1.1 On Periwinkle (Vinca minor L.)

I composed, from carbon dioxide gas and ordinary air that [p. 41] the phosphoeu-
diometer [see “eudiometer” in Glossary and discussion in footnote 9 of Introduction
to the translation] showed to be 21 hundredths oxygen, an artificial atmosphere
occupying 5.746 liters (290 cubic inches [original text erroneously says 290 cubic
centimeters]). Limewater showed it to be 7-1/2 hundredths carbon dioxide. [Note:
The amount of carbon dioxide in ordinary air is so small that virtually all the carbon
dioxide de Saussure measured in these experiments is the added gas.] The gaseous
mixture was confined in a receptacle closed by mercury that was wetted, or covered
with a very thin layer of water, to prevent contact of this metal with the air sur-
rounding the plants, for I have observed that such contact, as reported by the Dutch
chemists, is harmful to plant growth in prolonged experiments.
2.4 Of the Decomposition of Carbon Dioxide Gas by the Green Parts of Plants 17

Under this receptacle I placed seven periwinkle plants, each 2 decimeters

(8 inches) tall. They displaced a total of 10 cubic centimeters (1/2 cubic inch). Their
roots were immersed in a separate vessel containing 15 cubic centimeters (3/4 cubic
inch) of water. The amount of this liquid under the receptacle was insufficient to ab-
sorb appreciable carbon dioxide, especially at the ambient temperature, which was
never less than + 17◦ Réaumur [about 21 ◦ C, or room temperature].
This apparatus was exposed to direct sunlight for 6 consecutive days, from 5 to
11 o’clock in the morning, and shaded whenever the light became too intense. On
the seventh day, I removed the plants, [p. 42] which had not deteriorated in the least.
Their atmosphere, all corrections made [presumably for temperature and pressure],
had not changed in volume, at least as far as one could judge in a receptacle of
1.3 decimeters (5 inches) diameter, in which a difference of 20 cubic centimeters
(1 cubic inch) is almost imperceptible; but the error can be no greater than this.
Limewater showed no remaining carbon dioxide gas. The eudiometer indicated
24-1/2 hundredths oxygen gas. I set up a similar apparatus with pure atmospheric
air and the same number of plants at the same exposure. This atmosphere changed
neither in purity nor in volume.
From the eudiometric observations noted above, it follows that, before the
experiment, the mixture of ordinary air and carbon dioxide contained:

4199 cubic centimeters (211.92 cubic inches) of nitrogen gas

1116 cubic centimeters (56.33 cubic inches) of oxygen gas
431 cubic centimeters (21.75 cubic inches) of carbon dioxide gas
5746 cubic centimeters (290 cubic inches)

After the experiment, the same air contained:

4338 cubic centimeters (218.95 cubic inches) of nitrogen gas

1408 cubic centimeters (71.05 cubic inches) of oxygen gas
0 cubic centimeters (0 cubic inches) of carbon dioxide gas
5746 cubic centimeters [290 cubic inches—but de Saussure omitted this figure]

[p. 43] The periwinkles therefore elaborated, or removed, 431 cubic centimeters
(21-3/4 cubic inches) of carbon dioxide gas. Had the plants eliminated all of the
oxygen gas from it, they would have produced a volume of oxygen equal to that of
the carbon dioxide that disappeared. But they released only 292 cubic centimeters
(14-3/4 cubic inches) of oxygen. Therefore they assimilated 139 cubic centimeters
(7 cubic inches) of oxygen in decomposing the carbon dioxide, and produced
139 cubic centimeters (7 cubic inches) of nitrogen gas [an error; plants do not release
nitrogen gas.]
A comparative experiment showed me that the seven periwinkle plants that I used
had a dry weight of 2.707 grams (51 grains) before the decomposition of the carbon
dioxide, and yielded, through carbonization by fire in a closed vessel, 528 milligrams
(9.95 grains) of charcoal. The plants that decomposed the carbon dioxide were dried
and carbonized by the same technique and yielded 649 milligrams (12.23 grains)
of charcoal. The decomposition of the carbon dioxide therefore caused a gain of
120 milligrams (2.28 grains) of charcoal.
18 2 Influence of Carbon Dioxide Gas on Plant Growth

I similarly carbonized periwinkles that had grown in atmospheric air freed of

carbon dioxide, and I found that the [p. 44] proportion of their carbon had decreased
rather than increased during their stay under the receptacle.

2.4.2 Second Experiment

2.4.2.1 On Aquatic Mint (Mentha aquatica L.)

The mixture of ordinary air and carbon dioxide that served as atmosphere for two
mint plants, each 3.5 decimeters (13 inches) tall, and together displacing 10 cubic
centimeters (1/2 cubic inch), occupied 6.5 liters (328 cubic inches). Limewater indi-
cated 7-1/4 hundredths carbon dioxide. Before the addition of this gas, the ordinary
air contained 21 hundredths oxygen gas. The arrangement of the apparatus was the
same as in the previous experiment.
After 10 days the plants had elongated by 1 decimeter (4 inches) and had sprouted
long roots, but the volume of their atmosphere had not changed. Limewater indicated
no more than 2-1/2 hundredths carbon dioxide gas in the artificial mixture at this time.
After removal of the carbon dioxide, this atmosphere contained 23-1/2 hundredths
oxygen gas.
[p. 45] Ordinary air without mixture, in which two mint plants were grown at the
same time, did not change, either in purity or in volume.
The mints in the preceding experiment therefore removed 309 cubic centimeters
(15.6 cubic inches) of carbon dioxide gas. They eliminated 224 cubic centimeters
(11.26 cubic inches) of oxygen gas from it. They retained 86 cubic centimeters (4.34
cubic inches) of oxygen gas in elaborating the carbon dioxide gas, and replaced the
oxygen gas absorbed by a nearly equal amount of nitrogen gas.
Through carbonization, I found that these plants had increased their charcoal
content in this experiment, and that this increase was no greater in those that had
grown under a receptacle filled with pure atmospheric air.

2.4.3 Third Experiment

2.4.3.1 On Loosestrife (Lythrum salicaria)

The mixture of atmospheric air and carbon dioxide gas used for this experiment
occupied 1.486 liters (75 cubic inches). Limewater indicated 10/100 carbon dioxide
in it. Before the addition [p. 46] of the carbon dioxide, the ordinary air contained
21/100 oxygen gas. The loosestrife displaced 2.8 cubic centimeters (1/7 cubic inch).
The arrangement was the same as in the previous experiments. The receptacle that I
used here was 9 centimeters (3.5 inches) in diameter, and I could not be mistaken by
more than 5 cubic centimeters (1/4 cubic inch) in the estimation of volumes.
2.4 Of the Decomposition of Carbon Dioxide Gas by the Green Parts of Plants 19

After 7 days devoted to this experiment, not a single leaf of the loosestrife had
turned yellow. The atmosphere had decreased by 10 cubic centimeters (1/2 cubic
inch). It then no longer contained any carbon dioxide gas, and the eudiometer
indicated 27-1/4 hundredths oxygen gas in it.
Another loosestrife, grown at the same time under the same conditions, in pure
atmospheric air, changed the air neither in purity nor in volume.
Based on the eudiometric results reported above, the plant that stayed in the arti-
ficial mixture removed 149 cubic centimeters (7-1/2 cubic inches) of carbon dioxide
gas, and eliminated from it 121 cubic centimeters (6.13 cubic inches) of oxygen gas.
It assimilated 27 cubic centimeters (1.37 cubic inches) of oxygen gas from the car-
bon dioxide gas, [p. 47] and it produced 21 cubic centimeters (1.1 cubic inches) of
nitrogen gas.

2.4.4 Fourth Experiment

2.4.4.1 On Pine (Pinus genevensis)

The mixture of ordinary air and carbon dioxide occupied 5.549 liters (280 cubic
inches). Limewater indicated 7/100 carbon dioxide. For 18 days, I left in it a young
pine, 2.4 decimeters (9 inches) tall and 10 cubic centimeters (1/2 cubic inch) in
volume. After this period, the atmosphere had decreased by 39 cubic centimeters
(2 cubic inches), as far as I could estimate in a receptacle that was 1.6 decimeters
(6 inches) in diameter.
The eudiometers indicated 1-1/2 hundredths carbon dioxide gas there, and after
this was removed, 24-1/4 hundredths oxygen gas.
Another pine, grown for the same time under a receptacle filled with pure
atmospheric air, made no detectable change in it.
The plant in the artificial mixture removed 306 cubic centimeters (15-1/2 cubic
inches) of carbon dioxide gas. The plant disengaged [p. 48] 246 cubic centimeters
(12-1/2 cubic inches) of oxygen gas and retained 60 cubic centimeters (3 cubic inches)
of oxygen in the decomposition of the carbon dioxide. Finally, the plant produced
20 cubic centimeters (1 cubic inch) of nitrogen gas.

2.4.5 Fifth Experiment

2.4.5.1 On Raquette (Cactus opuntia)

The mixture of ordinary air and carbon dioxide gas occupied 3.012 liters (155 cubic
inches), and limewater indicated 10/100 carbon dioxide gas. The cactus displaced
22 cubic centimeters (1-1/10 cubic inches). It remained for 8 days under the recepta-
cle, exposed to the sun’s full intensity. I had moderated this effect for the other plants,
20 2 Influence of Carbon Dioxide Gas on Plant Growth

which would have been harmed without this precaution, but here there was not the
same risk, and without strong light, the carbon dioxide would have decomposed too
slowly.
When I withdrew the plant, the volume of its atmosphere had not changed de-
tectably. The eudiometers indicated 4/100 carbon dioxide gas in it, and after the
removal of this gas, 24/100 oxygen gas. A cactus identical to the [p. 49] preced-
ing, grown for the same time in an equal volume of pure atmospheric air containing
21/100 oxygen gas, made no detectable change.
The plant therefore removed 184 cubic centimeters (9.3 cubic inches) of carbon
dioxide gas from the artificial mixture. It eliminated from this gas 126 cubic cen-
timeters (6.4 cubic inches) of oxygen gas. The plant assimilated 57 cubic centimeters
(2.9 cubic inches) of oxygen gas in decomposing the carbon dioxide and replaced
the oxygen gas absorbed by a nearly equal amount of nitrogen gas.
It follows from all these experiments that plants, in decomposing carbon dioxide
gas, assimilate a part of the oxygen that it contains.

2.5 Plants Grown with Pure Water in the Open Air Take
Carbon from the Small Amount of Carbon Dioxide Gas
that Occurs Naturally in Our Atmosphere

The previous observations show that plants in closed vessels decompose [p. 50]
carbon dioxide gas when it is mixed with atmospheric air in much larger proportions
than occur naturally.
It is now appropriate to see whether plants carry out this decomposition in the open
air, which contains hardly more than 1/500 carbon dioxide by volume. Hassenfratz,
in a memoir on plant nutrition (Annales de Chimie, Vol. 13), sought to establish
that plants growing in pure water and the open air increase in volume with the aid
of the water alone, and that after they develop they contain an absolute amount of
carbon less than that which was present in the seed. On this subject, I did several
experiments, which yielded results contrary to those of this author. I will report two
examples.
First Experiment I immersed the roots of several peppermint plants (Mentha
piperita) in bottles filled with distilled water, and I grew these plants in the sun,
on an exterior window sill protected from the rain. By drying some similar plants
that had been uprooted at the same time and place, [p. 51] I ascertained6 that 100 parts
by weight of those that I grew in distilled water contained 40.29 parts of dry plant
substance, from which 10.96 parts of charcoal were withdrawn by carbonization.7
The 100 parts of mint, after 2-1/2 months of growth in the open air, had a green
weight of 216 parts, but this weight gain revealed nothing at that point, since it

6
The absolute weight of the plants grown in distilled water was 7.6 grams (3 gros).
7
For the procedure followed in this operation, see the note at the end of Chap. 5.
2.5 Plants Grown with Pure Water in the Open Air Take Carbon . . . 21

could have been due to the addition of water of vegetation, which always occurs
in plants when they are transplanted to a place that is wetter than the one in which
they had grown previously. By drying them at atmospheric temperature, they were
reduced to a weight equivalent to 62 parts. The plants had therefore increased their
dry substance, with the aid of air and water, by 21.71 parts. These 62 parts yielded,
by carbonization, 15.78 parts of charcoal, or 4.82 parts more than they would have
yielded before having grown in the distilled water. When I grew the same [kind of ]
plants [p. 52] under similar conditions in a weakly lit place, I found that they lost a
small amount of their carbon. Lack of light may be the reason for the results obtained
by Hassenfratz.
Second Experiment I placed four fava beans weighing 6.368 grams (120 grains)
among flint pebbles in glass capsules. I wetted them with distilled water. After
3 months of growth in the sun, under the open sky, the fava plants, immediately after
flowering, had a fresh weight of 87.149 grams (1,642 grains). They were reduced by
drying to 10.721 grams (202 grains). Thus they had almost doubled their dry plant
matter by growing in the open air. These plants yielded, by carbonization in a closed
vessel, 2.703 grams (51 grains) of charcoal. But four fava beans of the same weight
as those used in this experiment yielded 1.209 grams (22-3/4 grains) of charcoal.
The favas had thus more than doubled the amount of carbon in their substance by
developing with the aid of water in the open air, [p. 53] and there is no doubt that they
accomplished this by the decomposition of the carbon dioxide that they found in the
atmosphere, for we saw in the previous section that plants grown under receptacles
filled with air that was pure and not renewed did not increase their carbon.

2.5.1 Further Remarks on the Decomposition of Carbon Dioxide

Gas by Plants

Ingen-Housz8 observed that green plants to which he had applied, in darkness, an ar-
tificial heat that was ostensibly the same as that received from the sun by other, similar
plants yielded an impure air, while those exposed to the sun produced the opposite
effect. He concluded that the production of oxygen gas by the sun-exposed plants
was attributable to the light alone, abstractly considered, and not to the accompany-
ing heat. But this conclusion may be premature, because the heat that accompanies
light, [p. 54] and dark heat, act in very different ways in these experiments.
Luminous heat warms only the plant, due to the plant’s opacity. This heat hardly
warms the plant’s surrounding medium at all, because of the medium’s transparency.
The medium removes from the plant some of the heat that the sun conveys to it and
moderates an effect that, considered by itself, could destroy the plant.
Dark heat warms the transparent fluid and the plant to the same degree. The plant
is harmed because the medium surrounding it can no longer rid it of the heat that has
been conveyed to it.

8
Experiences sur les végétaux, Vol. 1, p. 39 [de Saussure cited p. 36, but it should have been p. 39],
and Vol. 2, p. 167.
22 2 Influence of Carbon Dioxide Gas on Plant Growth

It is very likely that plants in atmospheric air, without the intervention of light,
decompose a part of the carbon dioxide gas that they form themselves with the
surrounding oxygen gas. But this effect cannot be shown directly. I have seen marsh
plants, such as Polygonum persicaria and Lythrum salicaria, emit oxygen gas into
an atmosphere of nitrogen gas in a low, diffuse light, but they never produced this
effect in [p. 55] total darkness. It may be presumed, however, that plants decompose
carbon dioxide gas there, based on the following observations. I grew some peas,
loosestrifes, and fleabanes [“inules”] in profound darkness, beneath two identical
receptacles filled with atmospheric air. I renewed [presumably means replaced] these
plants each day so that they would not suffer harm. These experiments were done in
duplicate. One of the receptacles contained quicklime or potash, the other neither of
these. After 4 or 5 days, both atmospheres were corrupted, but I always found that
the receptacles provided with lime or potash contained less oxygen gas than those
that lacked alkali, and one imagines that it is because these plants found less carbon
dioxide gas to decompose in the receptacles that lacked lime or alkali. [This seems to
be a mis-statement; the plants should have found less carbon dioxide to decompose
in the receptacles that were provided with lime or alkali.]
Another effect also seems to indicate the decomposition of carbon dioxide in
darkness. It is the light green hue assumed by etiolated leaves that develop there.
Acording to Senebier, this color is a result of the decomposition of carbon dioxide.
[Senebier was mistaken; the decomposition of carbon dioxide requires the green
pigment plus light.] But as these signs are very weak and very indirect, we will not
generally accept, [p. 56] in the following, that the decomposition of carbon dioxide
can occur anywhere but in the light.
Senebier recognized that non-green plant parts, such as wood, roots, most petals,
white blotches on leaves, and leaves that have turned completely red or yellow in
autumn through deterioration of their juices, do not exhale oxygen gas. It is not
necessary, however, to infer from this that green color is essential to the plant parts
that decompose carbon dioxide gas, or a necessary result of this decomposition, for
the variety of Atriplex hortensis in which all the green parts are replaced by red or
dark purple parts produced for me, under spring water, in 5 or 6 hours, seven or eight
times its volume of oxygen gas, which contained only 0.15 of its volume of nitrogen
gas. The variety of the same plant having green leaves did not provide me oxygen gas
that was any purer or more abundant. [Wrong conclusion; the red or purple pigment
in the vacuoles of the mesophyll cells merely masks the green of the chlorophyll in
the chloroplasts, where photosynthesis takes place as usual.]
Not all kinds of leaves are able to decompose [p. 57] carbon dioxide gas to the same
degree. In several experiments, Lythrum salicaria was able in one day to decompose
up to seven or eight times its volume of it. Cactus opuntia and other succulents could
decompose only a fifth or a tenth of this amount. Without wishing to identify all the
causes of these differences, I will note that the green parts decompose carbon dioxide
in proportion to their surface area, but hardly at all in proportion to their volume.
Very thin leaves, and ones that are deeply cut and thread-like, decompose the most
carbon dioxide, under equivalent conditions. Fleshy plants, stems, and fruits, which
present little surface area, decompose, at the same volume, much less gas. [Further,
2.6 Summary 23

plants using the Crassulacean Acid Metabolism (CAM) pathway, which include many
succulents, absorb and store carbon dioxide at night and close their stomata by day,
so there is less gas exchange in the daylight; see Translator’s Introduction.].
Leaves, in exhaling oxygen gas, always release nitrogen gas. They produce nitro-
gen in appreciable amounts only in the light, and nearly in proportion to the carbon
dioxide that they decompose. [An error: plants do not release nitrogen gas.] Leaves
that can tolerate a very humid atmosphere the longest without damage are the ones
that produce the purest oxygen gas. Marsh plants are among these. [p. 58] What-
ever plant is placed in the unnatural conditions of these experiments, the oxygen gas
produced at the beginning of the experiment is always less corrupted than that at the
end. These observations indicate that the nitrogen gas arises from the very substance
of the plant. I am inclined to believe that this gas, which has an affinity for pure
oxygen gas, is removed in the interior of the leaf by oxygen gas in greater quantity
the more listless the growth is. It is understandable why leaves develop much less
nitrogen gas in atmospheric air to which carbon dioxide gas has not been added. It is
because the leaves there have much less contact with pure oxygen gas. The oxygen
gas contained by atmospheric air is saturated with the nitrogen gas that occurs there
naturally.

2.6 Summary

Carbon dioxide gas added in certain proportions to atmospheric air promotes plant
growth, but only insofar as the plants can decompose this gas. It is detrimental to
germination in the same proportions in which it is useful to developed plants. It
promotes the growth [p. 59] of developed plants in the sun and becomes detrimental
to them in darkness.
The presence, or rather the elaboration, of carbon dioxide gas is indispensable to
the growth of green plant parts in the sun. Sun-exposed plants die when the carbon
dioxide gas that they form with the surrounding oxygen gas is removed.
Green plants growing with the aid of distilled water in open atmospheric air
acquire a large amount of carbon there.
Plants, in decomposing carbon dioxide, assimilate a part of the oxygen contained
in the carbon dioxide.
[p. 60] Chapter 3
Influence of Oxygen Gas on Developed Plants

3.1 Of the Changes that Oxygen Gas Undergoes Through its

Contact with Leaves

In Chap. 6, I will treat the growth of green plants in environments previously depleted
of oxygen gas. I will show that plants develop there only because they emit this gas
there. The chemical results of the gas’s effects on green, herbaceous parts of plants
are the focus of my present research.
Some healthy leaves, cut after a serene summer day and placed for one night
under a receptacle filled with atmospheric air, caused the surrounding oxygen gas
to undergo changes that seemed to vary [p. 61] according to the nature of the plant.
1. The leaves of oak (Quercus robur), horse chestnut, false acacia, Sedum reflexum,
and most plants diminish the volume of their atmosphere. Its oxygen gas disap-
pears and it forms a volume of free carbon dioxide gas that is smaller than the
volume of oxygen consumed during the experiment.
2. The leaves or green parts of Cactus opuntia, Crassula cotyledon, Sempervivum
tectorum, Agavé americana, and Stapelia variegata diminish the volume of their
atmosphere by absorbing oxygen without forming appreciable carbon dioxide. In
this case, as in the previous one, it seems at first glance that the reduction in the
volume of the atmosphere is due to the assimilation of oxygen by the plant.
In order to observe, consistently, a condensation in the air around the leaves in
darkness, the leaves must be perfectly healthy and occupy between a 7th and a 20th
of the volume of the air in which they are enclosed. If they occupy a smaller space,
the effects will sometimes be [p. 62] undetectable; if they occupy a larger space, their
environment will be too depleted of oxygen for them to thrive. They must also be
placed in the experiment immediately after being cut at sunset and must not remain
under the receptacle more than 12 hours. Since my plants did not wither in any of
these trials, I cannot believe that plant deterioration affected my results. I should not
suppose that it did, especially in the case where no carbon dioxide was formed, for
all wilting leaves produce a certain amount.

J. F. Hill, Chemical Research on Plant Growth, 25

DOI 10.1007/978-1-4614-4136-6_3, © Springer Science+Business Media New York 2013
26 3 Influence of Oxygen Gas on Developed Plants

We may ask whether the oxygen gas absorbed by healthy leaves at night is as-
similated effectively by them for augmenting their dry matter, for we will see that,
if those leaves that absorbed this gas are exposed to the sun, they restore an approx-
imately equal amount of the gas, and that if the leaves have enough growth force to
remain healthy for several consecutive days under the same receptacle, they dimin-
ish their atmosphere each night and augment it each day in approximately the same
proportion.
[p. 63] I give to these alternating effects the names expiration and inspiration.
These expressions are perhaps inappropriate because they indicate in plant organs a
contraction and expansion that has not yet been established by observation. I accept
these designations only because the gaseous imbibition and emission by plants pro-
duces, in the final analysis, results analogous in some ways to those of respiration
in animals, although the appearance and the means are very different. Anatomical
comparision of plants and animals is misleading, but if only their overall physiolog-
ical traits, such as nutrition, secretions, reproduction, and the influence of oxygen
gas or respiration on their existence, are considered, without regard to the means by
which these functions are carried out, a striking analogy must be admitted between
these beings.
On matters that do not lend themselves to rigorous proof, accurate accounts of
observations are more instructive than are general conclusions, which often depend
on the observer’s point of view. For this reason, I will detail my experiments [p. 64]
on Cactus opuntia. Their results can be applied, with slight, easy-to-conceive, mod-
ifications, to the leaves of other plants and even to all herbaceous, green plant parts.
But these parts do not all have enough growth force to withstand the same tests.

3.2 Experiments on Inspiration [this section title supplied

by translator]

3.2.1 Inspiration by Cactus opuntia

I suspended, after sunset, some branches or leaves of cactus displacing 119 cubic
centimeters (6 cubic inches) in a receptacle containing 951 cubic centimeters
(48 cubic inches) of atmospheric air freed of its carbon dioxide. There was no water1
in this vessel, which was closed by mercury.
[p. 65] The next day at sunrise I found, after correcting for changes in temperature
and pressure, that the atmosphere of the plant had diminished by 79 cubic centimeters

1
Here is an example of the tremendous growth force of this plant. A cactus branch that had served
for 3 weeks in trials similar to those that I have just reported, either in the sun or in the shade, was
placed in darkness, without water or soil, in a cupboard for 14 months. There it endured cold of
−8◦ Réaumur in winter, and heat of +22◦ in summer. At the end of this period, it was wrinkled and
thinned from the loss of half of its water of vegetation, but it had sprouted stems and roots all over
its surface. No part of it had lost its growth force. When it was taken out of there, it produced, under
distilled water, in the sun, a third of its volume of an air containing 34/100 oxygen gas and 66/100
nitrogen gas. And when I planted it in vegetable mold, all its former substance swelled, recovered
its original dimensions, and continued to produce new shoots.
3.2 Experiments on Inspiration [this section title supplied by translator] 27

(4 cubic inches). I then examined this air. It contained only 14/100 oxygen, whereas
before the introduction of the cactus, it had contained 21/100 of this gas. Limewater
showed not an atom of carbon dioxide in the remaining air [presumably because
the plant retained the carbon dioxide it had formed with the surrounding oxygen
gas during the night]. Further, I will say that when I placed some limewater beside
the cactus under this receptacle for one night, no calcium carbonate [“carbonate
of lime”] was formed. It follows from the eudiometric observations communicated
above that the atmosphere of the plant contained, approximately:

Before inspiration 198 cubic centimeters (10 cubic inches) oxygen gas
After inspiration 119 cubic centimeters (6 cubic inches) oxygen gas
Inspiration 79 cubic centimeters (4 cubic inches) oxygen gas

[p. 66] The reduction in volume of the atmosphere during the night was thus
precisely equal to the reduction in oxygen gas. The plant therefore did not appreciably
absorb nitrogen gas, but only an amount of oxygen equal to three-quarters of its
volume. Not all cacti make such large inspirations. Inspirations during a night often
amount to only half the volume of the plant. These differences are due not only to
the condition of the plant, which absorbs more the more vigorous it is, but also to the
temperature at which the experiment is conducted. Inspiration is greater in a given
time at a temperature of 20◦ or 25◦ Réaumur than at 10◦ or 15◦ .
If these plants are kept in darkness for more than one night, they continue to
absorb oxygen, but always more slowly, until they contain about 1-1/4 times their
own volume of it. After this time, which arrives in 36 or 40 hours they neither
decrease nor increase the volume of their atmosphere as long as there is free oxygen
in it, whatever the capacity of the receptacle and the duration of the experiment, even
if it is prolonged for a month or until the death of the plant.

3.2.2 Leaves Saturated with Oxygen Gas Form Carbon Dioxide

Gas in Darkness [p. 67]

A cactus that is saturated with oxygen gas by inspiration is not, however, without an
effect on its surroundings. When the plant is at the point that it can absorb no more
oxygen, it begins to form, without deteriorating in the least, free carbon dioxide
gas from its own carbon and the surrounding oxygen. This combination, as I have
observed several times, does not change the volume of the atmosphere. It is noteworty
that leaves thus saturated consume, in a given period, by the formation of carbon
dioxide, around half as much oxygen as they consumed by inspiration.
Most leaves, particularly nonsucculent ones, form free carbon dioxide at the same
time as they inspire oxygen. They generally make lesser inspirations, but never larger
than that of the cactus.
28 3 Influence of Oxygen Gas on Developed Plants

3.2.3 A Cactus that has Inspired Oxygen Gas Cannot Expire it

Through the Effect of the Vacuum of an Air Pump

The oxygen gas inspired by a cactus or any other leaf is retained in the leaf by an
[p. 68] attraction so powerful that suppression of the weight of the atmosphere does
not release the gas appreciably.2 Six cubic inches of cactus that, in one night, had
inspired 4 cubic inches of oxygen were able to expire, in darkness, under a small
amount of water in a vacuum, only 1 cubic inch of air, containing 15/100 oxygen gas
and 85/100 nitrogen gas, and no or 1/100 carbon dioxide gas. The preceding day,
before the inspiration, this plant had yielded the same amount of air by this means,
but the air had been composed of 19/100 oxygen gas and 81/100 nitrogen gas.
The small difference in purity seen in the airs extracted before and after inspiration
supports the idea that the air contained in the leaves is always related to the purity of
the surrounding atmosphere. As the air in the vessel where the plant had stayed in the
dark was corrupted by inspiration, so it was in the plant. It is very likely that [p. 69]
the air pump obtains little except the free air present in the plants. I have always found
it, at all hours and exposures, to be less pure than atmospheric air, in plants growing
in the open air. When a cactus is completely saturated with oxygen gas, the effect of a
vacuum will extract from it an air containing two or three hundredths carbon dioxide
gas, which does not amount to a 50th of the oxygen that the plant has inspired.
A dark heat, moderate enough that it did not destroy the plant, was no more
effective than a vacuum at releasing the inspired gas. Six cubic inches of cactus that
had absorbed 4 cubic inches of oxygen gas at 15◦ Réaumur could not expire any
gas in the dark, under receptacles filled with atmospheric air heated to either 30◦ or
35◦ . In both cases, they only combined their carbon with the surrounding oxygen,
without changing the volume of their atmosphere. They turned yellow at 40◦ and
the results then became meaningless. When I placed, in the shade, some oxygen-
saturated leaves, in water heated to the preceding temperatures, they released an
amount of air equal to a sixth of their volume. [p. 70] This air contained 89/100
nitrogen gas and 11/100 carbon dioxide gas.

3.2.4 Leaves do not Inspire Appreciably in Gaseous Environments

that Lack Free Oxygen

A cactus (or any other leaf) cut at sunset and placed for one night in nitrogen gas,
hydrogen gas, or pure carbon dioxide gas does not decrease the volume of its atmo-
sphere. On the contrary, it expands the volume by adding carbon dioxide gas to it.
This increase, which is less the more vigorous the plant, ordinarily amounts, in one
night, to a third or a fourth of the volume of the cactus. There is no doubt, however,

2
Using this procedure, I removed from green fruits, such as pears and apples, even before they
ripened, an air containing a very large amount of carbon dioxide, but leaves never yielded a
significant amount, unless they had stayed in environments devoid of oxygen gas.
3.2 Experiments on Inspiration [this section title supplied by translator] 29

that these atmospheres penetrate in very small amounts into the interior of the plant,
for when I subjected cactuses that had passed the night in pure hydrogen gas to the
effects of the air pump, they yielded an amount of gaseous fluid equal to a sixth of
their volume. This air was composed of 40 parts hydrogen gas, 40 parts nitrogen gas,
and 20 parts carbon dioxide gas.
A cactus cut at sunset and [p. 71] placed for one night in hydrogen gas does not,
as I just said, make any detectable inspiration. But when it is removed from this
atmosphere and placed in darkness in a receptacle filled with ordinary air, it inspires
its own volume of oxygen gas, or very nearly the same amount as it would have if it
had not been in the hydrogen gas.
If carbon dioxide gas is substituted for hydrogen gas in this experiment, the plant,
as soon as it is placed in darkness in a receptacle filled with ordinary air, begins to
expand its atmosphere, emitting to it a small amount of carbon dioxide, equivalent
to around a sixth of the plant’s volume. The plant then inspires an amount of oxygen
gas about equal to the preceding emission, after which it changes the volume of its
atmosphere no further. It modifies the atmosphere only by combining its carbon with
the surrounding oxygen gas. This plant can remain 2 or 3 days in pure carbon dioxide
gas without perishing.

3.2.5 A Cactus Inspires Carbon Dioxide Gas in the Same

Proportion as Oxygen Gas, When the Carbon Dioxide
is Mixed with the Oxygen in a Small Amount [p. 72]

The experiments that I reported in Sect. 3.2.1, on the inspiration of oxygen gas,
were done in an air that had previously been freed of carbon dioxide gas, and the
question arises whether, in natural, atmospheric air, leaves absorb carbon dioxide
gas in preference to oxygen. Consequently, I placed a cactus under a receptacle
filled with ordinary air to which I had added carbon dioxide gas, and I found that
the carbon dioxide and oxygen gases were absorbed in the same proportion, but
that the inspiration was more rapid than it would have been if there had been no
carbon dioxide gas. Here are the details of one of these experiments: 153.6 cubic
centimeters (7-3/4 cubic inches) of cactus were placed after noon in 43-1/2 cu-
bic inches of air confined by mercury. This air, which I set equal to 100 parts,
contained 74 parts nitrogen gas, 19 parts oxygen gas, and 7 parts carbon dioxide
gas. After 12 hours in darkness, the volume of air was reduced to 81 parts. The
leaves had thus made an inspiration [p. 73] equal to around 1-1/4 times their vol-
ume [actually, closer to their exact volume; 1-1/4 times would be 9.69 cubic inches,
whereas this inspiration is 8.265 cubic inches]. I found by analyzing the remain-
ing air that the cactus had inspired 13-1/2 parts oxygen gas and 5-1/2 parts carbon
dioxide gas. They had thus inspired the two gases in the same proportion, or 3/4 of
each one. After this inspiration they were saturated. They could not continue to
reduce their atmosphere through a prolonged darkness.
30 3 Influence of Oxygen Gas on Developed Plants

3.2.6 A Cactus Never Appears Saturated with Oxygen Gas After

Staying for Several Days in Darkness, or an Indefinite Time
in the Open Atmosphere

I have indicated in Sects. 3.2.1 and 3.2.4 that a cactus placed in darkness in atmo-
spheric air, under a receptacle closed by mercury, absorbs in the period of 30 or
40 hours 1-1/4 times its volume of oxygen gas, [in Sect. 3.2.4, he does not give a
figure of 1-1/4; he says “its own volume”] and that after that time it does not reduce
the volume of its atmosphere further, however long the experiment. Based on this
observation, we should expect that a cactus cut at the same time as the previous one
and exposed to darkness for several days in the open air or outside a receptacle would
not diminish its atmosphere when subsequently placed, without water in darkness,
under a receptacle filled with atmospheric air. But [p. 74] in these circumstances
the plant makes an inspiration equal to a quarter of its volume in 24 hours. It is then
saturated and inspires no more. If the cactus is taken out of confinement and exposed
to darkness in the open air, and then returned to darkness under the receptacle, it
makes an inspiration equal to the previous one. By repeating the same trials several
times, I was able to make the same leaf absorb in a fortnight, in continuous darkness,
several times its volume, or an indefinite amount, of oxygen gas. In the following
[i.e., in Sect. 3.2.8], I will give an explanation of this phenomenon, which is common
to all plants or at least to all their succulent parts.

3.2.7 A Cactus Whose Structure has been Destroyed Makes no

Appreciable Inspiration in Atmospheric Air

Six cubic inches of cactus collected at sunset were cut into pieces and quickly reduced
to a paste by pulverization. Immediately afterwards, they were placed, for one night,
under a receptacle filled with atmospheric air closed by mercury. They did not change
the volume of their atmosphere. They did not absorb oxygen gas. But they [p. 75]
combined their carbon with this gas, forming carbon dioxide gas, and the juice of the
plant coagulated. The amount of oxygen gas consumed in the formation of carbon
dioxide gas in this experiment was much less3 than the amount that would have
disappeared through inspiration, for a healthy cactus would have consumed, under
conditions otherwise the same, at least 3 or 4 [cubic] inches of oxygen, whereas the
cactus whose structure had been destroyed removed only a quarter of this amount in
the formation of carbon dioxide gas.
A cactus reduced to paste and placed immediately afterwards either in pure nitro-
gen gas or hydrogen gas responds as an intact cactus does. It expands these gases by
emitting carbon dioxide gas. At later stages, it adds hydrogen and nitrogen gases.

3
This assertion is true only in the period immediately after the death of a plant. When it putrefies,
it consumes much more oxygen gas than through inspiration.
3.2 Experiments on Inspiration [this section title supplied by translator] 31

3.2.8 Inferences from the Preceding Observations: The Oxygen

Gas Inspired by Leaves in the Dark is Converted into Carbon
Dioxide Gas [p. 76]

It is very likely that the oxygen gas absorbed by cactus and by leaves in general
is retained in their parenchyma in the form of carbon dioxide gas. Although this
view has not been demonstrated by direct experiments, we must accept it because it
appears to be the only explanation for the principal facts that we have just observed.
The ability of leaves to expire and inspire oxygen gas appears to depend on their
ability to decompose carbon dioxide gas. The green parts of plants, which are in
general the only parts that carry out this decomposition, are also, as will be seen in
the following, the only ones that make successive inspirations and expirations through
the effects of sunlight and darkness. These two functions appear to be alternately
cause and effect, one of the other, as the fall of an oscillating pendulum seems to be
the cause of its ascent and this ascent the cause of a new fall.
A leaf that is placed in darkness [p. 77] immediately after being exposed to sunlight
contains no carbon dioxide gas because that gas was decomposed in the light. Air
freely penetrates the leaf, as we saw hydrogen gas do (Sect. 3.2.4). Atmospheric oxy-
gen gas is seized in passing by the carbon of the plant and forms carbon dioxide gas,
which loses its gaseous [“elastic”] state and its volume through its union with the wa-
ter of vegetation. Due to plant structure, the carbon dioxide undergoes a compression,
which is limited since the plants that absorb the most gas cannot acquire more than
1-1/4 times their volume of it. The effect of the oxygen gas thus only apparently
changes before and after saturation of the leaf. Before saturation (Sect. 3.2.1), the
gas appears to be absorbed, as long as the leaf can condense or absorb the carbon
dioxide gas produced by this oxygen gas; and after saturation (Sect. 3.2.2), the
oxygen gas forms free carbon dioxide gas because the leaf, being saturated, can
accept no more carbon dioxide gas.
But why are leaves never saturated with carbon dioxide gas when they have been
in darkness for an unlimited time in the open air (Sect. 3.2.6)? Why does alternating
exposure under a receptacle and in open air cause them to absorb [p. 78] many
times their volume, or an indefinite amount, of carbon dioxide gas in continuous
darkness? I respond that the latter phenomenon is only an illusion, produced by the
attraction that the open atmospheric air exerts on the carbon dioxide contained in the
leaves. Atmospheric air has a real chemical affinity for carbon dioxide gas [this is a
misinterpretation], and, like most combinations, has limits of saturation. It may not
be superfluous to recall here the evidence.
When pure water is placed under a receptacle filled with carbon dioxide gas, this
water becomes saturated, and it takes up neither more nor less than its own volume of
it. This water, saturated and confined in flasks with different amounts of atmospheric
air, sheds its gas in proportion to the amount of air contained in each flask, and the air
expands in proportion to the amount of carbon dioxide gas that it removes from the
water. If it is a question of destroying this gaseous combination, the substances used
will remove more or less carbon dioxide gas, according to their degree of affinity for
it. Limewater will remove more than distilled water, and potash more than limewater.
32 3 Influence of Oxygen Gas on Developed Plants

These effects and their causes have been indicated by Berthollet in his fine [p. 79]
memoir on affinities. (Mémoire de l’Institut National, sciences et arts, Vol. 3).
Pure atmospheric air, not being in equilibrium of saturation with a leaf that is satu-
rated, that is, contains 1-1/4 times its volume of carbon dioxide, removes only a small
amount of the leaf’s carbon dioxide. [This is confusing; the next sentence suggests
that the pure air should remove more than just a small amount of carbon dioxide from
the leaf.] Therefore we see that, as soon as a cactus that is saturated with oxygen
or carbon dioxide gas is put under a new receptacle filled with pure atmospheric
air, the atmospheric air undergoes an immediate increase in volume (Sect. 3.2.4),
due to an emission of the carbon dioxide gas that saturates the cactus. After this
emission, the plant forms, with the surrounding oxygen gas, new carbon dioxide
gas that it can inspire, because the saturated atmospheric air poses no more obstacle
to saturation of the leaf. When the leaf is removed from its confinement, the open
atmospheric air removes from the leaf the part of the carbon dioxide that the leaf
just acquired and puts the leaf in a condition to absorb anew, under a receptacle, that
which it has lost to the open air. The leaf thus does not accumulate carbon dioxide
or oxygen gas in this experiment.4
[p. 80] We cannot compare the phenomena produced in darkness by a plant that
enjoys all its growth capabilities in atmospheric air with those that it shows in oxygen-
free environments (Sect. 3.2.4).
In the latter case, plant growth remains suspended. All the effects that the plant
could produce as an organized body are interrupted. The plant begins to putrefy, to
decompose, producing from its own substance the two elements of carbon dioxide.
The water of vegetation, which is less than the weight of the plant, then imbibes this
gas only in very small amounts, because it is not retained by the pressure that the
growth force can exert on it. We have no idea of the limits of this compression. The
experiments [p. 81] of Hales show that the compression may exceed the weight of
our atmosphere. It is for this reason that the air pump was unable to extract the carbon
dioxide that the cactus had absorbed (Sect. 3.2.3). The disorganized cactus did not
inspire carbon dioxide gas or oxygen gas appreciably because this compression no
longer existed. The very small amount of carbon dioxide required to saturate the
water of vegetation under these conditions had been formed and absorbed in the act
of pulverization.
It generally follows from these considerations that leaves in darkness do not
assimilate the atmospheric oxygen gas that they have absorbed, at least insofar as they
do not decompose the carbon dioxide that results from this absorption. [He is presum-
ably saying that if they decomposed carbon dioxide, they would assimilate a small

4
The affinity that I have assumed between atmospheric air and carbon dioxide is contested by
Dalton. But let us confine ourselves to experiment and note that when 1 cubic inch of water is
introduced into a mixture composed of 90 cubic inches of atmospheric air and 10 cubic inches
of carbon dioxide, or any other amount, and this water is withdrawn, exposed to the open air in
darkness, and put back into the gaseous mixture, and this maneuver is repeated several times with
the same water, the atmospheric air is almost entirely depleted of the 10 cubic inches that had been
added to it. Thus the water produced precisely the same effect here as the cactus: It appeared to
have absorbed ten times its volume of carbon dioxide, although it was clear that the water never
accumulated one-tenth of this amount.
3.3 Experiments on Expiration [this section title supplied by translator] 33

amount of the resulting oxygen.] If this decomposition occurs in these circumstances,

it is only in a proportion that escapes any direct experiment. These conclusions would
doubtless be risky if they were based only on the results that cactus provided me,
but I confirmed them on more than 60 other plants of very different kinds. Several of
them consumed more oxygen gas in the same time, leaving more free carbon dioxide
gas in their atmosphere, but, in [p. 82] darkness that was prolonged indefinitely,
none was able to diminish the volume of this atmosphere by an amount greater than
the volume of the plant, and the oxygen gas that disappeared was always, with the
exception of this small difference, directly proportional to that which entered into
the composition of the carbon dioxide gas produced.

3.3 Experiments on Expiration [this section title supplied

by translator]

3.3.1 Expiration by Cactus in Atmospheric Air

The 119 cubic centimeters (6 cubic inches) of cactus that were used in the experiment
in Sect. 3.2.1, and that had corrupted and diminished their atmosphere during a single
night by inspiring 79 cubic centimeters (4 cubic inches) of oxygen gas, were placed
without water in another receptacle and exposed to the morning sun, without being
in contact with the walls of the receptacle or with the liquid that closed it. This air
contained 951 cubic centimeters (48 cubic inches) of atmospheric air freed of carbon
dioxide gas. In the evening, this atmosphere had increased by 87 cubic centimeters
(4.4 cubic inches). It contained no carbon dioxide gas. The eudiometer indicated
27-1/4 hundredths oxygen gas there, whereas before the introduction of the plants,
[p. 83] it had indicated only 21 hundredths. [Even though the cactus is a CAM plant,
it is releasing gas during the day.] It follows from these observations that the air of
the receptacle contained:

Before inspiration 200 cubic centimeters (10.1 cubic inches) of oxygen gas
+751 cubic centimeters (37.9 cubic inches) of nitrogen gas
After expiration 283 cubic centimeters (14.28 cubic inches) of oxygen gas
+755 cubic centimeters (38.1 cubic inches) of nitrogen gas
Difference, or expiration 83 cubic centimeters (4.18 cubic inches) of oxygen gas
+4 cubic centimeters (0.2 cubic inch) of nitrogen gas

The following night, I placed these same leaves in a new atmosphere of ordinary
air. There they inspired 74 cubic centimeters (3-3/4 cubic inches) of oxygen gas,
and the next day, in the sun, they expired 79 cubic centimeters (4 cubic inches) of
oxygen gas and 6 cubic centimeters (1/3 cubic inch) of nitrogen gas. When these trials
were continued for 7 days, the inspirations and expirations of oxygen gas steadily
decreased and the expirations of nitrogen gas steadily increased. I found, by adding
all the inspirations on the one hand and all the expirations on the other, that the same
cactus inspired in 7 nights 331 cubic centimeters (21-3/4 cubic inches) [conversion
not right; 21-3/4 cubic inches = 431 cubic centimeters; 331 cubic centimeters = 16.7
34 3 Influence of Oxygen Gas on Developed Plants

cubic inches; the correct figure seems to be the 21-3/4 cubic inches] of oxygen
gas, and expired, in the sun, during the 7 intervening days, 584 cubic centimeters
(29-1/2 cubic inches) of air [p. 84] composed of 23-1/4 cubic inches of oxygen
gas and 6-1/4 cubic inches of nitrogen gas. The final expiration was more than half
nitrogen, whereas the first yielded only 1/16 of its volume of this gas.
In this experiment, the emissions of pure oxygen gas exceeded the inspirations
[he wrote “expirations” here but presumably meant “inspirations”] a little, but the
difference is too small, and too insignificant compared with the volume of the plant,
to allow us to conclude anything about the decomposition of water.5

3.3.2 Expiration by Cactus Under Distilled Water

and in Nitrogen Gas

I varied the experiments on expiration by submerging, during the day, under a

receptacle filled with distilled water, a cactus that had passed the night without water
in a receptacle filled with ordinary air depleted of carbon dioxide. The general results
were the same [p. 85] as in the previous experiment. In this case, though, the plant,
since it was under water that was cooler than the air and, moreover, in an uncongenial
medium, carried out all its functions more slowly. It did not have time to decompose,
in one sunny day, all the carbon dioxide that it had formed during the night. Moreover,
its expirations were corrupted by a considerable amount of nitrogen gas, whereas
the earlier expirations [i.e., those in the preceding experiment] in ordinary air were
of almost pure oxygen gas and were never, in this regard, less than the inspirations.
Six cubic inches of cactus absorbed, in 7 nights, 17 cubic inches of oxygen gas, and
expired into [original text erroneously says “inspired” into] the water, in the form of
bubbles, 19 cubic inches of air that, according to an analysis with potassium sulfide
[“hydrosulfide of potash”], was composed of 13.3 cubic inches of oxygen gas and
5.7 cubic inches of nitrogen gas. The first expiration contained 80/100 oxygen gas and
20/100 nitrogen gas. The seventh was composed of 39/100 oxygen gas and 61/100
nitrogen gas. We cannot help but see that the nitrogen arises from the decomposition
of the plant. The plant produces it in a significant amount only when it is suffering
damage, either from the [p. 86] length of the experiment or from an uncongenial
medium.
A cactus that passes the night in atmospheric air and the day in nitrogen gas in the
sun makes larger expirations than it does under water, but less pure than in ordinary
air.

3.3.3 Expiration is Proportional to Inspiration

First Proof When cactus plants have inspired all the oxygen gas they can absorb
by being held for 36 or 40 consecutive hours in darkness under a receptacle filled

5
I will report other results on this subject in Chap. 7. The experiments will be much lengthier. The
plants will be nourished, in atmospheric air, by a small amount of water. They will not be exhausted
by the lack of this nutrient or plagued with repeated transfers.
3.3 Experiments on Expiration [this section title supplied by translator] 35

with atmospheric air, and they are then exposed to the sun, they make a much larger
expiration in 7 or 8 hours than when they have spent only a single night in ordinary
air. I performed this experiment several times, and in a manner that leaves no doubt.
I will give an example here: 6 cubic inches of cactus, having inspired 4 cubic inches
of oxygen gas during 12 hours in darkness, expired [original text erroneously says
“inspired”] immediately afterwards, in the sun, in 7 hours, 4.2 cubic inches of the
same gas. I then placed the same plant under a receptacle in the dark [p. 87] for
36 consecutive hours. It absorbed 7-1/2 cubic inches of oxygen gas. Next it was
exposed to the sun, and it exhaled, in 7 hours, the 7-1/2 cubic inches that it had
inspired in the previous 36 hours. The expiration is thus augmented by the amount
of the inspiration. I believe this result important because it shows that the emission
of oxygen gas, which is an effect of the inspiration, should not be confused with an
emission that might be attributable to the decomposition of water.
Second Proof I exposed to the sun, in atmospheric air freed of carbon dioxide gas,
some cactus plants that had been cut the previous evening and had passed the night
in either pure nitrogen gas or pure hydrogen gas. They exhaled oxygen gas into the
atmospheric air, but about half as much as they would have produced if they had
passed the night in a receptacle filled with ordinary air. The following night, I placed
these plants again in nitrogen gas, and the next day in atmospheric air. They produced
half as much oxygen gas as the day before. The third day they produced still less. The
fifth day they were dead and [p. 88] putrefying. The total oxygen gas that they were
able to yield in the sun in this entire experiment was less than their volume. The result
was the same when the plant was nourished with a small amount of water and, after
passing the night in nitrogen gas, was exposed under water to the sun. Cactus thus can
produce only a very limited amount of oxygen gas, less than its own volume, when it
has passed the night in a medium lacking oxygen gas and the day in atmospheric air.
I obtained very different results when I performed the converse experiment,
namely, placing the cactus during the night in a receptacle filled with atmospheric air
and during the day in a different receptacle, filled with nitrogen gas. I could then con-
tinue the experiment for a fortnight or more without the cactus wilting. The amount
of oxygen gas that the plant produced exceeded five or six times the volume of the
plant. The amount was, as it were, unlimited. These observations confirm the heading
of this section. Moreover, they prove that it is only during the night that leaves espe-
cially require contact with oxygen gas. [In CAM plants, the stomates are generally
open only at night.] The following inference can be drawn from this: When plants
must [p. 89] stay in a place that is dimly lit and in which the atmosphere is always
more or less corrupted, as in some greenhouses, they should be arranged so that, the
darker the place they occupy, the more frequently the air they receive is renewed.

3.3.4 Effect of Quicklime or Potash on Expiration by Cactus

Quicklime or potash, which will kill thin leaves growing in the sun in a few days if
it is placed in their atmosphere, does not have this effect on cactus or succulents in
36 3 Influence of Oxygen Gas on Developed Plants

general because, with their very thick parenchyma and less-porous epidermis, they
more tenaciously retain the carbon dioxide gas that they form. If limewater is put
under a receptacle full of atmospheric air in which a cactus passes a single night, no
calcium carbonate [“carbonate of lime”] is formed. But it is different if limewater is
placed in contact with the air in which a cactus is staying in the sun. The limewater
becomes covered with carbonate. [But stomates of CAM plants are usually closed in
the sun.] In this case, and especially when dry quicklime slaked with water [p. 90]
or a solution of potash is placed under the receptacle, the expiration of oxygen gas
is noticeably reduced. It is then never greater than the inspiration.

3.3.5 The Results Obtained on Expiration by Cactus are

Applicable to the Leaves of Other Plants

Experiments on expiration can give results that are dramatic and can withstand
scrutiny only with green plants that contain a large amount of green herbaceous
matter in a small volume and that have a strong enough growth force that they can
displace, in the sun, a tenth or a twentieth of the capacity of the receptacle without
crumpling and without touching the vessel walls, which are then warm enough to
destroy a plant that presses on them. Succulents are almost the only plants that are in
this category. Thin leaves are so delicate, so expanded, that they can grow in the sun
under a receptacle only if the plant bearing them displaces a 400th or 500th of the
receptacle’s capacity. They can be shown to be subject to the same laws, however, by
comparing the following two experiments: (1) If a thin-leaved plant, [p. 91] such as
a mint, an epilobe, or a Lythrum salicaria, is grown for a fortnight, in the alternating
effects of sunlight and night, under a receptacle full of ordinary air, this atmosphere
will be found to have changed neither in purity nor in volume after the allotted time.
These plants occupy too little space for the changes that they should produce there
to be appreciable. A cactus of the same volume and under equivalent circumstances
would not have produced more marked effects. (2) If the growth of thin leaves is
maintained in two receptacles, one designated for the stay of the plant in the sun and
the other for its stay in darkness, we find, after the successive transfers, that at the
end of a fortnight the air of the receptacle kept in darkness decreased in purity and
volume and that of the one in the sun improved.6 The opposing effects are too small
[p. 92] for their respective amounts to be very accurately compared, but the general
results cannot be doubted, and we can understand based on the first experiment that
the improvement obtained in the second is due to the oxygen gas transported by the
plant from the atmosphere in darkness into the atmosphere in sunlight.

6
This trial was performed by Ingen-Housz with cress plants (Lepidium sativum). Experiences sur
les Vegétaux, Vol. 2, p. 144. But this author did not investigate the cause of the result he obtained.
He states that the improvement is greater than the deterioration. I repeated this experiment on the
same plant. The deterioration of the air appeared to me to be greater than its improvement. My
eudiometric methods were more precise than his.
3.5 Relative Amounts of Oxygen Gas Consumed in Darkness by Different Leaves 37

Thin leaves always leave significant amounts of free carbon dioxide gas in their at-
mosphere in darkness. Due to their large surface area, part of the gas that they contain
is removed from them by the surrounding air more abundantly than in fleshy plants.

3.4 Plant Growth in Pure Oxygen Gas

Although all plants require oxygen gas in order to develop, they fare less well in the
shade when their atmosphere is composed only of oxygen than when the oxygen is
mixed with a certain amount of nitrogen or hydrogen gas. The latter gases do not, by
themselves, seem to have any noticeable effect on plant growth. Most plants do not
assimilate them if the gases are isolated or in a [p. 93] gaseous state. But the gases
can be useful to a plant by reducing its points of contact with oxygen gas.
In most of the experiments that I performed on partly developed pea plants (Pisum
sativum), the plants, during 10 days in the shade, with the aid of water and of
nourishment provided by their cotyledons, gained half as much weight in pure oxygen
gas as in ordinary air confined by receptacles. In pure oxygen gas, the plants formed
much more carbon dioxide gas, which, in itself, is always harmful to plant growth in
the shade. Moreover, the excess oxygen gas could be harmful to them by removing
too much of their carbon.
When the pea plants were exposed, in pure oxygen gas, to the direct effects of
solar rays, their weight gain was almost the same as in ordinary air, but their stems
seemed to be longer and thinner in the ordinary air. The shortened form of the
plants in the oxygen gas cannot be attributed with certainty to the purity of this gas,
however, because it seemed to me that this effect occurred in an artificial atmosphere
composed in the same proportions as ordinary air, [p. 94] with the oxygen gas
drawn from manganese [i.e., manganese dioxide], and the nitrogen gas from beef
muscle by nitric acid. The eudiometer revealed the same composition as for our
atmosphere.7

3.5 Relative Amounts of Oxygen Gas Consumed in Darkness

by Different Leaves

I tried to determine whether the amount of oxygen gas that different leaves of the same
volume consume, or remove, in darkness, either by inspiration or by the formation
of free carbon dioxide gas, is related to other circumstances of their growth.

7
Does our atmosphere contain substances that elude the eudiometer and that favor plant growth?
Or do artificial gases retain some ingredient of the substances from which they were extracted and
that inhibits this elongation? I cannot decide this. It is certain that well-washed nitrogen gas has
an animal odor if it was extracted from beef muscle, and a sulfurous odor if it was obtained from
ordinary air by a sulfide.
38 3 Influence of Oxygen Gas on Developed Plants

The leaves of succulent plants [p. 95] consume less oxygen gas than do most other
leaves. They retain it more tenaciously, or, in other words, they leave less free carbon
dioxide gas in their atmosphere. These effects occur because these leaves offer the
surrounding air many fewer points of contact, and their surface is, other things being
equal, pitted with many fewer pores, as microscopic examination reveals. Because
succulent plants growing under the influence of daylight and night-time can retain
almost all the carbon dioxide they form in darkness, they may lose only a very small
amount of their carbon and consequently can endure the lack of this nutrient for a
longer time, or require less of it through the roots for their growth. [Here is another
example of de Saussure’s view that plants do not obtain all of their carbon from
the atmosphere.] These plants grow naturally in sand, clay, or barren soil. Because
they consume very little oxygen, they can grow in a rarefied atmosphere. The large
number of sedum, saxifrages, and sempervivum growing in the mountains provide
examples. [Not a correct interpretation]
The leaves of deciduous trees generally (though there are numerous exceptions)
[p. 96] lose the most carbon, or consume the most oxygen gas. On high mountains,
we see that the dominion of trees ends long before that of herbs.
The leaves of evergreen trees consume less oxygen gas than do the leaves of
deciduous trees. Evergreens such as pines, juniper, and rhododendron can grow in a
sterile soil and in a rarefied atmosphere.
Marsh plants grow in an environment in which they are deprived of free access
to oxygen gas by the surrounding vapors. Experiment shows that, under equivalent
conditions, they consume less oxygen gas than do most other plants having herba-
ceous stems. From this, we can understand why herbaceous plants of the mountains
are often found in marshes of the plains.
We see that, in general, the amount of oxygen gas that leaves remove is related
to the site. Leaves growing in an unproductive soil, or in a rarefied atmosphere, or
in low, humid places consume, under equivalent conditions, less oxygen gas [p. 97]
than do leaves that grow only in a fertile soil, with ample access to atmospheric air.
I am attaching here the table of the observations from which I deduced these
results. The experiments were done in elongated receptacles closed by mercury that
was lightly wetted on its surface. The receptacles contained 991 cubic centimeters
(50 cubic inches) of atmospheric air and 20 cubic centimeters (1 cubic inch) of
leaves. The nonfleshy leaves filled, at this volume, almost the entire capacity of the
receptacle. I was obliged to keep them in the experiment for 24 hours in the dark
because, in a shorter time, several of them would not have produced, at the volume
that they occupied, effects appreciable enough to be measured exactly. I should note
that they did not wither or lose any of their general appearance or freshness in these
experiments, which were all done at a temperature of 15◦ to 16◦ Réaumur.
The numbers in the table are reported with the volume of the leaf taken as unity.
Thus, when I represent by the number 5.5 the amount of oxygen gas consumed by oak
leaves, I mean that, in 24 hours, the leaves [p. 98] caused 5-1/2 times their volume of
oxygen gas to disappear, partly by the formation of free carbon dioxide gas, partly
by inspiration.
In none of these experiments did the inspirations appreciably exceed the volume
of the leaves, and most often they were less.
3.5 Relative Amounts of Oxygen Gas Consumed in Darkness by Different Leaves 39

RelativeAmounts of Oxygen Gas Consumed by Different Leaves (Tables 3.1–3.5)

[p. 99]

Table 3.1 Leaves of evergreen trees and shrubs

Names of leaves Time of Oxygen gas consumed during
experiment 24 hours in darkness; reported with
the volume of the leaf taken as unity
Holly (Ilex aquifolium) September 0.86
Box (Buxus sempervirens) September 1.46
Cherry-laurel (Prunus laurocerasus) May, new leaves 3.2
September 1.36
Laurustinus (Viburnum tinus) September 2.23
Ivy (Hedera helix) September 1
Periwinkle (Vinca minor) June 1.5
September 0.93
Spruce (Pinus abies) September 3
Hare’s ear (Bupleurum fruticosum) May 4
Savin juniper (Juniperus sabina) June 2.6
Common juniper (Juniperus communis) June 2.4

Table 3.2 Leaves of deciduous trees and shrubs [p. 100]

Names of leaves Time of Oxygen gas consumed during 24 hours
experiment in darkness; reported with the
volume of the leaf taken as unity
Beech (Fagus sylvatica) August 8
Hornbeam (Carpinus betulus) May 5
September 6
Oak (Quercus robur) May 5.5
September 5.5
Horse chestnut (Aesculus September 4.8
hypocastanum)
Poplar (Populus alba) May 6.2
September 4.36
Apricot (Prunus armeniaca) September 8
Peach (Amydalus persica) June 6.6
September 4.2
Walnut (Juglans regia) May 6.6
September 4.4
Plane tree (Platanus occidentalis) September 3
Acacia (Robinia pseudo acacia) May 5
September 6.7
Lilac (Syringa vulgaris) May 3.36
September 2.2
Ash (Fraxinus excelsior) May 4.32
September 3.71
Pear (Pyrus) May 5.2
September 3.4
Rose (Rosa centifolia) June 5.4
Chestnut (Fagus castanea) July 5.6
40 3 Influence of Oxygen Gas on Developed Plants

Table 3.3 Leaves of nonaquatic, herbaceous plants [p. 101]

Names of leaves Time of experiment Oxygen gas consumed during
24 hours in darkness; reported with
the volume of the leaf taken as unity
Potato (Solanum tuberosum) September, before 2.5
flowering
Cabbage (Brassica oleracea) September, young 2.4
leaves
September, old leaves 2
Nettle (Urtica urens) September 2
Mercury (Mercurialis annua) September, during 2.33
flowering
Carrot (Daucus carotta) September, during 1.9
flowering
Fava bean (Vicia faba) Before flowering 3.7
During flowering 2
After flowering 1.6
Lily (Lilium candidum) May, before flowering 0.66
September, after 0.5
flowering
Nasturtium (Tropaeolum majus) September, during 3
flowering
Foxglove (Digitalis ambigua) July 2
Turnip (Brassica rapa) September, during 1.25
flowering
Oats (Avena sativa) June, before 2.7
flowering
Wheat (Triticum aestivum) May, before flowering 5
Pea (Pisum sativum) May, during 3.72
flowering
Rue (Ruta graveolens) August 2

Table 3.4 Leaves of marsh or aquatic plants [p. 102]

Names of leaves Time of Oxygen gas consumed during
experiment 24 hours in darkness; reported with
the volume of the leaf taken as unity
Water plantain (Alisma plantago) August 0.7
Fleabane (Inula dissenterica) September 1.6
Epilobe (Epilobium molle) September, during 1.9
flowering
Watercress (Sisymbrium nasturtium) September 1.6
Polygonum persicaria September, during 2
flowering
Veronica (Veronica beccabunga) September 1.7
Creeping buttercup (Ranunculus September 1.5
reptans)
Loosestrife (Lythrum salicaria) May, before 2.3
flowering
Caltha palustris May 1
Carex acuta May 2.25
3.6 Influence of Atmospheric Oxygen Gas on Plant Roots 41

Table 3.5 Leaves of succulent plants [p. 103]

Names of leaves Time of Oxygen gas consumed during 24 hours
experiment in darkness; reported with the
volume of the leaf taken as unity
Raquette (Cactus opuntia) August 1
Agave americana August 0.8
Sempervivum tectorum July 1
Sedum globosum September 1.5
Saxyfraga colyledon September 0.6
Sedum reflexum June 1.7
Stapelia variegata July 0.63
Mesembryanthemum deltoides July 1.7

3.6 Influence of Atmospheric Oxygen Gas on

Plant Roots [p. 104]

Plant roots that are buried in humus are not entirely deprived of atmospheric oxygen
gas. A certain amount of the gas can be withdrawn simply by boiling water that has
filtered through plant mold. If we pour water into a hollow made in the soil surface
and collect the air that the water displaces in penetrating the earth, we find that it
is less pure than atmospheric air but that it still contains a considerable amount of
oxygen gas. Hales already said it, but it seemed interesting to me to see whether
contact of this gas with roots is useful to plant growth.
I uprooted some young, leafy, horse chestnut trees, each weighing about 23 grams.
They had long roots of 2.5 decimeters, and stems of about the same length. I intro-
duced the roots of chestnut FG (Fig. III [see Plate, p. lix]), into the neck E of a
1.2-liter receptacle, and I luted its stem precisely with the neck E. [p. 105] After
I had filled this vessel with distilled water, I introduced, through its opening CD,
594 cubic centimeters of nitrogen gas, which was in contact with the entire upper
part of the roots, whose tips were submerged in water HI. The receptacle rested on a
bowl filled with mercury. I set up a similar apparatus with hydrogen gas and another
with carbon dioxide gas, which I supplemented every day because it was absorbed
by the water HI. Three other horse chestnuts were set up separately, in the same way,
with atmospheric air.
The plant whose roots were in contact with carbon dioxide gas died first, after 7 or
8 days. The plants whose roots were in contact with nitrogen gas and hydrogen gas
died nearly simultaneously, after 13–14 days. By adding carbon dioxide gas during
the entire time they were growing, they had increased the volume of their atmosphere,
which, after subtracting the carbon dioxide, had not decreased. The horse chestnuts
whose roots were growing in ordinary air were still vigorous after 3 weeks, when I
terminated the experiment. Their roots had reduced the volume [p. 106] of gaseous
fluid in the receptacle, either by absorbing the carbon dioxide that they had formed,
or because of the absorption of this same gas by the water HI. The nitrogen gas of
this atmosphere did not decrease.
One may conclude from this that contact of oxygen gas with the roots is useful to
vegetation. Several other, less direct, observations support this contention:
42 3 Influence of Oxygen Gas on Developed Plants

1. Duhamel noted (Physique des arbres, Part 1, Chap. 5) that the “lateral roots are
stronger and more vigorous the closer they are to the soil surface: so that if in
planting a tree in a uniform soil, several layers [de Saussure transcribed this last
term as “plants” (“plants” in English) but Duhamel’s original says “plans,” one
of the meanings of which is “planes,” which I have interpreted as “layers”] of
roots are retained, the layer that is closest to the soil surface will almost always
be more vigorous than one which is deeper.” Is this not because the deeper level
has less contact with atmospheric oxygen gas than the more superficial one? One
might believe that it is because the upper roots find more nutritive juices at the soil
surface, but the following observation shows that the effect cannot be attributed
to this cause alone: If common soil is piled on the roots and around a tree that
is thriving in a good soil, [p. 107] this tree will be harmed. If it does not die,
the lower roots, although planted in a better soil, will decay, and the tree will
sprout new roots upon the old ones, in the newly added soil. It is not necessary to
mention the benefits that accrue from opening or raising the surface of soil used
by vegetation.
2. Tap-rooted plants—so-named from their structure, as they have stout vertical roots
almost devoid of root hairs—fare better, other things being equal, in a dry soil
than in a damp one, and also better in a light soil than in a compact one, probably
because these roots, which have little surface relative to their mass, require more
contact with oxygen gas than do fine roots, which may have this contact even in
a heavy soil.
3. Tree roots that penetrate manure, mud, or water pipes divide indefinitely and
form foxtails because they can grow only to the extent that they multiply their
points of contact with the very small amount of oxygen gas that they find in these
environments.
4. A plant whose roots are [p. 108] suddenly covered by stagnant water is harmed
much more quickly than if it had met this fate in running water. The oxygen
gas of standing water will soon be used up, but that of running water will not
be exhausted. It should be noted that standing waters are more useful to plants
than pure water if the plants receive the impure waters by sprinklings that are
managed so as to provide contact with oxygen gas. From these observations, one
may conclude that most plants will be harmed if the soil is too wet, not only
because water as a nutrient is provided to them in too large a quantity, but also
because their roots have almost no contact with the external air.
Some writers have believed that infusions of humus and manure do not themselves
serve as plant nutriment, because plants that they uprooted and immersed by the
roots in these liquors did not fare as well as they did in pure water. But I doubt very
much that this observation is true as a generality. When infusions are not rich, or
approach spring water in richness, plant growth is better and the plants gain [p. 109]
more weight than in distilled water or in filtered rainwater. I repeated the very precise
experiments of Woodward on this subject, and my results are consistent with his.
Observations to the contrary were made with little regard for accuracy and with richer
infusions than can be obtained by means of a press from a naturally moist, fertile
3.7 Of the Changes that Oxygen Gas Undergoes Through its Contact with Roots 43

plant mold. But observations should be made on more dilute infusions, because the
interstices of the soil allow air to reach the roots, whereas in our total immersions, the
oxygen gas that could have reached the roots through the medium of distilled water
and of a less saturated infusion is entirely consumed by the extractive substances in
a more concentrated infusion.

3.7 Of the Changes that Oxygen Gas Undergoes Through its

Contact with Roots

If a healthy root, lacking its stem and having [p. 110] enough growth force that
the root will not decompose by putrefaction, is placed under a receptacle filled
with atmospheric air closed by mercury, the root decreases the volume of this air,
apparently by absorbing oxygen gas. Moreover, it forms carbon dioxide gas with
the surrounding oxygen gas. But whatever the capacity of the receptacle and the
duration of the experiment, the amount of oxygen gas that seems to be absorbed
by the root is always less than the volume of the root. If a root thus saturated is
transferred immediately into another receptacle filled with ordinary air, it does not
change the volume of this air, but simply forms carbon dioxide with the oxygen gas.
But if, before the transfer, the root is exposed for some time to the open atmosphere,
it absorbs the same amount of oxygen gas under the receptacle as it did in the first
experiment.
A recently uprooted yellow carrot root consumed its own volume of oxygen gas
in 24 hours, and absorbed a hundredth of this amount.
A potato tuber consumed, during the same period, 0.4 of its volume of oxygen
gas and appeared to absorb 0.08 of its volume of this gas. A lily bulb having true
roots consumed [p. 111] 0.39 of its volume of oxygen gas and absorbed 0.19 of its
volume.
A turnip [“rave”], in the same amount of time, always consumed its own volume
of oxygen gas. Absorption was a quarter of this amount.
The inability of a root to absorb more than its own volume of oxygen gas under a
receptacle from which it is not removed, and the ability it acquires to absorb additional
oxygen gas by exposure to the open air, shows that this organ does not permanently
assimilate the oxygen gas that it absorbs, but only converts it into carbon dioxide
gas, which atmospheric air, due to its affinity for carbon dioxide gas, can remove
from the root.
Roots behave, both in sunlight and in shade, very nearly as leaves growing in
darkness do, but the inspirations of roots are much less noticeable because, since
they do not decompose carbon dioxide gas during the day, they are never entirely
depleted of it.
In the experiments I have reported up to now, the roots were lacking stems, but
the results are very different when the roots have their stems and the experiment is
set up so that the root [p. 112] is confined in a receptacle filled with atmospheric
air, while the stem and leaves project into the open air. The roots then appear to
44 3 Influence of Oxygen Gas on Developed Plants

absorb several times their own volume of oxygen gas. I used the following apparatus
for this purpose with Polygonum amphibium or P. persicaria. It is uncommon, I
believe, to find plants that, through the length and suppleness of their stems and their
growth force, lend themselves to this arrangement, for lutes must be avoided as far as
possible. Lutes will not hold reliably enough on plants whose stems are always more
or less subject to contraction and expansion. On the shelf of a mercury bath CD [he
is evidently referring here to Fig. IV of the Plate, p. lix], I placed a narrow receptacle
AB filled with atmospheric air. Onto the surface of the mercury HI contained in the
receptacle, I passed a layer of water FG, 7 to 8 lines thick. Finally, I introduced the
root EK of the Polygonum LM across the mercury into the receptacle. The layer of
water FG, which nourished the plant, was soon drawn up by it. I then added water
to the receptacle for several days in a row. From the ascent of the mercury in the
bell jar, I saw that the air in this vessel gradually decreased. [p. 113] When all the
added water had been absorbed and the air of the receptacle had decreased by nine
or ten times the root volume, I ended the experiment, and I found that nitrogen gas
had not been absorbed, that the oxygen gas alone had disappeared, in proportion to
the decrease in the air volume.
The oxygen gas was not assimilated by the plant but was returned to the atmo-
sphere by the leaves, for I saw that, when I placed a Polygonum having its leaves and
roots under a receptacle filled with ordinary air, such that the roots were immersed
only by their tips in the water contained under the receptacle and were mostly in
contact with the atmospheric air that was confined there, the oxygen gas no longer
decreased, because the leaves restored on the one hand that which the roots had ab-
sorbed on the other. In these experiments, the plant displaced only about a 300th of its
atmosphere.
When one introduces, into a globe that is carefully closed and full of atmospheric
air, the tip of a leafy branch occupying around a 400th or 500th part of the volume
of the globe and is rooted in plant mold (Fig. VIII), the branch improves [p. 114] its
atmosphere very noticeably after a fortnight or 3 weeks. But this change does not
occur perceptibly when the same plant is completely enclosed, with its roots, in the
globe.
To study accurately the influence that plants exert, with the help of pure water, on
their surrounding atmosphere, it is necessary, as I indicated in Fig. I, that the roots
not be immersed in the water BD that is used to close the receptacle, for, if this water
(Fig. II) is considerable in amount, and if it is renewed often, the results obtained
are analogous to those given by a branch (Fig. VIII) of a plant rooted in humus. The
atmosphere provides to the water, and the water to the roots, an unlimited amount of
oxygen gas, which is partly released by the leaves into the receptacle.
It follows in general from these observations that oxygen gas is not assimilated
directly by the roots, but forms, with their substance, carbon dioxide gas that is
absorbed by the roots and elaborated by the leaves.
3.8 Of the Changes that Oxygen Gas Undergoes Through Woody Stems 45

3.8 Of the Changes that Oxygen Gas Undergoes Through

Woody Stems [p. 115]

In the springtime, I placed some woody branches, just before bud opening, under a
receptacle filled with ordinary air (see apparatus, Fig. V [of the Plate, p. lix]), with a
tip of each immersed in a small amount of water to nourish them. The branches leafed
out as in open air. But this development could not take place under a receptacle filled
with nitrogen gas or hydrogen gas. They rotted in these gases, emitting nitrogen
and carbon dioxide gases, without giving any sign of growth. Defoliated woody
branches, arranged as I have just described, corrupt ordinary air, both in the sun and
in the shade, without changing (other than by the volume of the stem) the volume
of this atmosphere. They always replace the oxygen gas that they cause to disappear
by an equal volume of carbon dioxide gas. Thus they do not assimilate oxygen gas.
Their green parts undoubtedly assimilate a small amount in the sun, in decomposing
[p. 116] the carbon dioxide gas that they form themselves with the surrounding
oxygen gas, but this effect is not detectable by these experiments.
Branches of willow (Salix alba), oak (Quercus robur), poplar (Populus nigra),
and hornbeam (Carpinus betulus), 7 millimeters (1–3 lines) in thickness, consumed,
in the spring and summer, at 15◦ Réaumur, an amount of oxygen gas equal to at least
half, and at most all, of their volume in 24 hours. Pear and apple, under the same
conditions, consumed two or three times their volume.
Defoliated woody stems show the phenomenon of inspiration of oxygen gas in
darkness, and, undoubtedly due to the green substance contained in their bark, that of
expiration of this gas in the sunlight. This emission is not detectable in atmospheric
air by eudiometric tests because the stems form, with the surrounding oxygen gas,
more carbon dioxide gas in sunlight than in shade, and because their green parts do
not have enough surface area to decompose all the carbon dioxide that they form in
the sun. It is very clear, though, that the atmosphere of the branch [p. 117] decreases
during the night and resumes its original volume in the light.
Woody stems do not replace the oxygen gas that they consume by an equal volume
of carbon dioxide gas when the plant has its roots in soil, its leaves in the open air,
and its stem alone confined in an atmosphere of ordinary air, as shown in the apparati
in Figs. VI and VII, which are sealed by luting and mercury. The carbon dioxide
gas that is left by the branch in its atmosphere is less, by an indefinite amount, than
the oxygen gas that the branch consumed. The carbon dioxide that is missing here
follows the course c a d of the branch, to be decomposed by its leaves in the open
air. I will elaborate on the details of one of these experiments.
At the beginning of July, at 7 o’clock in the morning, I luted (Fig. VI) to the
opening a of the tube ab, of which the opening e was immersed in mercury, the
tip of an apple branch from which I had removed the leaves and the last, purely
herbaceous, ramifications. This branch was 5 mm (2 lines) in diameter at its section
c [author mistakenly wrote e rather than c here], and it occupied 2 cubic centimeters
(0.1 cubic inch) in the apparatus. The tube itself contained 125 cubic centimeters (6.3
cubic inches) of atmospheric air [p. 118] in part ab, for I made the mercury climb to
46 3 Influence of Oxygen Gas on Developed Plants

b in the tube so that the confined air would be at the same pressure as the external
air.
Two hours after sunset, the mercury had risen, after corrections were made, by
3-1/2 millimeters (1-1/2 lines) inside the tube. This rise corresponded to about half
the volume of the branch. The next day, at daybreak, the mercury had dropped a
little, and 2 hours after sunrise, the air had regained the volume that it had had before
the experiment. This air contained 3/100 carbon dioxide gas, and after deduction
of that, 13/100 oxygen gas. The branch thus consumed, in 24 hours, 5-1/3 times its
volume of oxygen gas.8 The branch replaced it by barely three times its volume of
free carbon dioxide gas, and produced [p. 119] a large amount of nitrogen gas, which
prevented any volume change due to absorption of oxygen or carbon dioxide. I will
return soon to this development of nitrogen gas, which is a necessary consequence of
the experimental set-up. When I prolonged the experiment for 48 and 96 hours, the
effects were doubled and tripled. There is therefore no doubt that the confined part
of the stem absorbs unlimited quantities of the carbon dioxide gas that it produces.
But in none of these trials, however prolonged, did the mercury climb more than
1-1/2 lines, and it always descended the next morning to the point where it had been
the previous evening. I repeated these experiments, with the same results, on myrtle
and Lonicera xylostheum, with the apparatus arranged as shown in Fig. VII.
I did the same experiments (Fig. VI) by closing with water [rather than mercury]
a tube containing a column of atmospheric air 1.4 decimeters (5 inches) in height.
After 3 or 4 days, the water had climbed 2.7 centimeters [author erroneously wrote
“decimeters”] (1 inch) inside the tube. It did not climb higher when I prolonged
the experiment. The air in the tube, examined at this time, contained only nitrogen
gas (with the exception of about 2 or 3 hundredths [p. 120] carbon dioxide). Thus
the air had undergone the same decrease in volume as it would have if subjected to
the effects of a sulfide [which would have absorbed oxygen]. No nitrogen gas was
produced. Hales performed this last experiment9 and obtained the same results, but
they prove nothing regarding our subject because the carbon dioxide gas that was
produced, since it is absorbed by the water, cannot be compared with the oxygen gas
that disappeared at the same time.
This famous natural philosopher had an opinion that must be contested: He be-
lieved that woody branches absorbed a large amount of air during the day and expired
it during the night.10 He cemented, as in the experiment in Fig. VI, the defoliated tip
of an apple branch to the opening a of tube ae. But he filled this tube entirely with
water and immersed its lower opening in mercury. The branch, by rapidly sucking in

8
A branch of the same dimensions as the previous one was cut or separated from this apple tree
and placed for 24 hours under a receptacle closed by mercury, in the same exposure as the branch
that was connected with the ground. It caused four times its volume of oxygen gas to disappear and
replaced it with four times its volume of carbon dioxide gas. It produced no nitrogen gas. It follows
from these two experiments that a branch connected with the ground consumes more oxygen gas
than one that is separated from it.
9
Statique des végétaux, French edition, p. 130.
10
Ibid., pp. 76 and 77.
3.8 Of the Changes that Oxygen Gas Undergoes Through Woody Stems 47

this water, caused the mercury to climb several inches above [de Saussure mistakenly
wrote “below”] its level. This ascent soon stopped because air bubbles were disen-
gaged from the enclosed branch, [p. 121] preventing a further rise. This air came
(and the author, moreover, saw it correctly) from atmospheric air introduced from
d into c through the suction or vacuum that tended to make the mercury rise in the
tube, which was imperfectly sealed by the porous wood of the branch. He noted that
the mercury rose during the day and descended at night, and it was on this occasion
that he expressed his opinion on the diurnal inspiration and nocturnal expiration of
stems. But he wrongly attributed to vegetation an effect that was dependent only on
the arrangement of his apparatus and on the greater and lesser water uptake during
the day and at night.
During the day, when the tree transpired through its leaves, the volume of water
drawn up by the branch exceeded the volume of atmospheric air that was introduced
from d to c in the tube, and this air exited as it had entered, that is, through the
pores of the wood. At night, when transpiration was less or nonexistent, so was the
suction, and the air, penetrating the apparatus in greater proportion than the water
was aspirated, caused the mercury to descend.
About the same thing happens when the tube is full of atmospheric air, but the
effects are almost imperceptible [p. 122] to the eye because the branch draws in
carbon dioxide gas much more slowly than it does water. No sooner has the branch,
by drawing in some of this gas, made the mercury climb by 1-1/2 lines above its
level, than the mercury draws into the apparatus atmospheric air, from which the
branch consumes the oxygen gas and leaves only the nitrogen gas. We must attribute
to this cause the production of nitrogen gas in the first experiment.
No nitrogen gas is produced in a column of air 1.4 decimeters (5 inches) in
height, closed by water, because the rise of 2.7 centimeters [author erroneously
wrote “decimeters”] (1 inch) of water is insufficient to draw atmospheric air into the
apparatus. But when the column of water is higher, it produces the same effect as
mercury.
I luted the tip of an apple branch to a tube ae (Fig. VI) completely full of mercury,
and I dipped this tube deep enough in a mercury bath that the luted neck a was
submerged in it. The branch emitted no air into the tube in 24 hours, either in the
sun or in the shade. But when, by lowering the level of the external bath, I made the
branch support a column of mercury 2.7 decimeters [p. 123] (10 inches) in height, the
branch then produced, especially through its section c, countless air bubbles, which
lowered the mercury about 1 decimeter (3 or 4 inches) in 24 hours. After 60 hours,
the mercury had fallen to the level of the mercury in the outer vessel. The air that
was produced contained 86/100 nitrogen gas, 9/100 oxygen gas, and 5/100 carbon
dioxide gas, but in calculating the amount of oxygen gas that the branch consumed
in the apparatus, it seemed to me that the air produced must have originally been
about as pure as atmospheric air.
For 8 days, I exposed to the sun a young apple tree bearing its leaves, stem, and
roots, under a receptacle full of atmospheric air. It did not change its atmosphere
noticeably. The oxygen gas that the stems appeared to assimilate in some of the
preceding experiments thus circulated [as carbon dioxide] within the plant, to be
exhaled by the leaves.
48 3 Influence of Oxygen Gas on Developed Plants

I removed the bark from parts ac of the branches contained in the apparati in
Figs. VI and VII. These branches also absorbed and decomposed through their leaves
the carbon dioxide that they produced. When I performed this experiment under a
receptacle, and on branches [p. 124] cut as in Fig. V, they left in this vessel an amount
of carbon dioxide gas precisely equal to the amount of oxygen gas that they removed,
and they produced, at the same volume and surface, the same effects one to two times
faster than the nondebarked wood. The hardness acquired by debarked trees thus is
not attributable, as several authors have believed, to the absorption of oxygen gas by
the wood. It is due instead, as I will show below, to a superabundance of carbon.
In doing these experiments, I observed wood, especially that of oak, that at first
was perfectly white turn dark yellow or brown within a few hours. This coloration
was, as others have observed, more intense in the sun than in the shade. The conver-
sion of oxygen gas into carbon dioxide gas by debarked wood was also faster in the
light than in darkness.
When I placed fresh, recently debarked oak wood in nitrogen gas or under water,
in the sunlight, it retained its whiteness fully, and then colored immediately in the
sunlight under receptacles full of ordinary air. This coloration is thus an effect of
contact with oxygen gas. It is based, [p. 125] as Bethollet thought (Art de la teinture,
Vol. 1, p. 48), on a precipitation of carbon and formation of water. But this water
formation does not arise from a combination of atmospheric oxygen gas with the
hydrogen of the wood. This explanation would contradict experiment, which shows
that the oxygen gas consumed is found in an exactly equal amount in the carbon
dioxide gas produced. The precipitation of carbon is due to the fact that the wood,
in losing a small amount of its carbon through its removal by oxygen gas, loses,
at the same time, in much greater proportion, its oxygen and hydrogen in the form
of water (see Chap. 4, Sect. 4.3). The loss of oxygen and hydrogen leaves carbon
predominant in the woody residue.
Senebier made an observation similar to the one I just reported. He saw that the
green color of leaves, when prepared by alcohol [“spirit of wine”], turns yellow in
the sun only when in contact with oxygen gas. (Physiologie végétale, Vol. 3, p. 144.)

3.9 Of the Changes that Oxygen Gas Undergoes Through its

Contact with Flowers

According to my experiments, flowers, even those of aquatic plants, do not [p. 126]
develop in pure nitrogen gas. Their buds, ready to open, are as if paralyzed there,
and, whether opened or not, putrefy more rapidly than under receptacles filled with
ordinary air. Flowers such as white lilies or double roses that were able to displace,
without deteriorating, a seventh or an eighth of their surrounding atmosphere during
24 hours under a receptacle filled with ordinary air, replaced, except for the volume
of the flower, the oxygen gas that they removed, by carbon dioxide gas. I say, except
for the volume of the flower, because the flower retained in its parenchyma an amount
of carbon dioxide gas smaller than its volume. But when the flower was transferred
3.9 Of the Changes that Oxygen Gas Undergoes Through its Contact with Flowers 49

immediately afterwards into another receptacle filled with ordinary air, it did not
make this absorption. I will not return to the explanation for this observation.
It is rare for recently picked flowers confined in a vessel full of ordinary air to
change the volume of this atmosphere noticeably, because they very nearly replace the
carbon dioxide gas that they absorb [presumably, oxygen gas that the flowers convert
to carbon dioxide and retain] by nitrogen gas. They differ in this respect from other
plant parts, which, when growing in darkness, exhale much less nitrogen gas [p. 127]
and consequently almost always reduce the volume of the surrounding air.
Might not the production of nitrogen gas in darkness by flowers that do not appear
to be injured be related to the production of their odors? Even if we found that very
fragrant flowers produced this gas as abundantly as less fragrant ones, we could
conclude nothing against this supposition, because it is possible that not all flower
scents affect our sense of smell. It is regrettable that flowers live so briefly that we can
hardly prolong experiments beyond 36 hours, and that we must fear that the slightest
disruption of such a delicate structure will influence the results.
In the experiments that I am going to report, the flowers seemed not to have
deteriorated in the slightest. They lost nothing of their appearance or freshness.
Those in the bud stage bloomed under the receptacle. When the green parts could not
be removed, they were left on, but these parts were too small relative to the volume
of the petals to have any noticeable effect in the experiments on rose, lily, wallflower,
nasturtium, [p. 128] and chestnut catkins. The white lilies consumed [i.e., took in]
their own volume of oxygen gas in 24 hours in the shade in ordinary air, under a
receptacle closed by mercury, at a temperature of +17◦ Réaumur. They absorbed
[presumably, they converted to carbon dioxide and retained] an amount of oxygen
equivalent to 15/100 of their volume, and they replaced it by 15/100 of their volume
of nitrogen gas. [The rest of the oxygen that the flowers consumed presumably was
converted to carbon dioxide and released as such.]
Based on this account, we can understand the following data:

Oxygen gas Absorption Nitrogen gas

consumed in the produced
shade in 24 hours
Flowers of lily (Lilium album) 1.1 0.15 0.15
Umbels of carrot (Daucus carotta) 2.1 0.3 0.3
Flowers of nasturtium (Tropaeolum majus) 2.17 0.5 0.5
Raceme of false dittany (Dictamnus albus) 2.6 0.7 0.7
Raceme of gillyflower, double 1.2 0.7 0.4
(Cheyranthus incanus)
Cabbage rose (Rosa centifolia) 1.8 0.43 0.43
Male catkins of chestnut (Fagus castanea) 4.7 0.22 0.22

[p. 129] Petals consume more oxygen gas in the sun than in the shade. They give
off no hydrogen gas. I tried in vain to spark an explosion in the pure oxygen gas in
which, for 8 days, I had grown false dittony racemes, which I renewed [replaced]
often. The inflammation that can be produced in this plant appears to arise strictly
from the combustion of its essential oil.
50 3 Influence of Oxygen Gas on Developed Plants

I did the same experiments with the same results, either in the sun or in the shade,
on nasturtiums and marigolds. It has been said that these flowers spontaneously
produce flashes, but this phenomenon has been seen by only two observers, even
though these flowers are widespread in gardens. It is all the more doubtful because
their brightness could easily create an illusion.

3.10 Influence of Oxygen Gas on Fruits

The experiments that can be done with fruits, with regard to the questions that have
occupied me up to now, are far less reliable than are those with other plant parts,
because fruits retain almost no growth force [p. 130] when they are separated from
the plant that produced them.
I introduced some unripe grapes, whose vine was rooted in soil, into a globe that
was carefully closed and exposed to the sun. Before ripening, they improved their
atmosphere of ordinary air, in a fortnight, without emitting a detectable amount of
carbon dioxide, and they ripened. But when I put into the same globe some quicklime,
slaked and dried quickly by the heat of boiling water, they corrupted their atmosphere
under the same conditions and were unable to ripen. I obtained the same results with
unripe berries of Solanum pseudocapsicum. Green fruits thus seem to yield the same
results as leaves.
When I placed unripe grapes, Solanum berries, pears, and apples separated from
the tree under receptacles full of ordinary air, these fruits corrupted their atmosphere
in 24 hours, although they spent the day in the sun. Their inspirations and expirations
were hardly noticeable. During the day they could not decompose all the carbon
dioxide gas that they formed during the night. When they [p. 131] remained in
continuous darkness for several weeks, they assimilated no oxygen gas. The oxygen
that they removed reappeared in the same amount (except for the volume of the fruit)
in the carbon dioxide produced.

3.11 Utility of Oxygen Gas in the Nutrition of Plants

To assemble here the principal effects of oxygen gas on plant nutrition, I will an-
ticipate some functions that will be discussed in Chaps. 7 and 8. Humus contains
extractive juices that penetrate plants and promote their growth. When these juices
are exhausted, oxygen gas, by removing carbon from the humus, develops a new
extract that replaces the first one. The effect of oxygen gas on the soil is not limited
to this one influence. When the humus loses its carbon, it yields at the same time, in
the form of water, its oxygen and hydrogen. Absorption of this liquid may be useful
to plant growth under certain conditions.
Plants that develop with the aid of [p. 132] distilled water, under a receptacle
full of atmospheric air, do not have humus for nutriment, but their own substance,
3.12 Summary 51

at whose expense they grow, replaces the humus. The inner parts of the stem, or a
part of the roots or lower leaves, wither and empty their extractive juices into the
developing parts. All the influences of oxygen gas on humus relate to the case that
we will now examine.
I have shown in Chap. 2 and in this one that green plants combine their carbon
with the surrounding oxygen gas to form carbon dioxide gas, and that they wither if
they are prevented from decomposing carbon dioxide, by the removal of the gas with
lime or potash. The extractive juices thus probably assimilate their carbon into the
green parts only insofar as it is converted into carbon dioxide gas. For this compound
to be formed, oxygen gas is an absolute necessity to plant growth.
Oxygen gas itself serves as a nutrient to the green parts of plants. They do not
assimilate it directly, but once they have converted it into carbon dioxide, they retain,
in decomposing the carbon dioxide, a part of its oxygen. This effect is so small that
it [p. 133] can be appreciated only when they are given a large amount of carbon
dioxide gas to decompose. (See Chap. 2, Sect. 2.4, and Chap. 7, Sect. 7.3)
An atmosphere composed only of nitrogen and carbon dioxide gases is not favor-
able to plant growth (Chap. 2, Sect. 2.2). Free oxygen gas must be present. Oxygen
gas thus has an effect beyond that of presenting to plants, in the form of carbon
dioxide gas, elements that they can assimilate. One can presume that this second
influence consists not only in developing a nutritive extract and water in the humus
or plant, but also in producing a release of caloric [a precursor idea to the concept of
energy; see Glossary] by the union of oxygen gas with the carbon of the plant. The
release of caloric is a necessary result of this combination. If it usually escapes our
notice, it is because of its small amount and the opposing effect of evaporation.

3.12 Summary

Green plants that are exposed in atmospheric air to the successive effects of day and
night make alternating inspirations and expirations [p. 134] of oxygen gas mixed
with carbon dioxide gas. The oxygen gas that green plants inspire is not directly
assimilated by them. It is metamorphosed during inspiration into carbon dioxide
gas. The plants decompose the carbon dioxide gas in the act of expiration, and it is
only through this decomposition, which is only partial, that they are able to assimilate
the oxygen gas that they find in the atmosphere.
The roots, wood, sapwood, petals, and all parts that in general are not green do not
make successive inspirations and expirations. They do not assimilate atmospheric
oxygen gas either directly or indirectly. They transform it into carbon dioxide gas,
which is retained or dissolved in small amounts in their succulent parts, as it would
be in pure water. Otherwise, they do not change it.
I showed how deceptive it can be to perform these experiments on a single plant
part. The carbon dioxide gas that the roots, trunk, and woody branches form with the
surrounding oxygen gas follows the longitudinal course of the plant to be decomposed
by the leaves.
52 3 Influence of Oxygen Gas on Developed Plants

[p. 135] I have shown that contact of oxygen gas with the roots and the nongreen
parts in general is essential to plant growth. I will return to this subject in Chap. 6.
The leaves of marsh plants, succulents, and evergreen trees generally consume,
other things being equal, less oxygen gas than do the leaves of other plants.
The most noticeable influence that oxygen gas exerts on plants is to form carbon
dioxide gas and to present to plants, in this form, elements that they can assimilate.
[p. 136] Chapter 4
Influence of Oxygen Gas on Some Plant-Derived
Substances [“proximate constituents of plants”]

Several plant compounds that come in contact with oxygen gas during the early
stages of fermentation show changes whose origin we are going to trace. This inves-
tigation should clarify for us the changes that these compounds undergo during plant
growth, should it be true that we can sometimes compare the effects of a disorganized
substance to the effects occurring in a plant enjoying all its growth capabilities.

4.1 Use of Oxygen Gas in the Precipitation of Extracts

Fourcroy1 noted that, through exposure to air, extracts dissolved in water allow
[p. 137] pellicles or insoluble material to precipitate in the liquid containing them.
He attributed this precipitation to the combination of oxygen gas with the extract
itself, but I do not know if the assertion of this learned author is based on an analysis
of this precipitate or on eudiometric tests.
I placed some capsules containing extracts of cinchona, oak wood, and various
humuses under jars filled with atmospheric air and closed by mercury. After a few
days I found that a precipitate had formed in the solutions and that the air in the
receptacles had undergone a small reduction in volume. Eudiometric examination
of this atmosphere showed me that the oxygen gas that had disappeared had not
completely reappeared in the free carbon dioxide gas that was produced at the
same time. But an important fact struck me, namely, that the volume reduction
of the oxygen gas in which the experiment was done was always less than the
volume of the extractive solution, whatever its degree of liquidity or thickness. If
10 volume-parts of liquid extract are placed in contact with 1,000 volume-parts of
atmospheric air, the air undergoes a decrease in volume that does not exceed 10
parts [p. 138]. This invariable result made me suspect that the oxygen gas does not
combine with the extract, but that the oxygen removes carbon from the extract by
forming carbon dioxide gas, of which the liquid can absorb only its own volume.

1
Annales de Chimie, Vol. 5.

J. F. Hill, Chemical Research on Plant Growth, 53

DOI 10.1007/978-1-4614-4136-6_4, © Springer Science+Business Media New York 2013
54 4 Influence of Oxygen Gas on Some Plant-Derived Substances . . .

When, after this experiment, I subjected these extractive juices, which were appar-
ently saturated with oxygen gas, to the effect of a vacuum, I withdrew from them an
amount of carbon dioxide gas nearly equal to the volume of oxygen gas that they had
absorbed, whereas this release did not occur before saturation. But this test was not
exact enough to be decisive. It is not so with the experiment I am now going to report.
I passed a stream of carbon dioxide gas through the extractive liquids before
I put them in the experiment. They were then exposed for a few moments to the
open air (to rid them of excess carbon dioxide). Immediately afterwards, they were
introduced under a receptacle full of ordinary air, which rested on mercury. If they
did not cause their atmosphere to undergo any expansion in several hours, the test
was continued, but in the case of expansion, the air of the receptacle [p. 139] was
renewed. The extracts thus prepared and well depleted of excess carbon dioxide
underwent the same changes as they would have if they had absorbed nothing. They
became covered with an insoluble pellicle but they did not change the volume of their
atmosphere, did not absorb oxygen gas, and did not remove it without replacing it
with a volume of carbon dioxide gas exactly equal to the oxygen gas consumed.
It thus follows from my experiments that extracts, in the first stages of fermenta-
tion and the formation of pellicles, assimilate no oxygen gas, and that the effect of
oxygen is limited to removal of carbon.
When the extracts lose a small amount of their carbon due to the effects of oxygen
gas, they give up a larger amount of their oxygen and hydrogen in the form of water.
I will give here but one example of the experiments I performed on this subject. They
all yielded similar results. I mixed 1.91 grams (36 grains) of dry extract of oak wood
with 30 grams (1 ounce) of water impregnated with carbon dioxide gas, at the precise
point at which this liquid caused no expansion of the atmosphere of ordinary air
[p. 140] in which I placed it. This atmosphere, which occupied 792 cubic centimeters
(40 cubic inches), contained 21/100 oxygen gas and negligible carbon dioxide. A
fortnight later, a thick pellicle had formed but the air of the receptacle had not
changed in volume. It then contained 5/100 carbon dioxide, and, after deduction of
that, 16/100 oxygen. It must be concluded from these results that 40 cubic centimeters
(2 cubic inches) of oxygen gas disappeared and were replaced by 2 cubic inches of
carbon dioxide gas, which contain, according to Lavoisier, 0.4 grains of carbon.
The extract from which this element [“principle”] had been removed weighed no
more than 30 grains after being dried in a water bath. Thus it seemed to have lost
5-1/2 grains of water from its own substance. But it was possible that this water was
lost during drying. Therefore I dissolved 36 grains of dry extract of oak wood in the
same amount of water as was used in the previous experiment. This solution, dried
immediately afterwards for the same length of time and at the same degree of fire as
the extract subjected to the effect of air, yielded 34 grains of dry extract. The drying
process [p. 141] therefore volatilized 2 grains of extract, or rather 2 grains of water
belonging to the dry substance of the extract. It follows from all these trials that
36 grains of extract were divested of 1/2 grain of carbon by the oxygen gas under the
receptacle, and that they lost or formed, simultaneously, an amount of water equal
to 3-1/2 grains. I would not report such modest results with confidence had I not
confirmed them by a very large number of other observations.
4.2 Use of Oxygen Gas in the Conversion of Wine to Vinegar 55

The loss of oxygen and hydrogen from the extracts, in the form of water, in much
larger proportion than the carbon that the oxygen gas removes from them, tends to
increase the proportion of carbon in the extractive residue. In fact, I found that 100
parts of dry extractive pellicles yield, through the residue of their distillation in an
open fire, more charcoal than 100 parts of extract that was not separated from its
pellicles. Thus, these insoluble flakes, far from being an oxygenated extract, are an
extract that is deoxygenated, or supersaturated with carbon.
The juices of green plants, very rich in the extractive material that Fourcroy called
plant albumin [p. 142] because it may show some characteristics of animal albumin,
do not absorb oxygen gas. The juices of Cactus opuntia and Sedum telephium, upon
contact with oxygen gas, coagulate into a gelatinous and elastic substance, without
changing the volume of this gas, only transforming it into carbon dioxide gas.
But care must be taken not to put the extractive juices in vessels in which they
have little access to the surrounding air. The solution must be spread over a large
area. Without this precaution, the parts that are too far removed from contact with
oxygen gas could form carbon dioxide gas entirely from their own substance.
When the temperature at the end of the experiment is very different from that at
the beginning, it is necessary, before ending the experiment, to return the extractive
liquor directly back to the temperature it had had at the moment it was introduced into
the receptacle, and not to make, by calculation, the correction that is usual in such a
case, because the ability of water to combine with carbon dioxide gas increases with
the lowering of temperature.
In none of the experiments that I have described [p. 143] have the plant substances
begun to putrefy or reached the stage of fermentation required for them to generate,
under certain conditions, hydrogen and nitrogen gases.

4.2 Use of Oxygen Gas in the Conversion of Wine to Vinegar

Through the ages, it has been recognized that contact with air markedly favors
the conversion of wine to vinegar [“acetification”]. Lavoisier2 found this acid
fully formed, without the intervention of oxygen gas, in the fermentation prod-
ucts of sugar with yeast, but as experiment shows that a very small amount of
oxygen gas is enough to give vinous liquor pronounced signs of acidity, I do not
know but what the oxygen gas that was dissolved or interposed in the distilled
water used in this experiment contributed substantially to the formation of the
acid. We also cannot know if acetic acid exists [p. 144] fully formed in yeast,
combined either with ammonia or with the glutinous substance contained in
the yeast. Berthollet has already noted that this experiment was not conclusive.3

2
Traité élémentaire de Chimie, by Lavoisier, pp. 143 and following.
3
Statique chimique, Vol. 2, p. 525.
56 4 Influence of Oxygen Gas on Some Plant-Derived Substances . . .

Rozier found that oxygen gas was absorbed during the conversion of wine to
vinegar. He concluded this from the decrease in volume undergone by atmospheric
air in this process. But I showed in the preceding section that this decrease is not
a proof of the assimilation of oxygen gas with plant matter, if the volume of gas
absorbed is not compared with the volume of the liquid. I showed that the oxygen
gas absorbed may be limited to forming carbon dioxide gas, which remains dissolved
or interposed in the liquor.
For a full year, I held wine in contact with oxygen gas in receptacles closed by
mercury. This wine turned to vinegar, but I never found that the reduction in volume
of the oxygen gas exceeded the volume of the wine. It was always less than that.
When I performed this experiment with wine artificially impregnated with carbon
dioxide gas, [p. 145] at the point where the mixture no longer expanded atmospheric
air,4 this wine also turned to vinegar, but without changing the volume of its atmo-
sphere and by replacing the oxygen gas (which disappeared) by exactly the same
volume of carbon dioxide gas.
Three cubic inches of Bordeaux wine, lightly impregnated with carbon dioxide gas
and placed for 5 months in contact with 80 cubic inches of atmospheric air confined
by mercury in an oven heated to a constant 20–23◦ Réaumur, became acidic to the
point of being undrinkable and of eliciting a slight effervescence in potash. The
wine underwent this change without changing the volume of its atmosphere, but by
forming with oxygen gas about 6 cubic inches of carbon dioxide gas and causing
an equal volume of oxygen gas to disappear. Four experiments done under the same
conditions on different wines gave me similar results. In a fifth experiment, [p. 146]
I found a small volume reduction and a corresponding absorption of oxygen gas,
but, in this particular case, the wine had not been placed in, and withdrawn from, the
experiment at the same temperature. It was at +20◦ Réaumur when the experiment
began and 0◦ when it ended. I do not doubt that the attraction of wine for carbon
dioxide gas, which increases with the lowering of the temperature, was the cause of
this anomaly.
A freshly prepared dough of wheat flour does not assimilate oxygen gas when
acidifying in atmospheric air. The effect of this gas is limited to removal of carbon.
This experiment, done in the summer, lasted around 60 hours.
Three cubic inches of alcohol placed for 5 months in contact with 80 cubic inches
of atmospheric air underwent no change, nor did the surrounding air.
My experiments on acetic acid formation do not undermine the generally accepted
view that wines become acidified due to a larger share of oxygen, for they are formed
largely of oxygen, hydrogen and carbon, and the removal of carbon increases the
proportion of oxygen.
[p. 147] One could suppose that wines, on the one hand, assimilate oxygen, and,
on the other hand, return an exactly equal volume of carbon dioxide gas entirely from
their own substance; but admitting such a consistently exact compensation would
perhaps be giving too much weight to this hypothesis. Besides, the result would be

4
Wine by itself, deprived of air, always expands atmospheric air a little, due to the alcohol that is
dissolved in it and that assumes a gaseous state.
4.3 Influence of Oxygen Gas on Dead Wood 57

the same under this supposition: Wines become oxygenated or acidified only through
loss of their carbon, since they would lose, in the elimination of carbon dioxide gas,
as much oxygen gas as they would take from the atmospheric air.

4.3 Influence of Oxygen Gas on Dead Wood

Wood, moistened and placed in oxygen-free environments, add to them carbon diox-
ide gas formed entirely from its own substance. The formation of this gas does not
have the same origin when the wood is in the presence of oxygen gas.
I mixed 15.29 grams (1/2 ounce) of sawdust of an oak trunk, dried at a set reading
of the thermometer and hygrometer, with water so as to make a paste. [p. 148]
The mixture, placed in a glass capsule for 5 weeks under a capacious receptacle
full of ordinary air and closed by mercury, was exposed to a constant temperature.
After the allotted time, this air had not changed in volume. There were formed 198
cubic centimeters (10 cubic inches) [de Saussure wrote 1.98 cubic centimeters, but
presumably 10 cubic inches is correct, so the metric equivalent should be 198 cubic
centimeters] of carbon dioxide gas, which took the place of the same volume of
oxygen gas. This sawdust, which was then exposed to the open air under conditions
otherwise the same as those under the receptacle, was dried after 3 weeks, at the
same readings of the thermometer and hygrometer as before it was moistened. Its
dry weight had decreased by 796 milligrams (15 grains). Yet during its confinement
and drying, it could have lost only 159 milligrams (3 grains) of carbon, at most.
It had thus lost, in addition to this element, an amount of oxygen and hydrogen
represented by at least 637 milligrams (12 grains) of water. The sawdust was covered,
on the surface exposed to air, by a dark brown crust. One hundred parts of this crust
yielded,5 by carbonization, 20-1/2 parts of charcoal. [p. 149] One hundred parts of
sawdust, before its alteration by air, produced only 17-1/2 parts of charcoal.
From these experiments and several others, it follows: (1) that wood and sapwood,
after their death, do not assimilate atmospheric oxygen gas; (2) that the effect of
atmospheric oxygen gas is limited to removing carbon from them; (3) that these
plant substances lose oxygen and hydrogen, in the form of water, at the same time
and in greater proportion than carbon; (4) that the removal of oxygen and hydrogen
increases the proportion of carbon in the wood residue.
Woods that decompose through the effects only of water, without contact with
oxygen gas, as occurs in dead branches that retain a thick, tight bark, undergo
very different changes. Their substance whitens rather than darkening, and their
proportion of carbon decreases rather than increasing. I believe that this change is
due largely to the loss of their extractive and coloring substances, which are richer
in carbon than is the wood itself.
One hundred parts of oak wood that had decomposed to a white color without
contact with air [p. 150] yielded 16 parts of charcoal, excluding the ash.

5
The procedures followed in this operation are indicated in the note on carbonizations at the end
of Chap. 5.
58 4 Influence of Oxygen Gas on Some Plant-Derived Substances . . .

One hundred parts of oak wood, rotted to a black color with contact with air,
yielded 25-1/2 parts of charcoal, excluding the ash.
It is impossible to reduce wood, by multiple decoctions, to the point where it loses
no more extractive substances to water. Each decoction or maceration in contact with
air produces an extract that had not existed before, or that becomes soluble in water
through a change of substances brought about in the process of boiling or maceration.
I boiled 92 grams (3 ounces) of oak wood sawdust for a half-hour in 24 times its weight
of distilled water. The decoction, filtered and evaporated at a gentle heat, yielded a
dry extract weighing 4.7 grams (90 grains). A second decoction, similar to the first,
gave an extract weighing 1.5 grams (29 grains). These amounts decreased continually
up to the 9th decoction, which yielded an extract weighing 212 milligrams (4 grains).
The 11th decoction yielded a similar amount, and so did the 12th. The moistened
sawdust that had been subjected to these twelve operations [p. 151] was exposed to
air, but protected from dust, for 2 months. I ascertained that the wood only converted
oxygen gas into carbon dioxide gas, without assimilating the oxygen, during this
period. After the allotted time, the sawdust yielded, by a 13th decoction similar to
the previous ones, an amount of extract that weighed 292 milligrams (5-1/2 grains),
and that was, consequently, larger than the product of the 9th decoction. A 14th
decoction yielded an extract weighing 4 grains, but after 2 months of exposure to air,
the amount of extract produced by a single operation returned again to 5-1/2 grains.
Wood that is the most exhausted of its extractive substances always yields, when
macerated in cold water in contact with air, infusions that are rich in extracts. These
extracts are all soluble in a small amount of water. In a concentrated state, they
are clouded by limewater, potassium carbonate [“carbonate of potash”], and metal
solutions.
Oxygen gas has less effect on wood lacking its extractive substances than on wood
that has them.
Wood freed of its extract by decoction in a large amount of water yields [p. 152],
at the same weight, less charcoal than does natural wood.
The sap, in forming sapwood [sap was believed at that time to give rise to sap-
wood], probably undergoes changes somewhat similar to those of an extract that is
carbonized by the effects of oxygen gas and that becomes partly insoluble in water.
But there is undoubtedly this difference: In the change in the extract, carbon that is
more or less impure precipitates or separates, in a black form, from the rest of the
compound, whereas in the sap, through a particular disposition of this substance or by
the effects of plant growth, the carbon remains in combination to form the sapwood.
In this compound, insoluble in water, the proportion of carbon is at a maximum.
But when the sapwood is exposed to the effects of air, the combination is destroyed.
The combination parts with its carbon, which combines in part with oxygen gas
and in larger part precipitates in the form of humus or very impure charcoal. The
wood, through the separation of this carbon, returns towards its origin by passing to
a state of water-soluble juice. Fully formed wood itself is often perhaps no longer a
homogeneous combination, but a very condensed sapwood mixed with charcoal or
humus. See the note on carbonizations at the end of Chap. 5.
4.5 Condensation of Oxygen Gas by Putrefying Plant Substances 59

4.4 Condensation of Oxygen Gas by Oils [p. 153]

Oils cause oxygen gas to disappear, but not in the way that extractive juices, vinous
liquors, and wood do, solely by forming carbon dioxide gas. I found that turpentine
oil could absorb, in 4 months, twenty times its volume of oxygen gas, producing a
volume of carbon dioxide gas that was four times less than the volume of oxygen gas
absorbed. The results were the same when the oil being tested had been saturated
with carbon dioxide beforehand.6
Linseed oil could absorb more than twelve times its volume of oxygen gas in
[p. 154] 4 months without leaving a detectable amount of carbon dioxide in its
atmosphere. Of all the substances, either animal or plant, that I tested, it is the
only one that gave me this result. This property could make it suitable for use in
eudiometric tests. The linseed oil that I used in my experiments did not condense
nitrogen gas and did not emit any gas at room temperature in gaseous environments
devoid of oxygen gas.
The occurrence of water in oils, both animal and plant, that have been exposed
for a long time to the effects of air has been established by a very large number of
observations. It can be presumed that the oxygen gas absorbed is not assimilated by
these substances. I have not noticed the formation of water in the experiments that I
have detailed, but perhaps they were not continued long enough and the oil was able
to hold in solution or suspension the water that it contributed to forming. We must
therefore await further observations before deciding whether the formation of resins
by oils, through the effects of air, is due to a removal of substances or to an addition
of foreign oxygen gas, or to both effects together. We will be content [p. 155] to note
that the removal of carbon contributes little or nothing to this formation of resins.
Oils do not appear to absorb or assimilate oxygen gas in the process of plant
growth. They seem to have this property only when they have become a secretion,
or a body external to the plant that produced them. Thus the seeds of flax and hemp,
from which oils that condense oxygen gas can be expressed, do not absorb this gas
noticeably in the process of germination. They modify it, as do mucilaginous and
starchy seeds, only by converting it into carbon dioxide gas.

4.5 Condensation of Oxygen Gas by Putrefying Plant

Substances

Plants, regardless of their stage of fermentation, never increase the volume of the
atmosphere with which they are in direct contact if it contains a large proportion of
oxygen gas. But if it contains little or no oxygen, they increase it, in the early stages

6
In doing these experiments, I had occasion to observe a fact reported by Priestley: that turpentine
oil absorbs nitrogen gas. But it seemed to me that the volume of nitrogen gas that was condensed
never exceeded the volume of the oil. Priestley recognized that the oil lost this property over time
and regained it after being exposed to a vacuum. It therefore seems very likely that the nitrogen
does not lose its caloric in this combination.
60 4 Influence of Oxygen Gas on Some Plant-Derived Substances . . .

of fermentation, [p. 156] by pure carbon dioxide gas, and, during putrefaction or
the final stage of fermentation, by hydrogen or other inflammable gases mixed with
carbon dioxide and nitrogen gas.
I have, on several occasions, observed a fact already reported by Berthollet,7 with
his customary accuracy, namely, that a putrefying body, all of whose parts are in
contact with atmospheric air, never emits either hydrogen or nitrogen gas.
It seemed to me that as long as fermenting plant substances are not emitting
hydrogen gas into an oxygen-free atmosphere, they will not change the volume of an
atmosphere of oxygen gas, but that if they are put in an atmosphere of oxygen gas at
a time when they can emit hydrogen gas under water or in nitrogen gas, they cause
a condensation of the oxygen gas. I deprived some pea seeds of their growth ability
by submerging them for several moments in boiling water. These peas, swollen
and submerged in water, at first did not emit any gas. [p. 157] Carbon dioxide
appeared first, followed by hydrogen. Immediately after their death, I placed them
in an atmosphere of ordinary air. After 4 days the atmosphere had not changed in
volume. Six cubic inches of oxygen gas had disappeared and been replaced by 6 cubic
inches of carbon dioxide gas. After the experiment, I submerged these same peas in
water. After 2 days they began to release inflammable gas. I then put them in direct
contact with 100 cubic inches of atmospheric air closed by mercury. After 3 days,
this atmosphere had been condensed by 1 cubic inch, and I found by eudiometric
examination that 7 cubic inches of oxygen gas had disappeared and been replaced by
only 6 cubic inches of carbon dioxide gas. This atmosphere contained no hydrogen
gas. Haricots, favas, and potato roots provided, during the period when they emitted
hydrogen gas, the same results as the peas.
I placed 3.82 grams (1 gros) of fresh gluten under a receptacle containing 1,070
cubic centimeters (54 cubic inches) of atmospheric air. After 8 days at a constant
temperature [p. 158] of 10–12◦ Réaumur, and the gluten appearing to be softened,
the air of the receptacle had not changed in volume. It had lost 59 cubic centimeters
(3 cubic inches) of oxygen gas, which had been replaced by the same volume of
carbon dioxide gas. The gluten then had the sour smell of wheat dough beginning
to ferment. By drying for 3 hours, it was reduced to 1.35 grams. But before the
experiment, it had been reduced by drying to 1.48 grams (28 grains). Thus it had lost
2 grains of water from its dry substance.
I continued the experiment, under the same conditions, on 3.82 grams (1 gros)
of the same gluten, brought to the stage of fermentation at which I had left that of
the previous experiment. When placed under water, it began to release hydrogen
gas. After 8 days, the 54 cubic inches of atmospheric air had decreased by 3/4 cubic
inch. The air then contained 8/100 carbon dioxide, and after the deduction of this,
11-1/2 hundredths oxygen. Nine and one-half parts of oxygen gas had disappeared
and been replaced by only eight parts of carbon dioxide gas. [p. 159] The gluten
floated in the water that it had formed. I dried it at a gentle heat. It was reduced by
this operation to 1.38 grams (26 grains). It had thus produced 46 grains of water. This
liquid contained ammonia, but an insignificant weight of it. It did not turn plant dyes
[used as reagent] green, probably because the ammonia was combined either with

7
Statique chimique, Vol. 2, p. 546.
4.6 Summary 61

the gluten itself or with acetic acid. During putrefaction, the gluten was covered by
a black dust that did not appear when it putrefied under water without contact with
air.

4.6 Summary

It follows from the observations reported in this chapter that, in general (with the
exception of the oils), during the earliest stages of fermentation, oxygen gas is not
fixed in dead plants nor does it unite with their hydrogen to form water, and that
it removes only carbon from them. Thus, in the conversion of wine to vinegar, in
the precipitation of extracts, and in the coagulation of plant albumin, oxygen gas is
used only to remove this carbon. The proportion of carbon, however, [p. 160] is not
decreased in the combined residues from this removal, because the plant substances
that are concerned release, at the same time, a part of their oxygen and hydrogen in
the form of water.
But oxygen gas combines either with the hydrogen of dead plants, or with their
whole substance, when they begin to putrefy and at the stage of fermentation at
which they can release hydrogen gas. Since the time of this combination is exactly
that of the development of hydrogen gas, since hydrogen gas is no longer found
in the oxygen gas with which the putrefying plant is in direct contact, and, finally,
since a much greater amount of water appears at this stage of fermentation, we must
presume that the oxygen gas that has disappeared and that is not found in the carbon
dioxide gas produced at the same time is not assimilated by the plant substance but
is used to form water.
There is an important difference to note between the products of fermentation
that occurs without contact with oxygen gas and the products of fermentation that
occurs with this contact. In the first case, the carbon dioxide that is disengaged draws
[p. 161] its two elements from the fermenting plant, whereas, in the second case,
the carbon dioxide seems to draw only one of its elements (the carbon) from the
fermenting substance.
[p. 162] Chapter 5
Of Plant Humus

5.1 Research on the Composition of Humus

I mean by the term humus the black substance with which dead plants become covered
when exposed to the combined effects of oxygen gas and water. The experiments
that I reported in the preceding chapter tend to show that this substance is not the
result of the combination of oxygen gas with the dead plant, but that it is the residue
of the removal of some of the elements of this plant.
For most of my research, I used almost pure humus that was freed, by a fine
sieve, of most of the undecomposed plants that are [p. 163] always mixed with it.
This humus contained little but the mineral components that arose from the plants
that produced it. I collected the humus from high rocks or from within tree trunks,
where it could not have been modified by the foreign substances that are ordinarily
introduced into the soil through livestock access, manures, and deposits by spring
water. This humus seemed fertile to me, especially when mixed with a certain amount
of sand or gravel, which serves as support for the roots and provides access to oxygen
gas. I excepted, however, the humus that forms in the trunks of certain trees, such as
oak. When the water has no outlet, the humus is rich with excess extractives, which
block the vessels of the plants. These soluble constituents do not arise in this case
entirely from the humus itself, but partly from the living tree, and thus are not suited
to the nutrition of all plants.
The following operations provide an overview of the differences that may gener-
ally be seen between the composition of a humus and that of the plants from which
it originates.
Distillation of Oak Wood [p. 164] 10.614 grams (200 grains) of dry oak wood,
distilled at a heat up to red hot, in a luted glass retort, yielded 2,293 cubic centimeters
(116 cubic inches) of methane, + 575 cubic centimeters (29 cubic inches) of carbon
dioxide, + 4.25 grams (80 grains) of water holding in solution pyrolignate of am-
monia with excess crude acetic acid [de Saussure’s term for crude acetic acid was
“pyroligneous acid”], + 589 milligrams (13 grains) of bituminous empyreumatic oil
[liquid oil obtained from the distillation of some organic substances]. The charcoal
remaining in the retort weighed 2.23 grams (42 grains). It contained 26 milligrams
(1/2 grain) of ash.

J. F. Hill, Chemical Research on Plant Growth, 63

DOI 10.1007/978-1-4614-4136-6_5, © Springer Science+Business Media New York 2013
64 5 Of Plant Humus

Distillation of Brown Humus of Oak Wood 10.614 grams (200 grains) of dry
humus of oak wood, distilled as the previous wood was, yielded 2,456 cubic cen-
timeters (124 cubic inches) of methane, + 673 cubic centimeters (34 cubic inches)
of carbon dioxide gas, + 2.81 grams (53 grains) of water holding in solution py-
rolignate of ammonia and carbonate of ammonia, + 530 milligrams (10 grains) of
bituminous empyreumatic oil. The charcoal remaining in the retort weighed 3.13
grams (59 grains). It contained 424 milligrams (8 grains) of ash.
Distillation of Whole Rhododendron ferrugineum Plants [p. 165] 10.614 grams
of dry plants, distilled at a heat up to red hot in a luted glass retort, yielded 1,982
cubic centimeters (100 cubic inches) of methane, + 634 cubic centimeters (32 cubic
inches) of carbon dioxide gas, + 3.34 grams (63 grains) of water rich in pyrolignate
of ammonia with excess crude acetic acid, + 1.7 grams (32 grains) of bituminous
empyreumatic oil. The charcoal remaining in the retort weighed 2.813 grams (53
grains). It contained 159 milligrams (3 grains) of ash.
Distillation of Black Humus of the Above Rhododendron 10.614 grams (200
grains) of dry humus, distilled at a heat up to red hot, yielded 2,040 cubic centimeters
(103 cubic inches) of methane, + 673 cubic centimeters (34 cubic inches) of carbon
dioxide gas, + 3 grams (57 grains) of water rich in pyrolignate of ammonia and car-
bonate of ammonia, + 557 milligrams (11 grains) of bituminous empyreumatic oil.
The charcoal residue of the distillation weighed 3.45 grams (65 grains). It contained
689 milligrams (13 grains) of ash.
Distillation of the humus of grass and spruce, [p. 166] and of these plants them-
selves, yielded products showing the same differences [between plant and humus]. It
follows that undecomposed plants contain, at the same weight, more oxygen and less
carbon than do their humuses. We do not know whether the carbon of these humuses
is completely combined with their other constituents.
Nitrogen occurs in greater proportion in humus than in the undecomposed plant.
This result is not surprising since plants that ferment in contact with air evolve almost
no nitrogen gas. We cannot, however, attribute to this cause alone all the carbonate of
ammonia that I obtained from the distillation of humus. It undoubtedly arises partly
from the insects that live in the humus and that leave their remains in it.
The famous Klaproth obtained, by the distillation of peat, products1 that undoubt-
edly contained very little nitrogen, since crude acetic acid was very abundant in it.
But peat cannot be considered a true humus. It is the residue of plant decomposition
[p. 167] occurring in stagnant water and partly without contact with air. In this case,
the plants lose nitrogen in a gaseous form. Peat appears to contain less carbon than
is found in a finished humus. In the table at the end of this chapter, compare the
carbonizations of humus nos. 8, 15, and 19, with the carbonizations of peat nos. 32
and following.
Acids show no noteworthy effect when mixed with humus. They produce no
effervescence; they do not completely dissolve the humus; and they take up some
of the iron and earthy substances that the humus contains, but very little of the

1
Beiträge zur Chemischen Kenntniss. Vol. 3.
5.2 Extractive Substances of Humus 65

plant component. Concentrated hydrochloric and sulfuric acids reduce the plant
component to charcoal with the aid of heat, and they disengage acetic acid from it,
according to the comment of Vauquelin.
Alcohol does not dissolve humus. It usually separates from the humus a small
amount of extractive substances and resin, equivalent at most to two or three
hundredths of the weight of the humus.
Potash and soda dissolve humus almost completely. The humus releases ammonia
through their effects. This solution is decomposed by acids. [p. 168] They precipitate
from it a combustible, brown powder, which is not abundant compared with the
weight of the humus devoted to the operation.

5.2 Extractive Substances of Humus

Humus is essentially insoluble in water. This liquid separates from it extractive

substances that are not the humus itself. Via the following experiments, I will give an
overview of the amount of extract taken up by pure water that falls on a fertile soil.
I filled a large vessel with nearly pure grass humus and watered it with distilled or
rain water until it could absorb no more. After 5 days, it was subjected to the action
of a press. Ten thousand parts, by weight, of expressed and filtered fluid yielded, on
evaporation to dryness, a dry extract weighing 26 parts.
I performed the same experiment, for the same length of time, on the heavy soil
of a vegetable garden that had been amended with manure. Ten thousand parts, by
weight, of expressed water yielded a dry extract weighing 10 parts.
[p. 169] The same experiment, repeated under identical conditions on the light
soil of a field that bore a fine harvest of wheat [“blé”], yielded, from 10,000 parts of
water, 4 parts of extract.
The humuses were dry before the experiment, and the water used to moisten
them contained no carbon dioxide. But that was no longer the case when the water
was withdrawn. It then clouded limewater with calcareous carbonate, although not
much more than ordinary spring water does. One hundred cubic inches of water
from the humus, expressed into a retort in which I boiled the water immediately after
expression, yielded an air containing at most 2 cubic inches of carbon dioxide gas.
This determination may not be very precise, but other observations indicate that the
amount of carbon dioxide that roots draw from common soil is not substantial.
When the entire upper part of a green plant rooted in humus is put into a globe,
and the neck of this vessel is carefully closed at the origin of the stem, one may see
a small improvement in the air of the flask only after several days or even several
weeks, [p. 170] although a very large volume of water is given off by the leaves.
The amount of extract that boiling water can separate from humuses that are pure,
natural,2 and formed in the countryside is not considerable. I subjected such humuses

2
I mean by these terms a humus that leaves only a small amount of ash after it is burned, or no more
than a tenth of its weight. I also assume that the humus was not amended with manure, or formed
by an artificial accumulation of plants that died at the same time.
66 5 Of Plant Humus

to twelve successive decoctions, each lasting a half-hour, with an amount of water

weighing more than twenty-four times the weight of the humus. The amount of
extract that I could collect by all these operations was no more than an eleventh of
the weight of the humus, and often much less. It seemed to me that a pure humus
that yielded, by the twelve decoctions I mentioned, an amount of extract equal to an
eleventh of its weight, was, under the same conditions, less fertile for favas and peas
than the same humus containing only a half or two-thirds of that amount.
[p. 171] If the amount of extract that a humus should contain in order to support a
good crop must not be too large, it also must not be too small. I subjected an almost
pure humus to twelve successive decoctions, with renewal of the water. In two garden
pots full of the humus, I sowed seeds of fava, pea, and barley, and sprinkled them
with rainwater, comparable in purity to distilled water. Other, similar seeds were
sown at the same time in two pots like the preceding ones, and filled with the same
humus, except that this humus had not been depleted of its extract. The plants in both
experiments bore fertile seeds, but the weight of these plants and their seeds was one-
quarter greater in the humus provided with its extract than in the exhausted humus.
The action of decoction, however, had not changed the external characteristics of the
exhausted humus. It could not be distinguished, by sight or feel, from the humus that
still contained its extract. It seemed to me only that the exhausted humus was able
to retain or imbibe more water.
One hundred parts of humus, dry and depleted of [p. 172] most of its soluble
matter, were able to hold 477 parts of water.
Dry, unwashed humus could hold at most 400 parts.
Humus, like wood, cannot be entirely freed of its extractive substances by water,
at least detectably and in contact with air. The first macerations or decoctions remove
more extract than do the following ones, but soon a point is reached at which it yields
an amount that is constant or no longer decreases appreciably. If the moistened humus,
after it has reached this maximal exhaustion, is exposed for a long time to the effects
of external air, it changes in such a way that it can produce more extract than it did by
the preceding decoction. Ten thousand parts of dry, black humus of Rhododendron
ferrugineum, which yielded by combustion 65 parts of ash, were boiled with twenty-
four times their weight of distilled water. This decoction, after filtering, yielded
an extract weighing 250 parts. The extract of the 9th decoction weighed 40 parts.
The 10th and 11th decoctions each separately provided a similar amount. Humus
exhausted to this degree, and moistened, was exposed [p. 173] to the effects of
air, but protected from dust, for 3 months. After this period, it was subjected to a
12th decoction similar to the preceding, and it yielded an extract weighing 58 parts.
Maceration in cold water produces similar effects. These macerations, if conducted
for a long time and repeated more than 50 times, on the same humus already exhausted
by decoction, always produced, at least in contact with air, infusions stained by a
highly soluble extract.
Humus that has been partially freed of its extractive substances by decoction
yields, at distillation, very nearly the same products as unexhausted humus, but
the amount of charcoal residue from this operation is a little greater in the partly
exhausted humus. One hundred parts of partly exhausted humus yielded 33-1/4 parts
5.3 Of the Salts Contained in Humus 67

of charcoal, containing 5-1/4 parts of ash. One hundred parts of the same humus,
provided with its extract, yielded, by the same operation, 31 parts [de Saussure wrote
“grains” but clearly meant “parts”] of charcoal, containing 6-1/2 parts of ash.3 This
increase in the proportion of carbon in the humus is confined within very narrow
limits. [p. 174] When I subjected this same humus, exhausted by several successive
decoctions, to new decoctions, I could not continue to increase the proportion of
carbon, although I removed a large amount of extract by these operations.
Humus extract is not deliquescent. It yields carbonate of ammonia at distillation.
The aqueous solution of this extract, which is close to syrup in consistency, is neither
alkaline nor acid. It has a sweet taste; precipitates in air; and is clouded after a
few moments by limewater, carbonate of potash, and most metallic solutions. When
mixed with alcohol, the alcohol dissolves a small part of it and separates another
that is insoluble. The substance soluble in alcohol is very deliquescent.4 [Footnote
4, which appears at the bottom of p. 174, is not cited in text; it seems to belong
here.] The extract produced by the first macerations of humus in water contains, at
equal weight, a greater proportion of the deliquescent substance than does the extract
produced by the subsequent macerations.

5.3 Of the Salts Contained in Humus [p. 175]

Reagents do not usually reveal, if simply mixed with the infusion of a natural hu-
mus that formed in the countryside, significant amounts of potash, chlorides [called
“muriates” by de Saussure], or alkaline sulfates, if the substratum on which the
humus rests has added none. Most of the alkaline salts contained in plants mani-
fest themselves only in the residue of their combustion. It is the same for the salts
contained in humus.
Some writers have believed that plants themselves create the salts that they contain,
because the ash of most naturally occurring humus does not lose salts to boiling water.
This conclusion is undoubtedly premature. All the humus that I examined contained
alkaline salts, although the ash of the humus was often resistant to water. But these
salts were retained in the ash by semi-vitrification with the earthy substances, if the
latter were very abundant. One hundred parts of grass humus yielded, [p. 176] by
combustion, 50 parts of sand or ash that did not lose saline substances to boiling
water. But 100 parts of dry extract of this same humus yielded 14 parts of ash, and
100 parts of this ash formed, with boiling water, a lye containing 25 parts of salts
composed of free potash, chlorides, and alkaline sulfates. Further analysis showed
me that the water had extracted only half of the salts contained in this same ash.

3
For the procedures used in these carbonizations, see the note at the end of this chapter.
4
I do not know whether this deliquescent extractive substance is enclosed by the nondeliquescent
substance, which is superabundant, such that the deliquescent substance is prevented from drawing
humidity from the air, or whether the alcohol brings about a new combination in the extractive
substances.
68 5 Of Plant Humus

One hundred parts of Rhododendron humus contained 6-1/2 parts of ash. One
hundred parts of this ash lost to water only 1/2 part of alkaline salts. But 100 parts
of the ash of the extract of this same humus released to water one-third of its weight
of alkaline salts, and this liquid was far from extracting the salts completely.
I did the same experiments on six other, very different, kinds of humus. They all
provided me similar results.

5.4 Of Changes that Oxygen Gas Undergoes by its Contact with

Humus [p. 177]

Humus is the residue of a putrefied substance but is itself no longer subject to putre-
faction. It may even be considered an antiseptic, for the extractive substances that it
contains are likely to proceed to putrid fermentation if they are isolated, but not if
they remain united with the humus. For 1 year, I kept pure, unexhausted humuses in
receptacles full of water and closed by mercury. They did not emit any gas, except
perhaps for the small amount of carbon dioxide with which the water covering them
became charged.
There is no doubt that humus, except for its saline and earthy constituents, is
entirely destructible through the combined effects of air and water. Without recourse
to detailed research to prove it, I can do no better than to transcribe here the observa-
tions of my father (Voyages dans les Alpes, § 1319) on the plant mold that covers the
plains between Turin and Milan and [p. 178] whose cultivation dates back to remote
antiquity.
The thin layer of topsoil seen in these plains seems to me also to show that the amount of
this soil cannot be regarded as a measure of the time that has elapsed since the land began to
produce crops, for in this space of 10 leagues between Turin and Saint-Germano, I nowhere
saw a layer as thick as a foot. Now in my opinion the shallowness of this layer shows that
this soil is subject to a decomposition that sets a limit on its increase, for otherwise, why
would a flat land, fertile and cultivated for more than 3,000 years, not have a thicker layer?
This destructibility of the topsoil is a fact beyond exception, and farmers who have wished
to replace manures by too-frequent plowing have had a sad experience. They have seen their
soil gradually impoverished and their fields become barren by the destruction of the topsoil.
Thus, since this soil is destructible, the amount destroyed must be, [p. 179] up to a certain
point, proportional to its absolute amount, and as, on the other hand, only a limited amount
is produced annually, its increase must necessarily have certain limits.
The limits of this increase must vary according to the climate; the nature and location of
the substrate that serves as a base for the topsoil; the plants that grow there; the kind of
cultivation they are given; and, finally, the fertility of the land. But even if all the forces that
tend to increase the thickness of this soil layer were conjoined, this layer undoubtedly would
finally reach a certain maximum beyond which the destructive forces, having become equal
to the constructive forces, would allow no further increase.

Pure humus, imbibed with distilled water and enclosed under receptacles filled with
atmospheric air or oxygen gas confined by mercury, formed carbon dioxide gas
there, in causing oxygen gas to disappear. But the humus was never able to decrease
the volume of this atmosphere by an amount greater than the volume of the water
5.4 Of Changes that Oxygen Gas Undergoes by its Contact with Humus 69

used to moisten it, whatever the amount of humus and [p. 180] the duration of
the experiment, which was sometimes prolonged to more than a year. When this
water was impregnated with carbon dioxide beforehand, the humus did not change
the volume of its atmosphere. The oxygen gas consumed reappeared, in an exactly
equal amount, in the carbon dioxide gas produced, and the humus released neither
hydrogen gas nor nitrogen gas.
It is evident from these experiments that humus neither fixes nor assimilates
atmospheric oxygen gas. The effect of the oxygen gas is limited strictly to removing
carbon from the humus.
For the results given above to be obtained, the humus must not be rich in iron
or clay deposits. Incompletely oxidized iron contained in these deposits combines
effectively with oxygen gas, but this effect is not produced by humus or by pure
earths, or even appreciably by iron and manganese that are combined with the plant
part of the humus and that are revealed there by incineration.
Oxygen gas removes more carbon from humus that retains its extractive substances
than from humus that is [p. 181] depleted of them by decoctions. The latter, at equal
weight, formed half as much carbon dioxide gas with atmospheric oxygen gas as did
the same, but unexhausted, humus.
Moistened humus, contained in vessels that do not allow its extractive substances
to escape, loses, if considered in the dry state, some of its weight through contact with
oxygen gas, and this loss is greater than the weight of the carbon that is removed
by this gas. In a glass capsule, I mixed 30.57 grams (1 ounce) of Rhododendron
humus, dried5 in the shade at a set reading of the thermometer and hygrometer, with
water until it could imbibe no more. I put this capsule under a large receptacle full of
atmospheric air. This air was renewed several times and examined eudiometrically
each time. The experiment lasted 4 months. The humus passed 3 of them under a
receptacle, and 1 drying in the shade in open air to the same degree as it had been
before being mixed with water. I then found that its weight [p. 182] had decreased
by 849 milligrams (16 grains). During its confinement under the receptacle, it had
removed 476 cubic centimeters (24 cubic inches) of oxygen gas and replaced them
by the same volume of carbon dioxide. Thus, during the entire experiment, it had
formed about 32 cubic inches of carbon dioxide gas, assuming that the production
of carbon dioxide was the same during confinement and during drying.6 Now, as 32
cubic inches of carbon dioxide contain, according to Lavoisier, 6 grains of carbon,
the humus must have lost, besides this element, an amount of oxygen and hydrogen
represented by 10 grains of water.
The proportion of carbon increases by the removal of water from the residue of
plants that are being reduced to humus, but I believe that carbon is not increased by

5
For a long time this humus was completely dry to the touch, but although it was not deliquescent,
its weight varied with the readings of the thermometer and hygrometer.
6
Humus probably forms less carbon dioxide, other things being equal, under a receptacle than in
open air. But as it has almost no effect on oxygen gas during the last stages of desiccation, which
takes a long time, I believe that the amount of carbon dioxide that I attribute to drying is in excess
rather than default.
70 5 Of Plant Humus

this process in a finished humus. A finished humus must lose, due to the effects of
[p. 183] air and water, its oxygen, hydrogen, and carbon in very nearly the same
proportion. If it lost its oxygen and hydrogen in greater proportion than its carbon,
we would often find, in a soil long denuded of vegetation, residues of nearly pure
carbon or of charcoal. But we do not find this. They all yield, by distillation, products
of which carbon makes up half, at most.
Carbon is a very powerful antiseptic, and the state in which it occurs in humus
seems suited, in part, to give it this property. I filled several equivalent vessels with
various kinds of humus, pure or almost pure. In each of them, I placed the same
amount of beef flesh. The same experiment was done, under conditions otherwise
the same, with sawdusts of the woods from which these humuses had arisen; with
pure charcoal; with carcareous sand; with siliceous sand; with clayey sand; and
in the open air. The flesh stayed fresh a little longer, without deteriorating, with
the charcoal than with the humus, but much longer with the humus than with the
sawdusts or with the various sands, or in the open air. It is partly [p. 184] because
of this antiseptic property of humus that, for maintaining plant growth, there is a
big difference between nourishing plants with extractive solutions, isolated and not
renewed, and supplying them with humus. The insoluble part of the humus prevents
the undecomposed plant substances from furnishing plants with fermenting juices
that are always harmful to growth.

5.5 Summary

Carbon occurs in greater proportion in humus than in the plants from which the
humus arises. The proportion of carbon that a humus contains, however, does not
seem able to be increased significantly by the continued action of the causes that
formed it.
Humus that is moistened, but considered in the dry state, loses weight at room
temperature through contact with oxygen gas. Oxygen is not fixed in the humus,
and does not unite with the hydrogen of the humus to form water. The oxygen gas
removes only carbon from the humus. The humus, in losing this element, loses at the
same time, in the form of water, its oxygen and its hydrogen, and an extract soluble
in this liquid. [p. 185] Humus thus appears to be completely destructible at room
temperature, by the joint action of oxygen gas and washings.
The extractive juices of humus contribute to a certain extent to its fertility: Their
ash contains all the substances of plant ash.
Pure humus is antiseptic.

5.6 Note on the Carbonization of Various Plant Substances

When a plant or a substance derived from it [a “proximate constituent”] is distilled in

a luted glass retort, the degree of fire that the retort can withstand without melting is
5.6 Note on the Carbonization of Various Plant Substances 71

often insufficient to expel all the hydrogen that remains combined with the charcoal
and that could be disengaged at a higher heat. This procedure would not, however, be
inexact for estimating the relative amounts of charcoal contained in non-volatile plant
substances, if the various charcoals all retained the same proportion of hydrogen at
the highest degree of fire that the retort can withstand. [p. 186] But it is not so. Plant
substances such as certain seeds, that are dense and capable of softening through the
effects of fire, retain a larger proportion of hydrogen than do those that are less dense
and do not soften, as most woods.
In the results that I am going to give, the degree of fire used for carbonization was
higher than that which melts silver, and, as the weight of the various charcoals did
not change when they were subjected to a higher temperature, I have reason to think
that they are comparable among themselves. I was consistent in the way I managed
the fire and in the time taken to bring it to the highest level.
To carry out these carbonizations, I wrap the dry plant material in paper and form
it into a compact ball that is introduced into a cylindrical iron box, or a gun barrel that
is closed on one side and open on the other. It is 9 centimeters high and 4 centimeters
wide. I lay a small iron disk, with a vertical handle, on the ball. This handle is used to
remove the disk. The disk fits the inside diameter of the cylinder precisely. This plate
is luted into this position with clay. [p. 187] It is covered with a layer of charcoal
dust, and a second layer, of ash.
After the operation, I weigh the charred ball without defoming it, while it is
hot, and I subtract the weight of the charred paper, which is known from an earlier
operation.
The precautions taken in collecting, drying, and incinerating the plants are the
same as those indicated in Chap. 9.
In some cases, the relative amounts of charcoal produced by different substances
can be judged only by supposing them free of ash. It is this result that is entered in the
fifth column of the table of carbonizations. I will give an example: 100 parts by weight
of humus of spruce leaves (no. 19) produced 28 parts of ash; 100 parts of the same hu-
mus produced 52-1/2 parts of charcoal provided with ash. By subtracting the ash from
the humus and from the charcoal, and setting up the proportion 100 − 28: 52.5 − 28 =
100: x, I find that 100 parts of this humus devoid of ash would have yielded
34 parts of charcoal [i.e., 100 − 28 = 72; 52.5 − 28 = 24.5; 72: 24.5 = 100: x;
therefore x = 34] (Table 5.1).
 72

Table 5.1 Table of Carbonizations [pp. 188–192]

Carbo- Name of substance Weight of charcoal Weight of ash contained in Weight of charcoal sup-
nization supplied by 100 parts of 100 parts of dry substance plied by 100 parts of dry
No. dry substance substance, excluding ash
1 Wood of oak (Quercus robur) separated from the sapwood 19.75 0.2 19.69
2 Sapwood of the preceding wood 17.5 0.4 17.16
3 Wood and sapwood of the preceding oak, freed of their 16.75 0.2 16.54
extract by repeated decoctions
4 Bark of the trunk of the preceding oak 26 5 22.1
5 Phloem of the preceding bark 24.8 6.2 19.82
6 Bark of oak branches 1 centimeter (5–6 lines) in diameter 26.6 6 21.92
7 Debarked oak branches 1 centimeter (5–6 lines) in 17 0.4 16.66
diameter
8 Brown humus of oak wood 28.5 4 25.44
9 Oak wood that had become white due to decay without air 20 6.8 14.1
contact
10 Oak leaves collected in May 30 5.3 26.1
11 Oak leaves collected in Sept. 26 5.5 21.69
12 Levant gall-nuts 30 2 28.57
13 Cork 22 0.87 21.31
14 Entire Rhododendron ferrugineum plants 23.5 1.6 22.5
15 Black humus formed by the preceding plant 31 6.5 26.2
16 Same humus stripped of its extract by repeated decoctions 33.25 5.25 28.58
17 Debarked wood of spruce (Pinus abies) 20 0.34 (according to Kirwan) 19.72
18 Leaves of the preceding spruce 24.5 3 22.16
19 Humus formed by leaves of the preceding spruce 52.5 28 34
20 Wood of mulberry (Morus nigra) separated from the 23.25 0.7 22.7
sapwood
21 Sapwood of the preceding wood 17.25 1.3 16
22 Bark of the preceding tree 25 8.9 17.68
23 Phloem of the preceding bark 18.1 8.8 10.19
5 Of Plant Humus
Table 5.1 (continued)
24 Debarked branches of hazel (Corylus avellana); they were 16.5 0.5 16.08
1 centimeter (5 lines) in diameter
25 Barks of the preceding branches 25.6 6.2 20.68
26 Leaves of the preceding hazel, collected in May 29.25 5.8 24.9
27 The same, collected in June 29.5 6.2 24.84
28 The same, collected in Sept. 28 7 22.58
29 Wood of hornbeam (Carpinus betulus) 17.75 0.6 17.25
30 Sapwood of the preceding wood 17.5 0.7 16.92. In this tree, the
sapwood was very
indistinct from the wood
31 Bark of the preceding tree 30.5 13.5 19.74
32 Peat analyzed by Klaproth 38.75 18.5 23.62. This carbonization,
done in a glass retort,
could not be completed
33 Holland peat 34.25 21.3 16.45
34 Wheat plants collected May 1, a month before flowering 25 7.9 18.56
35 The same, in flower, June 14 25 5.4 20.71
36 The same, bearing mature seeds, July 22 26.6 3.3 19.95
5.6 Note on the Carbonization of Various Plant Substances

37 Straw of the preceding plants, separated from the seeds 23.6 4.3 20.13
38 Seeds of the preceding plants 20.5 1.3 19.45
39 Wheat flour 19.5 0.82 18.83
40 Bran 23.25 5.2 19.04
41 Starch 10.75 0.16 10.6
42 Gluten 22.75 1.25 21.77
43 Gum arabic 17.75 2.5 15.64
44 Gum tragacanth 16.75 2 15.05
45 Crystallized sugar 18.5 0.1 18.41
46 Unsized white paper 11 0.88 10.2
47 Holland linen washed several times 14 3 11.34
73
74 5 Of Plant Humus

[p. 193] Green plant parts appear to differ from the other parts in having a larger
proportion of carbon.
The proportion of carbon in the green parts decreases in autumn. They then deplete
themselves of their glutinous and extractive juices. These substances are very rich in
carbon, since, as we see, washed wood, linen, and paper contain only a very small
proportion of this element.
The wood contains more carbon than does the sapwood.
Bark usually contains more carbon than does the wood or sapwood. This result
is not consistent among all trees because the bark is not a homogeneous substance.
Its epidermis alone is carbonized by contact with air. The phloem and the inner parts
of the cork are not often changed by this cause, and the proportion of their carbon
varies in different plants, according to circumstances that it is impossible for us to
appreciate.
[p. 194] Chapter 6
Of Plant Growth in Oxygen-Free Environments

In focusing, in the two preceding chapters, on the decomposition that plants may
undergo after their death, I interrupted the train of my research on plant growth, but
this digression was necessary so that effects of a dead plant or its fermentation could
be distinguished from those of plant growth.
The development of some plants in oxygen-free environments presents phenom-
ena arising from a combination of these two effects.

6.1 Of Plants that Cannot Maintain their Growth

in Nitrogen Gas

Plant growth seems able to maintain itself, with the aid of water, in pure nitrogen gas
only [p. 195] through the oxygen gas that the green parts of the plants exhale there.
Plants lacking green parts or having them only in small amount cannot grow in
this atmosphere. Thus, seeds do not germinate in it, and if apparent exceptions to
this rule are found, it is because of the use in these experiments of too much water,
which, because it cannot be entirely depleted of the oxygen gas that is dissolved
or interposed in it, provides enough gas for development to begin. Not only does
germination fail to occur in nitrogen gas, but seeds that have already germinated
and are placed in it always die if they have sprouted only their radicles before being
put into this gas. I performed these experiments on seeds of pea, garden and water
cresses, and Polygonum amphibium. They all rotted without developing. But most
of the plants that developed from these seeds were able to survive and elongate
indefinitely in this gas, if they were placed in it after they had abundant green parts
or leaves.
Woody branches [p. 196] of poplar (Populus nigra) and willow (Salix alba), with
leaf buds ready to open, could not bring about this development with the aid of
water, in nitrogen gas, either in the sun or in the shade. They began to putrefy after a
fortnight. These same branches leafed out after 3 or 4 days when they were placed,
in conditions otherwise the same, under receptacles filled with ordinary air, and they
maintained their growth there for several weeks.

J. F. Hill, Chemical Research on Plant Growth, 75

DOI 10.1007/978-1-4614-4136-6_6, © Springer Science+Business Media New York 2013
76 6 Of Plant Growth in Oxygen-Free Environments

A wilted plant placed in a weakly lit spot, in a receptacle filled with ordinary air
or oxygen gas confined by water, always becomes covered with mold, but it does not
acquire mold in nitrogen gas. When I placed these molds, fully formed, in nitrogen
gas, they did not develop further. But the gas must be completely pure, for the least
amount of oxygen gas suffices to maintain the growth of these very small plants.
Roses, lilies, and carnations that were picked 2 or 3 hours before full bloom and
that did in fact bloom at the end of this period under receptacles filled with ordinary
air, could not accomplish this, with the aid of water, in nitrogen gas. They rotted
there, at the [p. 197] same developmental stage as when they had been cut, and more
rapidly than in ordinary air. These effects also occurred in a vacuum.
A rose has been said to last longer in a vacuum than in ordinary air, but appearances
are deceiving. It is true that the rose drops its petals sooner in ordinary air, but this
loss, which is a natural process in plant growth, does not indicate any decomposition
in the plant. The fallen petals give off a weak but agreeable odor. The opposite occurs
in a vacuum or in nitrogen gas: A rose appears to preserve its form and color there
for a long time, but when, after a fortnight, we expect to withdraw it still fresh, it
gives off a stench. Its petals are rotten, and it is clear that this apparent life conceals
a veritable death.

6.2 Of Plants that can Grow in Nitrogen Gas

As I have said, only plants that are amply provided with green parts can grow in
nitrogen gas, [p. 198] and even these are not all equally successful. It seemed to
me that they had to present a lot of surface area, and that they had to belong to the
class of plants that consume the least oxygen gas when growing in atmospheric air
in darkness.
Cactus opuntia, nourished by water, was able to maintain its growth in the sun for
3 weeks in nitrogen gas, but it was greatly harmed. In the shade it died in 5 to 6 days.
It was about the same with Sedum telephium. These plants grow indefinitely under
receptacles full of ordinary air.
Pea plants that were able to withstand the first 4 or 5 days in an atmosphere of
nitrogen gas (which is not always the case) continued to grow there, in the sun, for
months, but only sluggishly.
I will report an experiment made on these plants. The result can be considered an
average taken from several observations.
Three partially developed pea plants, which together weighed about 3 grams,
gained, under a receptacle full of ordinary air, in the sun, [p. 199] with nourishment
from pure water, 1.27 grams (24 grains) in 10 days. Similar plants that withstood
the effects of nitrogen gas, in the sun, for the same length of time gained no more
than 106 milligrams (3 grains). [There is a conversion error here; 106 milligrams =
2 grains.] These plants, when placed in nitrogen gas in the shade, always died within
the first 4 days. They survived for several weeks in ordinary air.
6.2 Of Plants that can Grow in Nitrogen Gas 77

Smaller periwinkles survived, both in the sun and in the shade, for as long in
nitrogen gas as in ordinary air, namely, about 3 weeks. In both cases, they died only
because they could not tolerate an excessively humid atmosphere.
Lythrum salicaria, Inula dysenterica, Epilobium hirsutum, E. molle, E. mon-
tanum, and Polygonum persicaria, which are all more or less marsh plants, grew
admirably in nitrogen gas and made good development there for an indefinite time,
similar to that under receptacles full of ordinary air. They could even sustain their
growth [p. 200] in nitrogen gas for months, if exposed to a weak light or protected
from the direct effect of sunlight. I turn now to the changes these plants make in this
atmosphere.
I grew Lythrum salicaria in the sun in 65 cubic inches of nitrogen gas, which
showed no decrease by nitric oxide [a test for oxygen; see “nitrous air test,” in
Glossary]. This plant displaced about 1/8 cubic inch, and it was immersed, only by
its roots, in an ounce of water. This liquid had no contact with the water that closed the
receptacle. In the course of the experiment, I had to renew [replace] the plant five or
six times because, in elongating, it pressed and burned against the walls of the vessel
that covered it. After 2 months, its atmosphere had increased by 3.4 cubic inches.
The eudiometer then indicated 5/100 oxygen gas there. I prolonged the experiment
for another month in this improved atmosphere but the plant did not continue to add
oxygen gas to it. Polygonum and other plants gave me similar results. I ascertained,
by several experiments, that in general the amount of oxygen gas that is developed
in nitrogen gas is not proportional to the length of stay of the plant under the [p. 201]
receptacle, but that the amount of gas usually produced in the first weeks does not
increase further in the following weeks, although growth is equally vigorous at all
times.
Similar plants that I grew for the same time under receptacles filled with ordinary
air never added oxygen gas there.
When I placed these plants in nitrogen gas in complete darkness,1 renewing them
every 12 hours so that their growth would not languish, they produced no oxygen gas.
They augmented their atmosphere with carbon dioxide gas that they formed entirely
from their own substance.
If the same experiment was performed in ordinary air, carbon dioxide gas was
also produced, [p. 202] but the volume of the atmosphere was neither increased nor
decreased by it. This gas thus had another origin. It was formed by the carbon of the
plant and the surrounding oxygen gas.
These observations show us the source of the oxygen gas that plants eliminate
in nitrogen gas. The oxygen comes from the decomposition of the carbon dioxide
gas that the plant forms entirely from its own substance. When the plant has thereby

1
Lythrum salicaria, Polygonum persicaria, and other marsh plants exposed to a weak or diffuse
light do not leave carbon dioxide in their atmosphere of nitrogen gas. They add oxygen gas to it.
But for them to produce this effect and maintain their growth for a long time at this exposure, the
temperature must not be excessively elevated, for plants, like animals, require and consume less
oxygen gas, the lower the temperature.
78 6 Of Plant Growth in Oxygen-Free Environments

made itself an atmosphere having sufficient oxygen gas, it releases no more of it,
because the carbon dioxide gas that it then produces and decomposes is the result
of the union of its carbon with oxygen gas that is already formed. Then it makes
almost all the oxygen that it has caused to disappear during the night reappear during
the day.
The small amount of oxygen gas that Lythrum and Polygonum formed in nitrogen
gas was necessary for the development of these plants, but it was much more than
they needed to maintain their growth without developing.
I suspended, at the upper part of a receptacle containing 60 cubic inches of nitrogen
gas, a mixture of 1 part iron filings, 2 parts flowers of sulfur [sublimed sulfur], and
1-1/2 parts water.2 [This mixture would remove oxygen gas] [p. 203] At the same time,
I introduced into this atmosphere confined by water two Lythrum salicaria plants,
which together displaced 1/8 cubic inch. Their roots alone were immersed under the
receptacle, in a small vessel containing 2 centiliters of water. The apparatus was ex-
posed, in a room, to the effects of sunlight moderated by window glass. Ten days later,
one of the plants was dead. The other continued to grow or maintain itself without
suffering harm, without the wilting of any leaf, for 4 months, from the third of ther-
midor [11th month (July–Aug.) of the French republican calendar] to the second of
frimaire [3rd month (Nov.–Dec.) of that calendar], at which time I withdrew the plant,
as healthy as before its introduction. The air of the receptacle underwent no reduction
by nitric oxide. The plant had made no development whatsoever during its confine-
ment. Its growth remained, somehow, suspended. This is the only effect produced by
the sulfide in this experiment. The same plant elongated by 5 or 6 inches after 10 days
[p. 204] in nitrogen gas in which this substance was not present. The removal of the
oxygen gas developed by the Lythrum presented the only obstacle to its development.
The sulfide vapors had no role in this effect, for similar plants elongated consider-
ably with iron sulfide under a receptacle filled with ordinary air that I renewed every
3 days.
For 5 weeks, a Polygonum persicaria plant responded very nearly as the Lythrum
had, under a receptacle filled with nitrogen gas in which I had suspended concentrated
potassium sulfide [“hydrosulfide of potash”]. The plant did not develop at all and lost
two leaves from the vicinity of its roots. It died after the allotted time, only from the ef-
fects of too-strong sunlight, from which I had not taken the precaution of protecting it.
Plants that I grew in nitrogen gas in the sun died much more quickly than did
plants in ordinary air, from the effects of quicklime or potash suspended in their
vicinity.
It is strange to see marsh plants withstand the effect of a sulfide, which removes
oxygen gas from them, and not withstand the effect of lime, which removes [p. 205]
carbon dioxide gas from them. But it should be noted that the sulfide removes oxygen
gas only after the oxygen is formed, whereas lime or potash removes this same gas

2
Each part here is equivalent to 11.5 grams (3 gros.) The correct proportion of water is essential
for the mixture to exert a strong effect on oxygen gas.
6.2 Of Plants that can Grow in Nitrogen Gas 79

from them before its release. [Lime and potash remove carbon dioxide before the
plant has a chance to break it down and release the oxygen contained in it.]
A superabundance of carbon dioxide gas is much more harmful to plants growing
in nitrogen gas than to ones growing in ordinary air. I have said elsewhere that carbon
dioxide gas mixed in the proportion of one-twelfth with atmospheric air, in which I
grew pea plants in the sun, favored their development. It did not harm marsh plants
in the same circumstances. But marsh plants were never able to tolerate this mixture,
in the proportions indicated, with pure nitrogen gas. They died there in a few days,
as did the pea plants. The elaboration of a certain amount of oxygen gas thus appears
necessary for the elaboration of a certain amount of carbon dioxide gas. Carbon
dioxide always becomes harmful to plants when they cannot decompose it.
Priestley believed that he had found that some plants have the ability to absorb
the nitrogen gas in which they are growing. He reported that an Epilobium hirsutum
plant, [p. 206] placed in a receptacle 10 inches high and 1 inch wide, absorbed, after
a month, seven-eighths of the atmospheric air that it contained.3
Ingen-Housz did not limit this faculty to a small number of plants. He observed4
that all the plants grown in nitrogen gas caused an appreciable decrease in the amount
of this gas in a few hours. I followed, with great care, the growth of Epilobium
hirsutum, both in ordinary air and in pure nitrogen gas, using the procedures indicated
by Priestley for this experiment,5 and prolonging it greatly, but I did not see any
decrease in the nitrogen gas after the removal of the oxygen gas that was formed
there. It was the same with all the other plants that I tested [p. 207]. Thus plants do
not condense nitrogen gas appreciably. The experiments of Senebier and Woodhouse
confirm this assertion.
If nitrogen is a simple substance, if it is not an element of water, we are forced
to recognize that plants assimilate it only from plant and animal extracts and from
ammonia vapors,6 or from other water-soluble compounds that they can absorb from
the soil and atmosphere. We must acknowledge that when they grow in an unrenewed
atmosphere, with the aid of a small amount of pure water, the parts that develop obtain
nitrogen only at the expense of the nitrogen that the other parts of the plant contained
before the experiment.

3
Experiments and Observations on Different Kinds of Airs, Vol. 3, p. 332.
4
Expériences sur les Végétaux, Vol. 2, p. 146.
5
The procedure consists of planting the plant in a pot full of plant mold, submerging this pot and
the origin of the stem in water underneath the shelf of a basin, and covering the rest of the plant
with a receptacle full of air. The plant then undergoes much greater development than it does when
it has its roots in pure water. For this reason, I was obliged to replace the plant several times.
6
The presence of ammonia vapors in the atmosphere cannot be doubted, when we observe that pure
aluminum sulfate [“sulfate of alumina”] is eventually converted, in the open air, into ammonium
aluminum sulfate [“ammoniacal sulfate of alumina”]. The superiority of animal manures to plant
manures seems, in large part, due only to a greater proportion of nitrogen in the former.
80 6 Of Plant Growth in Oxygen-Free Environments

6.3 Of Plant Growth in Carbon Monoxide [“oxide of carbon”]

Gas7 (Berthollet’s oxycarburetted hydrogen) [p.208]

I prepared this gas by distilling, in a gun barrel, at a red heat, a mixture of equal parts
calcite [“calcareous spar”] and iron filings. The gaseous fluid that was obtained after
having been freed of carbon dioxide contained 1/100 oxygen, which I removed by
potassium sulfide [“hydrosulfide of potash”].
Plants grew in the carbon monoxide gas as in nitrogen gas. Those lacking green
parts died in it. The growth of developed pea plants languished in it in the sun [p. 209]
and could not maintain itself in the shade. Epilobium hirsutum, Lythrum salicaria,
and Polygonum persicaria thrived in it as in ordinary air. After having grown in this
gas for 6 weeks, in the sun, they had not decomposed it. They had increased its
volume, as they had that of nitrogen gas, by a corresponding amount of oxygen gas.
In total darkness, they increased this atmosphere by carbon dioxide gas.

6.4 Of Plant Growth in Hydrogen Gas

All of the seeds that I tested, without exception, failed to germinate in hydrogen
gas, when placed in it with a small amount of water. Senebier observed that they
bring about a very considerable reduction in the volume of this gas, by putrefying.
The gaseous fluid that is a residue of this condensation is carbon monoxide gas. The
carbon dioxide gas that the seeds form from their own substance is decomposed by
the hydrogen gas, with the aid of the caloric [p. 210] disengaged in fermentation.
Water forms, and the carbon dioxide, freed of a part of its oxygen, is converted into
carbon monoxide gas.
Green plants grow in hydrogen gas very nearly as they do in nitrogen gas. Plants
that languish in nitrogen gas also languish in hydrogen gas, and those that thrive in
nitrogen thrive in hydrogen. If there is some difference in the vigor of plants growing
in these two gases, it seemed to me that it was in favor of growth in nitrogen gas.
Plants grown in hydrogen gas have been said to develop a darker green color, but I
did not notice this effect.
I consistently observed that marsh plants, such as Lythrum salicaria and
Polygonum persicaria, that I grew in the sun for 5 or 6 weeks in hydrogen gas left
little or no oxygen gas there, whereas in the same period in nitrogen gas they always
left fifteen or twenty times their volume of oxygen gas. This result is very likely due
to the fact that, in hydrogen gas, the plants cannot completely decompose [p. 211]

7
The idea that hydrogen is an essential component of oxide of carbon gas may be based on evidence
that is still too indirect to be recognized as certain.—I will note, however, in favor of oxycarburetted
hydrogen, that it is odd that plants do not decompose carbon monoxide gas and that they never reduce
carbon dioxide into carbon monoxide gas directly, or without the presence of hydrogen gas. [Even
after it was established that hydrogen was not a component of this gas, Berthollet continued to
believe that it was.]
6.5 Of Plant Growth in a Vacuum 81

all the carbon dioxide gas that they form there, because a large part of this carbon
dioxide is itself decomposed by the hydrogen gas. As I have already observed, water
and carbon monoxide gas are the products of this decomposition. The oxygen gas
that the plants would have emitted without the hydrogen gas is concealed in these two
compounds. One hundred parts (60 cubic inches) of hydrogen gas that had served as
atmosphere for 5 weeks to a Lythrum salicaria plant exposed to the sun could not be
diminished appreciably by nitric oxide. The gaseous fluid then contained no carbon
dioxide, but when I ignited it by an electric spark, with the correct proportion of
oxygen gas, it left a residue of water, 3 parts carbon dioxide gas, and 4 parts nitrogen
gas. Hydrogen gas in which there had been no plants and that had been placed along-
side the preceding, under the same conditions, did not produce a detectable amount
of carbon dioxide gas when burned.
The volume of the atmosphere of the Lythrum decreased during growth, but to
the same degree as hydrogen gas that was confined [p. 212] by water8 and not in
contact with Lythrum. Considering that this plant formed carbon monoxide gas and
that the addition of this gas was not detectable through an increase in the volume of
the gaseous fluid contained by the receptacle, there must have been a compensation
here, and the hydrogen gas was diminished by the effect of the plant growth. It does
not appear that the plants absorbed this gas. They condensed it by forming water
through an indirect route.

6.5 Of Plant Growth in a Vacuum

In the vacuum produced by the best air pumps, some seeds may give signs of the
beginning of germination. This result is not surprising, as it has been shown that this
vacuum cannot [p. 213] be complete, and because, moreover, even the most perfect
of these machines are never joined exactly enough to prevent the ingress of external
air entirely. The pump that I used produced a vacuum in which the barometer at
first stayed at 3/4 of a line, when there was no water in the receptacle. In 24 hours,
it climbed by 1 line, through the imperceptible introduction of external air. Peas
germinated in it after 12 days, even when I renewed the vacuum every day, but the
development never went beyond the first appearance of the radicle.
Fully developed, leaf-bearing pea plants, as well as favas and haricots, always
died after 3 days in a vacuum, either in the sun or in the shade. They also perished
in nitrogen gas in the shade, but they often survived in it in the sun. No thin-leaved
plant seemed to me able to maintain its growth in a vacuum in the sun. The thickest
joints or leaves of Cactus opuntia survived more than a month in a vacuum in the sun.
Only their epidermis dried out, in part. After this experiment, these leaves regained

8
For a year I kept hydrogen gas in a receptacle containing water and resting on mercury. The water
absorbed approximately its own volume of gas, but no more. When I withdrew the metallic fluid,
there was no limit to the absorption of the hydrogen gas by the water. It is quite probable that, as
Guyton conjectured, this gas was released by the water to the atmospheric air.
82 6 Of Plant Growth in Oxygen-Free Environments

their vigor [p. 214] when I planted them in plant mold. The thinnest cactus leaves
were dead after several days in a vacuum in the sun.
A one-foot-tall Polygonum persicaria plant, with its roots immersed in 1 ounce of
water, was placed in the vacuum that I mentioned and that I renewed each day. The
plant elongated several inches. It was only withdrawn after 6 weeks, as healthy as it
was before the experiment except for two or three leaves that had yellowed, in the
vicinity of the roots. I obtained the same results with Epilobium molle, E. hirsutum,
Lythrum salicaria, and Inula dysenterica. All of these plants thrived as well in a
vacuum as under a receptacle full of ordinary air. Their transpiration was the same
in both cases.
The experiments just described were carried out in full daylight but protected
from the direct effects of sunlight. The plants wilted as soon as they were exposed
to it, even if the rays were weak and produced no effect on similar plants confined
in receptacles filled with ordinary air or pure nitrogen gas. [p. 215] It is likely that
plants survive in a vacuum only with the aid of the oxygen gas trapped in their
parenchyma, and that the sun harms the plants in expelling this gas by the expansion
it causes in the gas. Solar rays do not exert the same effect on plants in nitrogen gas
because the oxygen gas that the plants contain is compressed by the full weight of
the atmosphere.
Plants do not seem to survive and develop in a vacuum, except with the aid of the
oxygen gas eliminated by their green parts. Seeds that have only their radicles die in
it. Woody plants in the springtime were unable to open their leaf buds in it.
The buds of rose, lily, and carnation were as if paralyzed there. It is evident
that, in many respects, plants behave in a vacuum as in nitrogen gas, hydrogen gas,
etc. The suppression of the weight of the atmosphere, or the expansion that the plant
necessarily undergoes by this suppression, does not appear to have a very appreciable
effect on its growth. Only the removal of oxygen gas is harmful to it.

6.6 Summary [p. 216]

Only plants provided with their green parts seem to be able to grow in oxygen-free
environments, because these plants release this gas there. When this gas is removed
as the plants form it, their development is arrested. The amount of oxygen gas that
some require to survive, without developing, is inappreciable.
Plants do not absorb nitrogen gas. Neither do they absorb hydrogen gas. They
reduce the amount of the latter a little, but this decrease occurs because the hydrogen
gas decomposes the carbon dioxide gas formed by the plant. The result of this
decomposition is water and carbon monoxide gas.
The green parts emit less oxygen gas in hydrogen gas than in nitrogen gas.
Green plants growing in carbon monoxide gas in the sun do not decompose this
gas. They add oxygen gas to it.
Green plants grow in the vacuum produced by an air pump as in nitrogen gas,
provided that the experiment takes place away from direct sunlight.
[p. 217] Chapter 7
Of the Fixation and Decomposition of Water
by Plants

7.1 Investigations on the Fixation of Water by Plants Growing

in Atmospheric Air Freed of Carbon Dioxide Gas

Those authors who have addressed the question of the decomposition of water by
plants have advanced this subject only by conjectures unsupported by any direct
experiment. Senebier never saw a plant that had been deprived of contact with carbon
dioxide gas produce a volume of oxygen gas exceeding the volume of the plant itself
(Physiologie végétale, Vol. 3, pp. 228 and following). This effect seemed to him
insufficient to prove the decomposition of water. Moreover, he recognized that this
small amount [p. 218] of oxygen gas, less than the volume of the leaf, was due to
the carbon dioxide gas lodged in its parenchyma. This learned physiologist does not,
however, reject the decomposition of water by plants. He considers it likely based on
very scholarly considerations and on inferences drawn from the germination of some
seeds in pure water without apparent contact with oxygen gas. But this observation,
even if correct, would still not be a proof. I have shown that this result should be
attributed to the air dissolved in the water and foreign to its composition. Ingen-Housz
regarded water as a simple substance. Nevertheless some of his observations seem
to suggest the decomposition of this fluid. He observed succulent plants ameliorate
an atmosphere of ordinary air that was not renewed, but, as his eudiometric methods
were unable to give absolute amounts, and as the volume of the plant and that of
the gas produced are unknown, it is still conceivable that the volume of oxygen gas
eliminated is less than the volume of the plant used in the experiment. Spallanzani,1
who obtained the same results [p. 219] with succulents submerged in limewater,
also leaves us uncertain about the volume of the plant. Senebier, in analyzing and
repeating these last experiments, showed clearly that they were not conclusive, for
the same reasons as I gave above.
The famous Berthollet, whose opinion always carries such great weight, accepted
the decomposition of water in vegetation, but by reasoning rather than by new

1
Journal de Physique, 1799. [De Saussure cited the date as “pluviose, an 7” (the 5th month of
year 7 of the French Republican calendar).]

J. F. Hill, Chemical Research on Plant Growth, 83

DOI 10.1007/978-1-4614-4136-6_7, © Springer Science+Business Media New York 2013
84 7 Of the Fixation and Decomposition of Water by Plants

experiments. Several authors, whom it would be superfluous to cite, accepted this

decomposition, but without proofs and without even discussing the question.
Plants growing with the aid of pure water, in oxygen gas or in ordinary air that was
washed in limewater before the experiment, may gain in fresh weight under these con-
ditions, without losing or letting dry any of their parts. This result proves neither the
decomposition of water, nor even the fixation of the oxygen and hydrogen of water in
the plant. The plant may gain weight solely by the introduction of liquid water into the
sap vessels, or of water of vegetation into the cellular tissue, for experiment [p. 220]
showed long ago that this water may increase with soil moisture and with etiolation.
We can judge whether the dry or solid substance of plants is increased through
the fixation of the constituent parts of water, by drying at room temperature a plant
similar to, and of the same weight as, the one grown in a closed vessel with the
aid of pure water and oxygen gas, and noting whether the dry weight of the plant
grown under the experimental conditions is greater than it would have been if the
plant had been dried before the experiment, as was the dried plant that serves as a
comparison. It is superfluous to add that the two plants must be collected at the same
stage of maturity, from the same soil, and that they must always be weighed at the
same readings of the thermometer and hygrometer.
The numerous experiments that I conducted using these procedures showed me
that plants grown with the aid of water alone, in a closed vessel, in atmospheric air
freed of its carbon dioxide, do not increase their dry weight appreciably, and that,
if they do increase it, it is by a very small, very limited amount, or, in other words,
by an amount that cannot [p. 221] be further increased if their growth is continued
for a longer time. From the numerous experiments I conducted on this subject, I will
select those that gave me the most marked results.
In the month of June, in a receptacle containing 4.95 liters (250 cubic inches) of
ordinary air freed of its carbon dioxide, I placed three loosestrife plants (Lysimachia
vulgaris), whose roots were immersed, under the receptacle, in 1 centiliter of distilled
water. They were exposed to the alternating effects of darkness and sunlight that was
moderated when too intense. These plants weighed, in the green state, 6.96 grams
(129-1/4 grains), and they displaced 10 cubic centimeters (1/2 cubic inch).
I collected other loosestrifes, of the weight I just indicated, and I dried them at
room temperature. Their dry plant substance weighed 2.05 grams (38-1/2 grains) at
a set reading of the thermometer and hygrometer. These plants had been uprooted at
the same time and place as those grown under the receptacle.
After 8 days I removed the experimental plants from their confinement. They
were perfectly healthy. [p. 222] They had elongated, and they had not detectably
changed the air around them, either in purity or in volume. They then weighed, in
the green state, 7.43 grams (141 grains) and, after being dried to the same degree
as the previous plants, 2.159 grams (40-1/2 grains). They had thus increased their
dry matter by 2.159 − 2.05 = 109 milligrams (2 grains).2 If these 109 milligrams had

2
These 2 grains include the oxygen that the plant assimilated from atmospheric air, but the amount
of carbon dioxide that the plant formed and decomposed for this assimilation is so small that I
cannot count this increase in my results.
7.2 Fixation of Water by Plants Growing in a Mixture of Ordinary Air . . . 85

been formed only from the hydrogen of water, the plants would have had to eliminate,
under the receptacle, all the oxygen with which these 109 milligrams of hydrogen
were combined, that is, at least 436 cubic centimeters (22 cubic inches) of oxygen
gas, an amount that would have been very obvious, either through an increase in the
volume of the air or by a eudiometric test. Now, as they did not emit an appreciable
amount of oxygen gas in this experiment, it follows that they assimilated, in addition
to the hydrogen [p. 223] of the water, almost all of its oxygen.
The experiment that I just reported was repeated for double and triple the length
of time. That is, the plant was grown in a closed vessel for a fortnight and for a
month. But loosestrifes of the same weight as the previous ones were never able to
increase their substance by more than 2 grains, and often did not increase it at all,
although they elongated by several inches.
Seven periwinkle plants (Vinca minor L.), having a combined weight of
7.855 grams (148 grains) and containing 2.375 grams (44-3/4 grains) of dry plant
matter, were placed under a receptacle, in the same circumstances as those in the
preceding experiment. They displaced 10 cubic centimeters (1/2 cubic inch). These
plants did not deteriorate in the least. They did not change their atmosphere, either
in purity or in volume. After drying, they weighed 46-1/2 grains. Their dry plant
matter had therefore increased by 93 milligrams (1-3/4 grains), which could not be
attributed solely to the assimilation of hydrogen gas from the water, since this result
would have presumed the [p. 224] elimination of 19 cubic inches of oxygen gas,
which did not occur. I repeated this experiment several times, prolonging it greatly,
but the assimilation of water by the plant was never any greater.
Two aquatic mint plants, whose green weight was 157 grains, increased their
green weight by 17 grains by growing in a closed vessel, with the aid of pure water,
in atmospheric air [presumably freed of its carbon dioxide, as in the preceding
experiments]. But they did not increase their dry plant matter by more than 1 grain.
When the experiment was prolonged, on other, similar plants, the increase was no
greater. In this experiment, as in the previous ones, the atmosphere of the plant
changed neither in purity nor in volume.
I must note that I have considered only the results in which the plants did not
suffer harm, for if growth languishes, the plants, far from gaining weight, lose it.

7.2 Fixation of Water by Plants Growing in a Mixture

of Ordinary Air and Carbon Dioxide Gas [p. 225]

The previous experiments on the fixation of water yielded such trivial results that
they are scarcely beyond the limits of experimental error, but I believe that this is
not hard to explain. It is very likely that the amounts of oxygen and hydrogen in
plants cannot be increased beyond certain limits without correspondingly increasing
the proportion of their carbon. Consequently, I grew plants in a mixture of ordinary
air and carbon dioxide gas, so that they could assimilate carbon. In all of the cases
in which the plants were not harmed, the results were then more pronounced. The
86 7 Of the Fixation and Decomposition of Water by Plants

plants clearly increased their dry matter by an amount exceeding what they drew
from the elements of the carbon dioxide.
I will now return [p. 226] to the experiments that I reported in Sect. 2.4, on the
decomposition of carbon dioxide gas. I did not give all the details then. We saw in
the First Experiment that seven periwinkle plants, grown with the aid of pure water
in a mixture of ordinary air and carbon dioxide gas, assimilated the carbon contained
in 431 cubic centimeters (21-3/4 cubic inches) of carbon dioxide, or an amount of
carbon equal, according to Lavoisier, to 217 milligrams (4.2 grains). The plants also
appropriated, from the same gas, 139 cubic centimeters (7 cubic inches) of oxygen
gas, but I must disregard this last assimilation, because they lost, in turn, an equal
amount of nitrogen gas. These two opposing effects nearly cancel each other. Be-
fore the experiment, these plants weighed, green, 8.955 grams (168-3/4 grains) and
contained 2.707 grams (51 grains) of dry plant matter. After decomposition of the
carbon dioxide, they yielded, upon drying, 3.237 grams (61 grains). They thus in-
creased their dry matter by 531 milligrams (10 grains), of which only 217 milligrams
(4.2 grains) can be attributed to the carbon dioxide. [p. 227] The periwinkles in this
experiment thus fixed or solidified 315 milligrams (5.8 grains) of water.
Two aquatic mint plants (Sect. 2.4, Second Experiment) decomposed 309 cubic
centimeters (15.6 cubic inches) of carbon dioxide gas, which contain, according to
Lavoisier, 159 milligrams (3 grains) of carbon. They yielded, before the experiment,
1.698 grams (32 grains) of dry plant matter, and after the experiment, 2.016 grams
(38 grains) of the same substance. They thus increased their dry plant matter by
318 milligrams (6 grains), of which 159 milligrams (3 grains) must be attributed to
the fixation of water.
These results require conditions that are rarely encountered: The plant must remain
healthy in all its parts. If a single leaf wilts under the receptacle, or if the roots
suffer harm, the dry weight of the plant often decreases, even though the plant has
decomposed a large amount of carbon dioxide gas.

7.3 Of the Decomposition of Water by Plants [p. 228]

Since plants appropriate the oxygen and hydrogen of water, we may suppose that,
under certain conditions, they exhale the oxygen that was a component of this liquid.
Dead plants in a heap, fermenting without contact with air, form carbon dioxide
entirely from their own substance. In this process, and with the aid of the heat
produced during fermentation, the oxygen of the water that was fixed or solidified
during growth unites with the carbon to produce carbon dioxide gas. And growing
plants, in decomposing the latter, eliminate the oxygen that had originally belonged
to the water. But in no case do they appear to decompose water directly, appropriating
its hydrogen and immediately releasing its other element in the form of oxygen gas.
Green plants growing day and night in nitrogen gas emit to it several times their
volume of oxygen gas because, having lacked [p. 229] contact with oxygen gas
during the first stages of their growth, they form, entirely from their own substance,
7.3 Of the Decomposition of Water by Plants 87

the carbon dioxide gas that they decompose. But the same plants, or at least those that
are not fleshy, do not add oxygen gas to an atmosphere of ordinary air or of oxygen
gas in which they are growing day and night, and it is only in these atmospheres that
we can judge whether they decompose water directly, because the carbon dioxide gas
that they form there results from the combination of their carbon with the surrounding
oxygen gas and is not a product entirely of their own substance.3
[p. 230] I grew Lythrum salicaria with the aid of water for 4 months in the sun,
in 40 cubic inches of atmospheric air closed by water and mercury. I renewed [i.e.,
replaced] this plant several times so that it would not deteriorate. After this lengthy
trial, the air of the receptacle contained 1/100 less oxygen gas than it had before
having been subjected to the effects of plant growth, and yet growth was still very
vigorous there.
Likewise, I grew Polygonum persicaria plants under equivalent conditions for
3 months, and the ordinary air surrounding them did not improve or change
detectably.
All the other thin-leaved plants that I subjected to the same trials gave me the
same results.
The only result that might be surprising is that the plants did not corrupt their
atmosphere more appreciably by absorbing its oxygen gas in the decomposition of
the carbon dioxide gas that they formed there. But it must be noted [p. 231] that
the roots, submerged in water that was not renewed, being deprived of contact with
oxygen gas, must produce, entirely from their own substance, a small amount of
carbon dioxide gas, and that the decomposition of the latter by the leaves may very
nearly offset the absorption of oxygen gas.
Indeed I often noted in these experiments that the roots released a small amount of
air, especially when they were exposed to the sun. In the shade, this gas was absorbed
by the plant in the proportion in which it was produced.
If plants found excess oxygen gas in the water, they would, in decomposing the
carbon dioxide gas, eliminate all of the oxygen of the latter gas, but the experiments
reported in Chap. 2 showed that they assimilated a part of this oxygen. Thus the
oxygen found in the water that nourished them did not suffice for their nutrition.
At first glance, Cactus opuntia and probably other succulents4 seem [p. 232] to
give results that are inconsistent with those I just reported. Cactus growing day and

3
The results of the following experiments do not contradict those I just mentioned. I grew two
aquatic mint plants in the sun for 3 weeks, with the aid of water, under two identical receptacles
filled with ordinary air. In each receptacle, beside the growing plant, I placed a mint plant that was
wilted but not desiccated and not completely dead. But, in one of the receptacles, this wilted mint
was covered by the water that served to close the receptacle, whereas in the other receptacle, the
wilted mint was suspended in the air contained therein. The plant growing near the wilted plant
in air did not improve its atmosphere, whereas the mint growing with the submerged plant added
several times its volume of oxygen gas to the ordinary air surrounding it. The difference in the origin
of the carbon dioxide produced by the wilted plants under these two circumstances explains these
results. The submerged plant produced carbon dioxide entirely from its own substance, whereas the
one that was in air supplied only the carbon component of the carbon dioxide.
4
I did the same experiments on Sempervivum tectorum, Stapelia variegata, and Sedum telephium,
but they could not tolerate a very humid atmosphere long enough without being harmed.
88 7 Of the Fixation and Decomposition of Water by Plants

night under the same receptacle filled with atmospheric air previously freed of its
carbon dioxide gas may emit several times its volume of oxygen gas. I performed the
following experiment on this plant. I repeated it four times under the same conditions
and with the same results. The details that I am going to report will show the exactness
of my methods.
At sunset, in June, I cut a leaf or joint of cactus that displaced 15 cubic centimeters
(3/4 cubic inch). It was placed in a stemmed glass containing, at its bottom, 10 cubic
centimeters (1/2 cubic inch) of distilled water. The leaf touched the surface of this
liquid only by its tip, or wedge-shaped extremity.
All of this was covered by a sufficiently elongated receptacle that a change of
1/4 cubic inch in the volume of the atmospheric air it contained would be de-
tectable. This air, freed of carbon dioxide gas, occupied 847 cubic centimeters
(42-3/4 cubic inches). When the water in the glass was so depleted that it was
no longer in contact with the cactus, I was able to replenish the water, without
[p. 233] removing the cactus, by dipping the receptacle in boiled water and lift-
ing it out. The receptacle was closed by this liquid, which rose inside it, and the
bottom of the basin on which it rested was filled with mercury. After 31 days, I re-
moved the plant, which seemed as healthy as when I had placed it in the experiment.
It had grown roots 2.7 centimeters [text erroneously says “decimeters”] (1 inch)
in length, and its atmosphere had increased by 69 cubic centimeters (3-1/2 cubic
inches). The eudiometer indicated, in the 917 cubic centimeters (46-1/4 cubic inches)
of air (taking the mean of five observations), 25/100 oxygen gas, and, before the in-
troduction of the plant, 21/100 of this gas. This atmosphere contained no carbon
dioxide. It follows from these data that the cactus eliminated, in 1 month in the sun,
3-1/2 times its volume of oxygen gas, which might be attributable to the decomposi-
tion of water.
But the plant does not appear to have brought about this decomposition directly, or
to have immediately appropriated to itself the hydrogen of the water, in eliminating
its oxygen. An in-depth examination leads us to believe [p. 234] only that it produced,
in the sun, carbon dioxide entirely from its own substance, and that it decomposed
this gas.
It should be noted (1) that the inner parts of the cactus, due to the plant’s small
surface area and the low porosity of its epidermis,5 are not in direct contact with
oxygen gas when the sun has expelled the free air that had penetrated there. After
this expulsion, they are in about the same condition as they would be if they had
been subjected to distillation in a closed vessel or had grown in nitrogen gas. They
form carbon dioxide gas entirely from their own substance. It is not the same with
thin leaves. All their parts are in contact with the surrounding air, not only because
their parenchyma is not as thick, but also because their epidermis is, other things
being equal, pitted with many more pores. The improvement made by the cactus in
its atmosphere is very slight. [p. 235] Under the best circumstances, it amounts to
only 1/13 of the volume of the leaf in 24 hours.

5
The epidermis of the leaves of succulent plants has many fewer pores than that of the leaves of
other plants. We owe this curious observation to DeCandolle.
7.4 Summary 89

(2) A direct experiment showed me that cactus emitted oxygen gas only through
the decomposition of carbon dioxide gas. I grew some cactuses for 1 month under the
same conditions as those under which they had produced 3-1/2 times their volume of
oxygen gas, but I suspended at the upper part of the receptacle a vessel full of potash.
The cactus then added no more oxygen gas to its atmosphere. It did not change
the atmosphere at all, and the potash became effervescent. The plant, however, was
not harmed at all. It grew roots, as in the previous experiment. The potash, in this
trial, did not remove from the plant the carbon dioxide gas that the plant formed in
darkness with the surrounding oxygen gas. The cactus retained this carbon dioxide
in its thick parenchyma. The potash removed from the plant only the gas that the
plant formed from its own substance. When I put limewater under the receptacle, I
saw that the latter carbon dioxide was formed only in the sunlight.
(3) I showed in Sect. 2.4, Fifth Experiment, that a cactus growing with the aid
of [p. 236] water, in a mixture of ordinary air and carbon dioxide gas, retains, in
decomposing the latter gas, a considerable part of the oxygen it contains, as do other
plants. This plant thus does not find enough oxygen in water alone. Therefore, under
the conditions most favorable to its growth, the plant is far from having to reject this
element of water as superfluous.
(4) I investigated, with the utmost care, whether cacti that had emitted 3-1/2 times
their volume of oxygen gas in growing for a month in atmospheric air freed of carbon
dioxide gas had, in this process, increased their dry matter,6 but I found no increase.
It even seemed to me that the weight of this substance always decreased slightly.

7.4 Summary

Plants appropriate the oxygen and hydrogen of water, causing the water to lose its
liquid state. [p. 237] This assimilation is pronounced only when the plants incorporate
carbon at the same time.
Water fixed or solidified by plants probably cannot lose its oxygen in a gaseous
form except after the death of the plant or one of its parts. When plants that have
assimilated the oxygen and hydrogen of water begin to ferment without having free
contact with oxygen gas, they form carbon dioxide gas entirely from their own
substance. The oxygen of the fixed water can unite with their carbon to form this
carbon dioxide gas, and plants or their growing parts, in disengaging the oxygen from
this latter gas, indirectly release an element that had originally belonged to water.
Thus, through the combination of plant growth and fermentation that proceeds
without contact with air, the water may release its principal element in the form of
oxygen gas.
But in no case do plants decompose water directly, assimilating its hydrogen and
eliminating its oxygen in a gaseous state. They exhale oxygen gas only through the
direct decomposition of carbon dioxide gas.

6
To dry the cactus, I immersed it for several moments in boiling water and then exposed it to the sun.
90 7 Of the Fixation and Decomposition of Water by Plants

[p. 238] Thin-leaved plants, growing with the aid of pure water in a mixture of
oxygen and nitrogen gases, under the alternating effects of sunlight and darkness,
do not add oxygen gas to the atmosphere and give no external sign of the direct
decomposition of water. The oxygen gas that the plants emit in pure nitrogen gas or
under water cannot be attributed to the direct decomposition of water because they
form carbon dioxide gas entirely from their own substance whenever they are in an
oxygen-free environment.
Some succulents, when growing in ordinary air freed of its carbon dioxide gas,
add to this atmosphere an amount of oxygen gas that exceeds several times their
volume. But this gas, although it could have originally been a component of water,
arises, in the final analysis, only from the decomposition of carbon dioxide that the
plants form, in the sun, entirely from their own substance. For when a substance that
can absorb this carbon dioxide is placed near them, they no longer add oxygen gas
to the environment in which they are growing day and night. They no longer give
any indication of the direct decomposition of water, although their growth [p. 239]
is vigorous. The exclusive property that succulents have of forming carbon dioxide
gas from their own substance arises from the low porosity of their epidermis, or the
limited contact of their inner parts with the surrounding oxygen gas.
It cannot be doubted that most of the hydrogen that annual plants acquire in
developing in the open air, with the aid of distilled water, originates in this liquid,
which the plants solidify. We can say as much of their oxygen, for we can judge,
whether by the amount of carbon dioxide gas that these plants can decompose in a
given time, or by the small change that they make in ordinary air, that the amount
of oxygen that they obtain from atmospheric gases is insufficient to account for the
oxygen that they acquire in the short period of their development. We must not forget
that water is the most abundant product of the decomposition of most dry plants, or
that oxygen is their main element.
[p. 240] Chapter 8
Of the Absorption of Solutions by Plant Roots

8.1 Water and Gas are Insufficient Nourishment to Bring about

the Full Development of Plants

Plant roots, as filters, are too fine to be able to absorb substances other than fluids.
If they admit solids, these must be so diluted, so divided, that their dispersion in
the liquid has all the characteristics of a true solution.1 [p. 241] Research on the
absorption of solutions by plants is very important for the theory of their nutrition
because these studies help us assess the amount and kind of nourishment that plants
can assimilate from the soil through their roots.
Tull, van Helmont, and even some modern naturalists have tried to show that
plants draw only water from plant mold,2 and that manures [p. 242] only act on
the soil, by providing plants a support more or less suitable for retaining heat and
moisture. These authors supposed that the vital force, animal as well as plant, could,
by decomposing or combining atmospheric air and water in various ways, produce
all the substances, even the salts, earths, and metals, that analysis and incineration
reveal in plants. This confused idea is no more provable than is that of making
gold from substances that contain none. Before we have recourse to transmutations
that are unintelligible, miraculous, and contrary to all known observations, we must
make certain that plants are not finding these substances, already formed, in the

1
For a month I nourished thirty Polygonum persicaria and peppermint plants with distilled water
to which I had added a certain weight of finely divided silica, which remained partly suspended
with the aid of a small amount of sugar dissolved in the liquid. After the experiment, I did not find,
either by incinerating the plants or by closely examining the residue of the absorbed liquor, that
this earth had penetrated the plant appreciably. Bonnet caused some plants to absorb ink, but only
an imponderable amount of the undissolved coloring matter was absorbed. It would have passed
much more profusely through the most perfect filters we can make.
2
I think it is unnecessary to list here the experiments of van Helmont, Tillet, Bonnet, and Duhamel,
who were the main supporters of this theory. The flaws in their methods were sufficiently demon-
strated by Bergman, Kirwan, and Hassenfratz. They noted especially that the vessels in which Tillet
and van Helmont grew plants were porous and buried in plant mold, which could influence the
results these authors obtained. Duhamel’s oak was moistened with water naturally rich in extractive
material. Bonnet’s plants had for support plant substances more or less soluble in water.

J. F. Hill, Chemical Research on Plant Growth, 91

DOI 10.1007/978-1-4614-4136-6_8, © Springer Science+Business Media New York 2013
92 8 Of the Absorption of Solutions by Plant Roots

environments in which the plants develop, and that they do not draw the substances
from that source.
Other authors have come closer to a plausible explanation in supposing that all
plant constituents, except water, were provided to plants in a gaseous form. But,
based on the results of direct observation, this hypothesis is not wholly tenable.
Oxygen and carbon dioxide are the only known gaseous substances with which
plants can [p. 243] nourish themselves in our atmosphere. Experiment shows that
most plants do not assimilate nitrogen gas, yet nitrogen composes an essential part
of plants. It is consistently found in wood, extracts, and the green coloring matter.
Plants contain earths that can, as in animals, contribute to forming their solid or
bony parts. We cannot prove that these substances exist in a gaseous form in our
atmosphere, although we see them there in a suspended or vaporous state. But the
earths have been shown to be present in springs that have washed over plant molds
and that enter the roots. It has also been shown that these same spring waters hold
in solution extractive substances of which nitrogen is an esssential part, and that
soil fertility is, in some respects and within certain limits, dependent on the amount
and nature of the water-soluble substances it contains. Finally, it is evident that, by
appropriating these substances, plants eventually exhaust the soil or make it sterile.
Plants that grow naturally on dry rocks or in pure sand find, in the remains of
plants and animals [p. 244] floating in our atmosphere, the nutriments that the gases
composing the atmosphere cannot provide. These bodies settle on the leaves, dissolve
in the water that condenses there, and penetrate the interior. Rain water absorbed by
the roots is rich in the same materials.
Very few plants can develop fully by taking all their sustenance from our atmo-
sphere in this way. Such scant nutrition suffices only for those whose structure is
adapted to their situation, only for those that are perennial and whose very slow
development is proportionate to the small quantity of nutrients assimilated. Most of
them, such as the mosses, ferns, succulents, and pines, are evergreen. Their leaves
decompose carbon dioxide gas all year round. They continuously offer stopping
places for the corpuscles floating in our atmosphere. They transpire little. They are
either leathery or succulent, and in these two forms they lose only a small amount of
their carbon through the effects of the surrounding oxygen gas. But almost no annual
plants occur in a soil [p. 245] that lacks plant mold. They must grow or develop too
quickly to be able to find in our atmosphere the large amount of nutrients that they
consume.
Annual plants languish as soon as they have exhausted the nourishment contained
in the substance of their seeds. I tried to make seeds of fava, haricot, pea, and cress
develop by giving them for support pure sand or horsehair contained in funnels that
drained away the excess distilled water with which I wetted them. They usually
bloomed, but their seeds were never able to mature. Nevertheless, I carried out
variations of these experiments with all possible care for 5 consecutive years. Giobert,
Hassenfratz and other naturalists preceded me in this research without obtaining any
better results.
Those who believe that atmospheric air and water are the sole constituents
and nutrients of plants object that sand, hair, and other, similar supports are
8.2 Do Plants Absorb the Substances that are Dissolved . . . 93

not suitable media for providing the amount of water needed to maintain plant
growth.
This objection would be justified if [p. 246] many observations had not shown
that a soil may be largely stripped of its growth-promoting properties yet still be
indistinguishable from the most fertile soil by physical qualities, namely, retention
of heat and moisture, root penetrability, and serving as root support. Thus, it is well
known that a soil that is washed-out by spring or rain waters loses its fertility, yet
retains the same appearance, color, weight, and consistency. When I (as described in
Sect. 5.2) stripped an almost-pure humus of most of its extractive materials, it retained
its external characteristics. Its fertility, however, decreased. If it were possible to
divest a humus of its soluble plant parts entirely, it might be compared to pure
sand with respect to its growth-supporting properties, but complete despoliation is
impossible, and we can reason here only by approximation.
If manures promoted plant nutrition principally by the gases that they develop, a
field that is unsown and produces no crop should exhaust itself to the same extent as
one that nurtures a rich harvest. Farming operations, however, all demonstrate the
contrary. [p. 247] We find that harvests impoverish the soil, and do so more or less
according to their nature. In general, annual plants that are rich in plant substance
and transpire profusely exhaust the soil more than do perennials, which develop less
rapidly, and more than annuals having succulent leaves, which transpire little, such
as peas, favas, and buckwheat.3
We can make another observation, which is a consequence of the foregoing and
which should serve to prove it. It is that, other things being equal, the plants that are
the most exhausting are those that require soil that is richest in nutrients.

8.2 Do Plants Absorb the Substances that are Dissolved in

Water in the Same Proportion as they Absorb the Water?

I prepared several solutions, each containing, in 793 cubic centimeters (40 cubic
inches) of distilled water, [p. 248] 637 milligrams (12 grains) of the substance
indicated below. I will assume these 12 grains equal to 100 parts.
1. Solution containing 100 parts potassium chloride [“muriate of potash”]
2. Solution containing 100 parts sodium chloride [“muriate of soda”]
3. Solution containing 100 parts calcium nitrate [“nitrate of lime”]
4. Solution containing 100 parts powdered sodium sulfate [“powdered sulfate of
soda”]
5. Solution containing 100 parts ammonium chloride [“muriate of ammonia”]
6. Solution containing 100 parts calcium acetate [“acetate of lime”]
7. Solution containing 100 parts copper sulfate
8. Solution containing 100 parts crystallized sugar

3
Bibliothèque Britannique. (Mémoires sur la culture du Blé. Agriculture, Vol. 5, p. 499).
94 8 Of the Absorption of Solutions by Plant Roots

9. Solution containing 100 parts gum arabic

10. Solution containing 100 parts 25 parts humus extract4
I soaked root-bearing plants of Polygonum persicaria or Bidens cannabina in each
of these solutions. I primarily used marsh plants in these studies, because they would
be less harmed by [p. 249] the superabundant water that nourished them. I will add
that, before putting them into the experiment, I kept them for several days in distilled
water, until their roots began to lengthen.
The Polygonum plants grew in the shade for 5 weeks in the solutions of potassium
chloride, calcium nitrate, sodium chloride, sodium sulfate, and humus extract, de-
veloping their roots in these solutions. The plants always languished in ammonium
chloride, without any development. They could maintain themselves in sugar water
only if the solution, which putrefied very rapidly, was renewed. They died in 8 or 10
days in the gummy water and in the solution of calcium acetate. They could survive
no more than 2 or 3 days in the solution of copper sulfate.
The Bidens plants showed very nearly the same responses in these different
solutions. In general, the Bidens tolerated them less well than did the Polygonum.
When I investigated the proportions in which the solutes were absorbed relative
to the water by root suction, [p. 250] I used the same solutions, but I ended the
experiment as soon as the plants had taken up precisely half of the liquid nourishing
them, that is, 397 cubic centimeters (20 cubic inches) of solution. The plants were
numerous enough that this absorption was completed in 2 days. If absorption had
continued for a longer time, the roots would have putrefied in the solutions that were
unsuited to them. Moreover, the sugar, gum, and humus extract could have lost most
of their constituents through fermentation.
Analysis of the 397 cubic centimeters (20 cubic inches) of solution that the plants
left in each vessel after suction showed me the amount of salts5 that they had taken
up. [p. 251] They would have taken up 50 parts if they had absorbed the saline

4
I did not dissolve previously prepared and dried extract in water because, if this is done, the
solution still holds in suspension the part of the extract that was precipitated during evaporation.
Instead I made a cold infusion of humus in water. The evaporation of a part of this infusion showed
me that the infusion in which I grew the plants contained 25 parts of extract. The results obtained
with salts are more accurate than those with extract, sugar, and gum because these plant compounds
are always a little corrupted by contact with the roots.
5
The saline solutions were analyzed more accurately by reagents than by any other method.
I ascertained that the 100 parts of potassium chloride that I used formed, by silver nitrate, a
precipitate equal to 187-1/2 parts
100 parts of sodium chloride, by the same reagent 232-1/2 parts
100 parts of ammonium chloride, by the same reagent 270 parts
100 parts of calcium nitrate, by potassium oxalate [“oxalate of potash”] 78 parts
100 parts of calcium acetate, by the same reagent 81-1/4 parts
100 parts of powdered sodium sulfate, by barium acetate [“acetate of Baryta”] 167-1/3 parts
100 parts of copper sulfate, by the same reagent 94-1/2 parts
The solutions of gum, sugar, and humus extract were evaporated to dryness and the residue was
weighed.
8.2 Do Plants Absorb the Substances that are Dissolved . . . 95

substances in the same proportion as the water. But I found that the Polygonum, in
taking up half the liquid in the experiments, absorbed only
14.7 parts Potassium chloride
13 parts Sodium chloride
4 parts Calcium nitrate
14.4 parts Sodium sulfate
12 parts Ammonium chloride
8 parts Calcium acetate
47 parts Copper sulfate
9 parts Gum
29 parts Sugar
5 parts Humus extract

The Bidens absorbed:

16 parts Potassium chloride
15 parts Sodium chloride
8 parts Calcium nitrate
10 parts Sodium sulfate
17 parts Ammonium chloride
8 parts Calcium acetate
48 parts Copper sulfate
32 parts Sugar
8 parts Gum
6 parts Humus extract

[p. 252] It is evident that, in general, the plants absorbed all the substances that
I presented to them, but that they took in water in much greater proportion than the
solutes. It is also evident that the plants did not always take from this water more
of those nutrients that were most beneficial to them. Copper sulfate, which was the
most harmful, was absorbed the most. Gum and calcium acetate, which were very
unfavorable to growth, entered the plants only in small amounts.
I repeated these experiments several times, either in the same proportions or
in different ones, and I always obtained the same general results. The plants always
absorbed more chlorides and alkaline sulfates than calcium acetate or calcium nitrate.
They always absorbed more sugar than gum. But the results varied in detail. The
absolute amounts of salts, gum, and sugar that were absorbed were never the same in
two similar experiments. I soon noticed that the anomalies arose from the different
conditions of the roots, which absorbed more solutes if they were less vigorous. I
tried cutting off the roots, and the plants then very quickly suffered harm [p. 253] in
all the solutions. They absorbed two or three times more solute than in the previous
experiments.
I think that we must attribute the very great absorption of copper sulfate, especially,
to the disorganization that it causes in the roots. From the following results, in fact, it
will be evident that the addition of this salt to a solution of calcium acetate or calcium
nitrate causes these latter substances to penetrate the plants in larger amounts than
they would by themselves or if mixed with a salt less harmful to growth than copper
sulfate.
96 8 Of the Absorption of Solutions by Plant Roots

8.3 Do Plants Absorb Certain Substances in Preference to

Others, from a Liquid Containing Several Substances in
Solution?

So far, I have presented to the same plant only a single salt in solution. I will
now present several of them and see if the plant will make particular secretions
of them. [“Secrétion” is used here in the sense of absorption of substances by a
plant; see Glossary, Appendix 2.] I dissolved, [p. 254] in 793 cubic centimeters
(40 cubic inches) of water, two or three different salts, each weighing 637 milligrams
(12 grains). As in the preceding experiments, I set these 12 grains equal to 100 parts,
and I analyzed the residue of the solution when it had been reduced by suction to
precisely half its volume. The amount of salt contained in this residue, subtracted
from the amount that the liquid contained before the plants were introduced, indicated
to me the amount of salt that the plants had absorbed. The Polygonum absorbed 11.7
parts of sodium sulfate and 22 parts of sodium chloride, in absorbing up to half
of a solution containing 100 parts (637 milligrams) of each of these salts. From a
similar solution, the Bidens absorbed 7 parts of sodium sulfate and 20 parts of sodium
chloride. Based on this example, the results that I present in the following table can
be understood.

[p. 255] Weight of substances Weight of substances taken Weight of substances taken
dissolved in water before the up by Polygonum in absorb- up by Bidens in absorbing
experiment ing half the water of solution half the water of solution
100 parts of powdered sodium sulfate 11.7 7
[“powdered sulfate of soda”]
100 parts of sodium chloride [“muri- 226 20
ate of soda”]
100 parts of powdered sodium sulfate 127 10
100 parts of potassium chloride 17 17
[“muriate of potash”]
100 parts of calcium acetate [“acetate 8-1/4 5
of lime”]
100 parts of potassium chloride 338 16
100 parts of calcium nitrate [“nitrate 4-1/4 2
of lime”]
100 parts of ammonium chloride 16-1/2 15
[“muriate of ammonia”]
100 parts of calcium acetate 31 35
100 parts of copper sulfate 34 39
100 parts of calcium nitrate 17 9
100 parts of copper sulfate 34 36
100 parts of sodium sulfate 69 13
100 parts of sodium chloride 10 16
8.3 Do Plants Absorb Certain Substances in Preference . . . 97

Weight of substances dissolved in Weight of substances taken Weight of substances taken

water before the experiment up by Polygonum in absorb- up by Bidens in absorbing
ing half the water of solution half the water of solution
100 parts of acetate of lime Inapprec.quant. Inapprec.quant.
100 parts of gum 26 21
100 parts of sugar 3410 46

6
I precipitated the residue of root suction by barium acetate [“acetate of Baryta”]; the decanted
liquor was precipitated by silver nitrate
7
The analysis was done by the same procedure. It is evident that I made plants take in salts that
mutually decompose. But this decomposition was not carried out here because the solutions were
too dilute. Berthollet showed that the exchange of substances took place only when caused by
the force of crystallization, or, what amounts to the same thing, by the insolubility of the new
combination
8
The residue of the root suction was divided into two equal parts; one was precipitated by potassium
oxalate [“oxalate of potash”] and the other by silver nitrate
9
The residue of root suction was divided into two equal parts; one was precipitated by potassium
oxalate, and the other by barium acetate. After the separation of the barium sulfate [“sulfate of
Baryta”], the decanted liquor was precipitated by silver nitrate
10
I evaporated the residue to the consistency of syrup and then threw on it alcohol, which dissolved
the sugar and precipitated the gum. A comparative experiment showed me that the precipitated
gum obstinately retained about a quarter of its weight of sugar. The results that I have given were
corrected according to this principle.

[p. 256] I repeated some of these experiments on peppermint, Scotch pine, and
common juniper. The results were generally the same. The salts that were absorbed
the most by Bidens and Polygonum also were by these other plants. There were
differences in the absolute weights of salts absorbed. Differences are to be expected
for plants that are dissimilar, but plants of the same species present, in this regard,
such frequent anomalies that I could not attribute these differences with certainty to
the kind of plant that I was testing.
The excision of roots, their decomposition, and a general languor of growth favor
the introduction of solutes from the water into the plant.
I let plants whose roots I had cut absorb solutions. They then carried out the
above-mentioned [differential] secretions in a less marked way. They almost indis-
criminately absorbed all the salts that I presented to them. They absorbed all of them
in large amounts and nearly in the same proportion as the water of solution. Gum,
calcium acetate, and copper sulfate killed the plants in 7 or 8 hours, [p. 257] and I was
able to complete the trial in the other solutions only by repeatedly replacing the wilt-
ing plants with new ones. Thus, root-bearing plants absorb, from the same solution,
certain substances in preference to others. For example, they always absorb more
sodium chloride and potassium chloride than calcium acetate and calcium nitrate.
From a solution of sugar and gum, they take more sugar than gum, etc. These sub-
stances do not all enter the plant in proportion to their influence on growth. They are
absorbed in a much smaller proportion than is the water in which they are dissolved.
I am inclined to think that the plant, in absorbing one substance rather than
another from the same liquid, does not do this by virtue of a kind of affinity, but
98 8 Of the Absorption of Solutions by Plant Roots

rather based on the degree of fluidity or viscosity of the different substances [an
erroneous interpretation]. In fact, we find that distilled water is more fluid, or passes
more easily or rapidly through a filter, than does water in which chloride [p. 258] or
sulfate of sodium [“muriate or sulfate of soda”] is dissolved.11 We find that calcium
acetate and calcium nitrate form, with the same amount of water, more viscous
solutions, which pass less easily through a filter than do the chlorides and alkaline
sulfates. These last were always absorbed more abundantly by the plant than were
the first. Gum, which is more viscous than sugar, was absorbed in a smaller amount.
But we must assume that roots are exceedingly fine filters, much more perfect than
the filters that we commonly make. For if a solution containing calcium acetate and
potassium chloride is poured on a filter made of several double layers of paper, and
this filtration is interrupted as soon as half the liquid has drained through, [p. 259]
this half is not found to contain less calcium acetate than the half that remains on the
filter. The same is true for solutions of sugar and of gum.
The roots did not noticeably decompose the salts that I let them absorb. I never
found that an acid or an alkali had become free in the residue of the root suction.
Through incineration, I convinced myself that the salts had actually penetrated
the plant substance. In distilled water, I grew some root-bearing Polygonum plants
weighing, in the green state, 173.13 grams (3,262 grains). I grew other Polygonum
plants, of exactly the same weight, in 1.58 liters of water in which 3 grams (56-
1/2 grains) of potassium chloride were dissolved. As the plants used up the solution,
I replaced it with distilled water. The plants fared as well in the solution as they did
in the pure water. After 3 weeks I ended the experiment, and I found, by analyzing
the residue of the suction after drying it at a red heat, that the plants had removed
1.59 grams (30 grains) of potassium chloride.
After drying, [p. 260] the plants weighed 39.8 grams (750 grains), and they
produced, by combustion, 4.246 grams (80 grains) of ash.
The Polygonum plants grown in distilled water weighed, after drying,
38.851 grams (733 grains) [the author made an error in conversion: he wrote
132 grains; since the context strongly suggests that the “38.851 grams” figure is
correct, the conversion should be 733 grains] and produced 2.76 grams (52 grains)
of ash [another conversion error; the author wrote 151 grains instead of 52].
The plants thus increased their ash by an amount very nearly equal to that of
the potassium chloride [“muriate of potash”] that disappeared from the solution. I
analyzed these ashes and found, by silver nitrate, all the hydrochloric acid contained
in the added salts. The potassium chloride had not been decomposed. The absolute
amount of free potash was no greater in the ash of the Polygonum that had absorbed
pure water than in the ash of the Polygonum that had absorbed potassium chloride.
[De Saussure probably meant to say this the other way around: that the absolute
amount of free potash was no greater in the ash of the Polygonum that had absorbed

11
For these effects to be detectable, the solutions must be more concentrated than the ones that I let
plants absorb. It seems likely to me that a dilute aqueous saline solution is not always a homogeneous
combination, but a mixture of saturated and unsaturated water. It is well known that a dilute aqueous
solution of sodium chloride that is left completely at rest becomes denser and saltier at the bottom of
the vessel than at its surface. (See the memoir of Leblanc on Cristallotechnie. Journal de Physique,
year 11 [of the French Republican calendar; i.e., 1802].
8.4 Considerations on the Saline or Mineral Substances . . . 99

potassium chloride than in the ash of the Polygonum that had absorbed pure water.]
This experiment, repeated on Bidens, yielded very similar results. It is clear that
the same [kinds of ] plants may contain very different amounts of salts and that we
cannot rely too closely on constructed tables for assigning to each [kind of ] plant a
certain proportion of ash or salts. These determinations [p. 261] can be exact only
insofar as the plants grew in the same soil, for the same time.

8.4 Considerations on the Saline or Mineral Substances

that Enter into the Composition of Plants

Some authors have argued that the mineral substances found in plants are only there
accidentally and are not at all necessary to their existence, because the plants contain
them only in very small amounts. This opinion, undoubtedly true for substances that
are not always found in plants of the same kind, has not been demonstrated for those
substances that are invariably present. Their small amount does not indicate that they
are useless. The calcium phosphate [“phosphate of lime”] contained in an animal
perhaps composes not 1/500 of its weight, yet no one doubts that this salt is essential
to the constitution of its bones. I have found this same salt in the ash of all the plants in
which I have looked for it, and we have no reason to assert that they can live without it.
[p. 262] It has often been concluded, from the fact that some salts, in certain
proportions, are harmful to certain plants, that all salts, in all proportions, are harmful
to plant growth. But observation rarely confirms these systematic and general ideas.
Observation shows that many plants require a saline nutrient but that the nutrient must
be modified in its amount and characteristics, according to the nature of the plant
that must absorb it. Duhamel recognized that marine plants languish in a soil that
lacks sodium chloride [“muriate of soda”]. This salt is harmful to wheat [blé] in the
same proportions in which it is useful to marine plants. Pellitory, nettle, and borage
thrive only where they find calcium nitrate [“calcareous nitrate”] or potassium nitrate
[“nitrate of potash”]. Calcium sulfate [“sulfate of lime”] accelerates the development
of alfalfa, clover, and sainfoin. It has no effect on some other plants. Salts were
believed to favor plant growth only because they hastened the putrefaction of dead
plant matter spread on the soil or because they drew humidity from the air. But
calcium sulfate is not deliquescent, and if it were useful in accelerating putrefaction,
its salutary effect would not be limited to [p. 263] such a small number of plants.
Farmers use too little of it for it to produce the septic effects that have been attributed
to it or to be able to change the physical properties of the soil, considered simply as
a support for plants.
Thouvenel and Cornette believed they had found that saline solutions did not
enter plants and that the salts acted as astringents or corrosives, by closing the vessel
openings and resisting the absorption of water. Observation has shown, however,
up to now, that plants contain salts that occur in the soil in which they are growing.
Pellitory, nettle, etc., are rich in nitrate. Marine plants are rich in sea salt and no longer
yield it if they are forced to languish far from the seaside. De Bullion (Mémoires
d’Agriculture, 1791) sowed some seeds of sunflower (Helianthus annuus) in a sandy
100 8 Of the Absorption of Solutions by Plant Roots

soil lacking niter. The plants arising from the seeds yielded, at analysis, no trace
of this salt. He watered other, similar plants in this same soil with a solution of
potassium nitrate [“nitrate of potash”], and they became enriched in it.
Although experiment shows that certain [p. 264] saline substances are favorable
to the growth of some plants, it reveals at the same time that this nourishment suits
them only if provided in very small amount. Calcium sulfate probably owes its
salutary influence on the development of some plants partly to its low solubility. The
ineffectiveness of salts that are more soluble, such as potassium nitrate, potassium
carbonate, and sea salt, when used in their pure state and applied directly to the soil,
is recognized for most of the common plants. The usefulness of ash that contains
some of these salts is not disputed, but in the ash they are combined with earths
through a semi-vitrification that decreases their solubility and makes them penetrate
the plants slowly and in repeated doses, rather than all at once. In certain plants,
excess salts accumulate on the surface of the leaves and form there an incrustation
that kills them by impeding their transpiration. This, in large part, is the origin of the
white malady that attacks the Cucurbitaceae12 and many garden vegetable plants.
Plants do not obtain all of their mineral substances from saline solutions similar
to those that can be formed artificially by dissolving, for example, chlorides of lime,
iron or manganese in pure water, but they absorb them largely in combinations that
we cannot compose, such as ones in which the elements of these salts are chemically
united with oxygen, hydrogen, nitrogen, and carbon [p. 266] in humus extract, and
in which the salts can be detected only by incineration of the extract.

8.5 Application of the Foregoing Observations to Research on

the Amount of Nutrients that the Humus Substance Alone
Provides to Plant Roots

We are too far from knowing all the effects of the plant on humus, and of humus
on the plant, to be able to calculate and weigh all the elements that the plant takes
from humus. In the study of nature, we can do no better than to follow this maxim13
of the famous Haüy: Things are supposed to be such in themselves as they present

12
In gourd plants, this malady begins with viscous beads that appear especially on the upper surface
of the leaf, near the petiole. These beads dry and form prominent, powdery white blotches that spread
and multiply successively, out to the perimeter of the leaf. I detached such an incrustation. It was
only partially soluble in water and in alcohol. These solutions, evaporated to dryness, yielded a
deliquescent salt that had all the properties of calcium chloride combined with a negligible amount of
magnesia. The salt was precipitated in abundance by silver nitrate, potassium oxalate, and alkaline
carbonates, but not at all by barium hydroxide fully dissolved in water [“water of baryta”], and it
was almost unalterable by fire. The saline and earthy part of the incrustation made up about a third of
its weight. It was enveloped by a white plant substance insoluble in water and alcohol, and abundant
enough that the incrustation itself did not noticeably draw humidity. This malady especially attacks
old plants growing in soil that is very rich in animal manure, and in hot-beds where the leaves are
not washed by rainwater.
13
Traité de Minéralogie, by Haüy, Vol. 1, p. 7.
8.5 Application of the Foregoing Observations to Research . . . 101

themselves to our observation. If we rely on the observations made either in this

chapter or in previous ones, we see that humus extract, carbon dioxide gas, oxygen
gas, and all the water-soluble compounds that enter from the soil into a green plant
through its [p. 267] roots are not nearly sufficient to make up the greater part of the
weight of this plant, considered in the dry state.14
In Sect. 5.2, we saw that rain water, after having remained for several days in
well manured garden soil, formed an infusion that contained 1 part of dry extract
in 1,000 parts of water. We saw that a plant that absorbed this infusion would take
only a quarter of the solid extract that it contained. If the humus extract were its
sole nourishment, this plant would then gain only 1/4 pound of dry weight [p. 268]
in absorbing 1,000 pounds of infusion. An annual plant, such as a sunflower, that
grew in this garden would be able to acquire, in the 4 months after it germinated,
a weight of 4 kilograms (8 pounds) in the fresh state, or 1/2 kilogram (1 pound)
in the dry state.15 If we accept the experiments of Hales showing that the amount
of water taken up and transpired in 24 hours is equal to half the fresh weight of
the sunflower, then, according to my weighings of the sunflower at different stages
of its growth, it could not have transpired and absorbed more than 100 kilograms
(200 pounds) of water or infusion in 4 months. The amount of dry extract contained
in these 100 kilograms is 100 grams, or 1/5 pound. The plant took only one-quarter,
or 25 grams, of it, but the carbon dioxide contained in 100 kilograms of infusion must
be added to it. [Here de Saussure is again saying that some carbon dioxide enters
plants by way of the roots.] I estimate that this gas weighs 3.7 grams (70 grains),
based on the experiments [p. 269] reported in Sect. 5.2. Only half of the weight
of this gas was assimilated by the sunflower, which, in decomposing it, exhaled a
part of its constituents. This plant thus drew from the humus, disregarding the water,
25 + 1.85 grams of plant substance, which makes up only about 1/20 of the dry
weight that the sunflower gained, based on direct observation.
The calculation that I have just made is undoubtedly far from exact, but, supposing
that my estimate of the amount of nutrients that the plant draws through its roots from
the soil is two or three times too large or two or three times too small, the essential,
general results that I have in mind nevertheless remain the same. They show equally
that the humus extract, the gases, and all the water-soluble substances that come from

14
I could rely on the experiment of van Helmont, who, after having grown a willow for 5 years
in 200 pounds of humus, found that this plant had gained, in the green state, 164 pounds and that
the humus, which was dried in an oven before and after the experiment, had lost only 2 ounces.
But Kirwan made some very valid reflections on these results: He observed (1) that the humus
was placed in an unglazed, porous earthen vessel, which was itself buried in plant mold, and that
this last conveyed extractive juices to the plant; (2) that the oven drying might not have been the
same before and after the experiment; (3) that the willow left, in the soil, root fibrils whose weight
could not be estimated; (4) that the rain water that was used for watering the plant must, through
its impurity, have contributed to the plant’s nutrition. (Mémoire sur les Engrais, by Kirwan.)
15
Hales says that a sunflower in full growth loses only a quarter of its weight by drying. This result is
certainly incorrect, as are the inferences that this author deduced from it. [De Saussure misinterprets
Hales, who wrote, on p. 8 of Vegetable Staticks (1727), that the sunflower, upon drying, lost 3/4 of
its weight as water.]
102 8 Of the Absorption of Solutions by Plant Roots

the soil and enter the roots of a green plant do not compose the major part of the plant’s
dry weight, if the water itself is excepted. These [i.e., the water-soluble substances
from the soil] will always, however, be found to enter the plant appreciably and, to
have, as nutrients, a very powerful influence on growth, despite their small amount. It
is clear that the water [p. 270] acquired and solidified by the plant, whether from the
soil or from the atmosphere, makes up the greater part of the plant’s dry matter, by
weight; that the carbon is provided to the plant in a gaseous form, by the atmosphere
more abundantly than by any other source; but that the nitrogen, the salts, and the
earths, which are the least abundant elements in the plant, come from: (1) extractive
and saline solutions drawn by the roots from the humus; and (2) plant and animal
matter held in suspension in the atmosphere and deposited on the plant.

8.6 Summary

1. Plant roots absorb salts and extracts, but in smaller proportions than the water in
which these salts and extracts are dissolved.
2. The cutting away of roots, their decomposition, and sluggish plant growth in
general promote the introduction of salts and extracts into plants.
3. A plant does not absorb, in the same proportion, all the substances contained
simultaneously in the same solution. It makes particular secretions from it. In
general, it absorbs [p. 271] more of those substances whose separate solutions
are less viscous.
4. If the weight of the extract that the most fertile soil can produce is compared to
the weight of a dry plant that developed there, it is evident that the plant could
have drawn only a very small part of its substance from that source.
[p. 272] Chapter 9
Observations on the Ash of Plants

9.1 Observations Made by Various Authors on the Amounts

of Ash Produced by Plants

Experiment has long shown that plants of the same weight vary, by species, in the
amount of ash that they can produce. But the general reason for this phenomenon
has not been traced. The origin of this ash and the reason for the differences have
not been investigated. Before telling of my observations, I will briefly review what
is already known.
The inspector-commissioners of saltpeter manufacture in France, [p. 273] and
Kirwan and Rückert,1 found that, after drying, herbaceous plants yield more ash than
the same weight of woody plants. This result has been confirmed by all who have been
concerned with incineration products. Perthuis provided a beautiful demonstration
of this principle, when he reported that the trunks of trees produce less ash than
the branches, and the branches less than the leaves (Annales de Chimie, Vol. 19).
But we must stop there, regarding the results of this author. His other assertions,
(1) that plants that are burned when they are dry produce less ash than when they are
burned green; (2) that decayed wood produces less ash than healthy wood; (3) that
plants burned when mature produce more ash than when burned before or after
maturity; and (4) that plants generally yield more water-soluble salts the more ash
they contain, seemed to me to need to be modified. It is regrettable that, of the 60
incinerations carried out by Perthuis, only four were [p. 274] done on plants that were
dry and not mixed with unknown plants. The uncertainty left by incomplete drying,
estimated vaguely by fractions, such as one-half, one-quarter, and three-quarters,
limits the inductions that can be drawn from these experiments, especially if, as
must be suspected based on the results, the plants were not protected from the rain
after being cut. Despite these minor inaccuracies, which are inevitable in the course
of a lengthy work, we cannot disregard the usefulness of the research of this author,
whose object was not physiological observations but the large-scale extraction of
alkaline salts.

1
Mémoire sur les Engrais, by Kirwan, Société royale d’Irlande, Vol. 5, p. 129; and Rückert’s
Feld-Bau.

J. F. Hill, Chemical Research on Plant Growth, 103

DOI 10.1007/978-1-4614-4136-6_9, © Springer Science+Business Media New York 2013
104 9 Observations on the Ash of Plants

9.2 Principle by Which Ash Varies in Amount in Woody

and Herbaceous Plants

In a subject as new and complicated as the one that occupies me now, the explanations
that I give will, undoubtedly, very often be guesswork. But I have reason to believe
that the observations on which they are based are not so, at least for the species
[p. 275] that I examined, for although my incinerations were numerous, they are
perhaps still not sufficient for us to draw general conclusions.
I will consider only plants that are dry and of the same weight. The water of
vegetation varies so greatly according to the particular constitution of each individual
of a species, the age of the plant, and the climate, that it is advisable, above all, to
remove this source of error.
The research of the authors that I cited in the preceding section, and my own (as
can be seen in the table of incinerations (Table 9.1) at the end of this chapter), are
consistent in showing that woody plants contain less ash than do herbaceous plants.
If, as we saw in Chap. 8, plants allow earthy and saline substances into their interior
only in a liquid state, they should contain more ash the more abundant the suction
or transpiration is, for these two functions are always dependent, one on the other.
Hales2 and Bonnet3 demonstrated [p. 276] that herbaceous plants transpire more
water than do woody plants. The latter should therefore contain less ash.
According to the observation of Hales, the leaves of evergreen trees transpire
less than do those of deciduous trees. The latter are, for this reason, richer in ash.
Compare, at the end of the chapter, the incinerations of plants burned in the dry state,
nos. 1, 2, 16, 31, with nos. 67, 71, 74.
The leaves of evergreen trees transpire year-round, it is true, but in winter this
effect is very slight or almost nonexistent, and at this season they probably lose, due
to washing by rain water, as much ash as they acquire.
When a saline solution is contained in a vessel that allows evaporation through its
pores, more salt is deposited on the parts where evaporation is most abundant. The
distribution of ash in a plant generally follows this principle. Transpiration being
less abundant through the stem than through the leaves, the leaves are richer in ash.
Compare incineration no. 29 with the following one; no. 67 with no. 69; and no. 70
or 71 with the following one [this last should be no. 71 with no. 73].
[p. 277] Leaves, according to the observation of Hales, transpire more than do
fruits. Fruits yield much less ash. Compare, in the incinerations, nos. 30, 31, or 32
with no. 34; nos. 51, 52, or 53 with no. 54 [this last presumably should be no. 55,
not no. 54]; and nos. 57, 58, or 59 with no. 61.
The bark is the immediate site of the transpiration of the trunk and contains much
more ash than the inner parts. Compare, in the incinerations, nos. 5 and 6 with no. 7;
see nos. 14 and 15, nos. 20 and 21, nos. 22, 23, 24, and nos. 26, 27, 28.

2
Statique des végétaux, French edition, pp. 3 and 43.
3
Recherches sur l’usage des feuilles, p. 77, 8◦ edit.
9.3 General Composition of Ash; Influence of Soil 105

Ash steadily increases in tree leaves from the moment they emerge from the bud
until they turn yellow and fall off. See nos. 1 and 2, nos. 12 and 13, nos. 16, 18,
and 19, and nos. 30, 31, 32. Plants should accumulate ash in all the parts that do
not change their form or growth force. At all times, I chose leaves that were entirely
green and healthy. In summer and autumn, however, I took care to collect only those
that appeared the oldest.
The results that I have just noted [p. 278] cannot be applied to annual plants
collected and burned in their entirety, with any dead parts that they may have. The ash
of these plants diminishes as their growth advances. See, in the table of incinerations
(Table 9.1), nos. 35 and 36, nos. 37, 38, 39, nos. 43, 44, 45, nos. 46, 47, 48, nos. 51,
52, 53, and nos. 57, 58, 59. The reason for this difference is that annual plants, as
they age or produce new growth, lose their lower leaves, which are the oldest and
consequently the most laden with ash. If these leaves do not drop off, they languish
or dry, and in this withered state they give up their soluble substances to rain water,
dew, and even to their transpiration. If we could monitor the proportion of ash in a
single leaf of an annual plant before the leaf deteriorated, we would probably see an
increase, as in the leaves of the trees that I cited.
Ash appears to increase in annual plants as they age, if the plants are considered in
the green state. See nos. 37 and 38, and nos. 47 and 48. But this is [p. 279] an illusion
produced by the water of vegetation. The greater loss of this water with advancing
growth appears to increase the proportion of ash in the plant as the plant approaches
the dry state.
Ash does not accumulate indefinitely in tree trunks. The sapwood contains more
ash than the wood. See, in the table of incinerations (Table 9.1), nos. 5 and 6, nos. 22
and 23, and nos. 26 and 27. When the layers of sapwood harden and become wood,
they lose, to the ascending juices, the ash that they had accumulated during their
growth.
I took fresh hazelnut leaves, dried some of them and washed the rest several times
in cold, distilled water. The latter were dried as the former were. One hundred parts,
by weight, of the washed leaves yielded less ash than 100 parts of the unwashed
leaves. This result may serve, in part, as support for the preceding interpretation.
A putrefied plant yields, at the same weight, more ash than the same plant unpu-
trefied. One hundred parts, by weight, of rotten wood provided me more ash than
did 100 parts of healthy wood. But this result presumes, in some cases, [p. 280] that
the plant has not been washed by running water during putrefaction.

9.3 General Composition of Ash; Influence of Soil

The alkaline salts of potassium [“potasse”] or sodium [“soude”], the earthy phos-
phates of lime or magnesia, free or carbonated lime, silica, and the oxides of iron
and manganese make up, together or separately, the most notable components of the
ash and the only ones with which I will be concerned. Ash contains many others that
we do not usually notice, due to their small amount. Ash holds perhaps all the known
substances that cannot be volatilized by the effects of the fire used for combustion.
106 9 Observations on the Ash of Plants

For it is possible that our atmosphere holds in suspension all the elements, and that
a thorough analysis would reveal traces of them in all the soils.
Analysis shows that all the dominant substances in the ash are contained in humus,
and that its soluble part, which alone penetrates the plant, contains these substances
in greater proportion than does [p. 281] the insoluble part. (See the table of analyses
(Table 9.1), nos. 10 and 11, nos. 76 and 77, and nos. 78 and 79.) [The pair 78/79
does not illustrate his point, because the table shows much more ash in the humus
than in its extract. He notes, in the table, that the large amount of ash in this humus
is due to sand and detritus of gneiss mixed with it.] Their presence in the plant is
therefore only natural; their absence would be more surprising.
The nature of the soil has, other things being equal, an appreciable influence
on variations in the amount of ash in most plants. I gave (Chap. 8) details of an
experiment in which I grew Polygonum persicaria with a saline solution, and other,
similar plants with distilled water. The plants in the saline solution yielded, at equal
weight, almost twice as much ash as the ones in distilled water. I allowed fava seeds
to develop by three different methods. The first ones were protected from the rain
and nourished with distilled water. One hundred parts, dry, of the plants that these
seeds gave rise to contained, at flowering time, 3.9 parts of ash. Other, similar seeds
were sown in glass capsules filled with gravel and placed on the ground under the
open sky. They were watered, both naturally and artificially, with rain water. One
hundred parts, dry, of these plants yielded, at flowering, 7-1/2 parts of ash. [p. 282]
Finally, fava plants in flower, grown in open ground in a vegetable garden, yielded
12 parts of ash.
The proportion of the elements in the ash is almost always related to that of the
elements composing the soil. Plants grown in a humus originating from a siliceous
mountain yield, other things being equal, ash that contains less lime and more silica
than plants grown in a carcareous humus. (See, in the table of analyses (Table 9.1),
nos. 67 and 68, nos. 71 and 72, nos. 74 and 75.) The soil of le Bréven [modern
spelling “Brévent”], the granitic mountain [in the Alps] on which I made the siliceous
harvests, was much richer in iron oxide than was the soil of the calcareous mountain
[presumably la Salle, in the Jura; de Saussure (1800) was an analysis of the minerals
in plants of various mountain soils]. The same difference was seen in the ash. All
the plants were harvested at the same season, but growth was a little retarded in
the siliceous soil, and this circumstance tended, as I will show in the following, to
decrease the proportion of silica and iron oxide in the ash of the plants that grew there.
The differences between the productions of calcareous and siliceous soils are
detectable only insofar as the extractive substances that [p. 283] nourish the plants
contain different amounts of silica and lime. Plants grown in a calcareous sand and in
a granitic sand, amended by the same manure or by the same humus, would contain
similar ash. One may judge by the experiment of Lampadius, recorded in the Journal
des Mines, no. 65 [actually, no. 55; see Appendix 3, Bibliography of References Cited
by de Saussure (as expanded by the translator)]. This author prepared five garden
beds, each 4 [square] feet in surface area and 1 foot deep. They were separated by
boards. Each square was filled with a very pure earth and 8 pounds of cow manure.
These earths were alumina, silica, lime, magnesia, and garden humus. Rye was sown
9.4 Of the Alkaline Salts in Ash 107

there, and its ash was found to contain the same substances in the five different earths.
The author concludes from this that silica forms during plant growth, regardless of the
nature of the soil in which the plant is grown. But as Lampadius did not analyze the ash
of the cow manure, which alone nourished the rye, we cannot draw this conclusion.
The only certain conclusion to which this experiment can lead is that earths, if not
in a soluble state, do not penetrate plants. [p. 284] Furthermore, I will show in the
following that the open atmosphere introduces a small amount of ash into plants.
I will now investigate why plants of different species, growing in the same soil,
do not contain all their ash constituents in the same proportion, and why this ash
also varies in the different parts of the same plant. The explanations that I can give
on this subject are sometimes far from fully satisfactory. They often depend on a
knowledge that I lack, namely, that of plant structure. But they are less absurd than
those that ascribe to plants a creative force for all of its elements, and, if my reflections
are incorrect, I hope at least to draw attention to new observations. I will examine
separately the principal ash constituents.

9.4 Of the Alkaline Salts in Ash

First Observation The alkaline salts4 are, without comparison, the most abun-
dant constituent [p. 285] in the ash of an herbaceous green plant, all of whose
parts are growing. I was very surprised to find that the ash of young goldenrod
plants, no. 43, grown in a barren, uncultivated terrain, and the ash of fava no. 37,
sunflower no. 46, wheat [“froment”] no. 51, and maize [Zea mays] no. 57, con-
tained at least three-quarters of their weight as alkaline salts. [The entries in the
water-soluble-salts column are actually only 55–69 %, but adding in the major
part of the “deficit” (see discussion in Sect. 9.11 G), which is listed in a sepa-
rate column, they mostly reach three-quarters or more.] We can say almost the
same about the ash of tree leaves emerging from their buds: At least one-half and
sometimes three-quarters of their weight consists of these salts. I found one-half
in the ash of leaves of hazelnut and poplar trees on 26 May; and three-quarters in
oak leaves on 10 May, before their full development. And all these trees grew in
a soil that was uncultivated and of poor quality. This result should not be surpris-
ing, since we have seen that the ash of an extract of an uncultivated humus yielded
at least half its weight as alkaline salts. (See Sect. 5.3, and the table of analyses
(Table 9.1), nos. 76 and following.) [He probably means, instead of no. 76, no. 77,
with the alkaline salts that are included in the “more precise analysis” and the
“deficit” columns added in.] [p. 286] But even if this product composed much less
of the extract, there could be much more in the plant, if, as we must presume,
the alkaline salts are the finest, least viscous of all the fixed parts of the extract.

4
I adopt here the oldest accepted meaning, which confers the name of alkaline salts only on those
in which potash, soda, or ammonia is one of the constituents. We must exclude only ammonia,
because it cannot enter into the composition of ash.
108 9 Observations on the Ash of Plants

I showed in Chap. 8 that the different substances that could be contained in the same
solution did not enter the plant in the same proportion, but in inverse relation to their
viscosity.
Second Observation The proportion of alkaline salts never increases appreciably,
and most often decreases, as the plant develops and ages in the same soil. This
observation applies not only to annual plants, but also to tree leaves during the course
of a season. A plant emerging from the ground, a leaf emerging from its bud, contains
ash that is as rich, and usually richer, in alkaline salts at this time than at any other
stage of its growth. See, in the table of analyses, nos. 1 and following; nos. 12 and
following; nos. 16, 18 and following; nos. 30 and following; nos. 35 and following;
nos. 43 and following; 46 and following; 51 and following; 57 and following. The
more advanced the growth, the greater is this removal of saline substances. [p. 287]
A plant emerging from the ground in May loses less salt between May and July than
between July and September.
When a fresh plant is washed with water (table of analyses, nos. 16 and 17), the
liquid removes the alkaline salts in greater proportion than all the other constituents
of the ash. The liquid removes less of the salts, or the plant retains them more
tenaciously, the greater the plant’s growth force or the less advanced its development.
The water that circulates around the roots, the rain water that falls on the leaves, and
indeed the water that the leaves transpire carry away the salts, as I have just said, and
in greater proportion the more water the plant receives.
Third Observation The ash of bark contains a much smaller proportion of alkaline
salts than does the ash of wood or sapwood. See no. 4, no. 7, no. 15, no. 21, no. 24,
no. 28. [These are all entries for barks, which are to be compared to entries for wood
or sapwood listed nearby in the table.] The bark renews itself only very slowly. It is
exposed year-round to washing by rain and dew. It is provided with a dead substance,
the epidermis or cork. More than any other part, it must be depleted of its [p. 288]
most soluble salts. It does not seem, however, that this effect can be attributed solely
to the causes I have indicated.
Fourth Observation The ash of fully formed wood is almost as rich in alkaline
salts as the ash of the sapwood that is adherent to it. See, in the table of analyses,
nos. 5 and 6, nos. 22 and 23, nos. 27 and 28. [This last should be 26/27; also, pairs 5/6
and 26/27 show the fully formed wood having slightly more alkaline salts than the
sapwood, even after the deficit is added back in.]
This result is peculiar. It is contrary to the reduction of ash in the wood and to the
other components of this same ash. [But the table shows, for 5/6, 22/23, and 26/27,
that some ash components are less abundant, some more abundant, in the sapwood
than in the fully formed wood.] The well formed woods in healthy, vigorous trees
such as the woods that I burned are not a dead substance. They serve as channels
or conduits for the sap, and it is likely that these channels, which are much nar-
rower or denser than those of the sapwood, admit only the most fluid juices, such as
those carrying saline substances, and that these same channels are an insurmountable
obstacle for the other substances. Moreover, the wood is far removed from atmo-
spheric influences, and loses through these influences almost none of the salts that it
acquires.
9.4 Of the Alkaline Salts in Ash 109

It seems to me that, other things being equal, the harder a wood is, the richer it
should be in saline substances, [p. 289] because the ducts of hard wood are narrower
and because all their substance allows less rain water to penetrate. But the slow
growth rate of trees having hard wood undoubtedly requires frequent modifications
of this principle. If soft woods are more quickly depleted of the salts that they acquire,
they receive, on the other hand, a greater quantity of salts in the same time due to
their rapid absorption and growth.
Fifth Observation The ash of seeds is richer in alkaline salts than is the ash of the
plant that bears them; see, in the table of analyses, nos. 29, 32, 34 [compare these
three as a group], 40, 41 [these are all right as a pair, if the “more precise analysis” in
the far right column is taken into consideration], 54, 55 [these are all right as a pair],
60, 61 [no. 61 shows no value for any substance; he probably means no. 62 instead,
but the total for alkaline salts in these seeds is less than in the stem, no. 60, whereas it
should be more], 63, and 64 [these are all right as a pair if the deficit and “more pre-
cise analysis” are taken into consideration]. Most seeds are enclosed in integuments
of very low porosity, which protect them from atmospheric influences. They transpire
little and should therefore retain their salts. Moreover, they are attached to the rest of
the plant only by very fine, very slender channels, which admit only the most fluid
juices. The famous Vauquelin made an observation contrary to the one I just reported.
He observed (Annales de Chimie, Vol. 29), in comparing the ash of oat grains with
the ash of a stack of grain-bearing oat plants [p. 290] accidentally burned at Ecouen
[a canton on the outskirts of Paris], that the seed alone contained no alkaline salts,
but that the whole plant did contain them. I will note, in general: (1) that the time of
harvest introduces such large variations in the composition of the ash that the differ-
ent parts of the same [kind of ] plant cannot be compared unless they were collected
at the same time and from the same soil; (2) that this chemist did not burn pure oat
seed, but oats with its chaff, for I obtained, as he did, 3.1 parts of ash in 100 parts
of unhulled oats, but I found only 1.7 parts of ash in 100 parts of oats stripped of its
chaff or calyx. The chaff composes about one-third of the weight of unhulled seed,
and 100 parts of this envelope contain about three times more ash than do 100 parts
of naked seed. Thus, in the ash that Vauquelin examined, more than three-quarters
of the weight was associated with the chaff.
I analyzed the ash of oat seeds bearing this envelope. This ash yielded almost no
alkaline salts by a purely aqueous leaching, but when I dissolved the ash [p. 291] in
nitric acid, separated the silica by a filter and the earthy phosphates by ammonia, and
then subjected the remaining liquor to a heat-driven evaporation pushed by degrees
to the highest temperatures, I obtained, as a residue, 15 parts of alkaline salts in
100 parts of ash of oats provided with its chaff.
The difficulty of obtaining enough naked oat seed, without spoilage, prevented
me from making a precise analysis of its ash compared with that of its straw, but I
have reason to believe that the results would confirm those that I obtained with other
seeds, and would show that the ash of naked oat seed contains more than half its
weight as alkaline salts. I have devoted more time to this discussion than it needs. The
results of Vauquelin are no doubt completely accurate, but the timing of the harvests,
or even the exposure of the seed to rain, are sufficient to explain the differences in
our analyses.
110 9 Observations on the Ash of Plants

The ash of seeds contains very nearly the same amount of alkaline salts as the
ash of the plant that arises from them, in the same soil, at the time when the plant
yields the most, that is, in the first stages of growth. [p. 292] I obtained only very
slight differences in this regard, and these are largely attributable to the fact that my
analyses were made on entire seeds. Their whole substance is not used as nutriment
by plants. Only the inner part of the seed is used for this function. Now, I found that
the bran, or the outer part of the wheat grain, contained a somewhat smaller amount
of alkaline salts than the grain itself, and this bran was mixed with a certain amount
of meal.
I went into great detail about the distribution of alkaline salts in plants because
the distribution of the other solid substances in plants is almost entirely subordinate
to it.

9.5 Of Earthy Phosphates in Ash

Earthy phosphates, namely, those of lime and magnesia, form a water-soluble com-
bination in humus extracts, as only nature can produce. We do not know whether
these salts are essential to the extract. Be that as it may, when I speak of the solubility
of these phosphates in water, I mean [p. 293] only the solubility of the extractive
part that contains them. It is the same with silica, lime, and the metallic oxides.
The earthy phosphates are, after the alkaline salts, the most abundant constituent
of the ash of an herbaceous green plant all of whose parts are growing and developing.
We must presume that, after the alkaline salts, they are the finest, least viscous part
of the solid substance of the extract. These two kinds of salts almost always follow,
except for their amount, and by the same causes, the same course in the ash.
When a plant is washed with water, the earthy phosphates are removed by this
liquid in greater proportion than all the other constituents of the ash except for the
alkaline salts. See, in the table of analyses, nos. 16 and 17.
A tree leaf contains ash that is richer in earthy phosphates when it emerges from
its bud than at any later stage of growth. See, in the table of analyses, nos. 1, 2,
etc. [The pair 1/2 shows this pattern, but the immediately following ones are not
relevant.]
The proportion of earthy phosphates decreases in the ash of annual plants from
the time of germination up to [p. 294] that of flowering. See nos. 37, 38, 43, 44, 46,
47, 51 and 52. But these phosphates seem to increase at the time of seed maturation,
if the seeds are abundant. (See the numbers that follow all those I just cited.) This
reverse course is an illusion produced by the effect of a change of substances that
occurs during the process of incineration. The phosphates are decreased in the plant
and increased in its ash. The ash of seeds is abundantly provided with phosphate of
potash and lacking in carbonate of lime. The ash of the stems and leaves, at the time
of fruit maturation, contains little or no phosphate of potash, and, on the other hand,
some carbonate of lime. See analyses nos. 40, 41, 54, 55, 60, 62, 63 and 64. When
the ashes of the stems and seeds are mixed during incineration, the free or carbonated
lime decomposes the phosphate of potash, forming earthy phosphate. It changes an
alkaline salt into an earthy salt, and thus makes the earthy salt seem more abundant
9.6 Of Free or Carbonated Lime in Ash 111

in the ash than it is in the plant. This result is very striking in the analyses of the ashes
of fava plants having mature seeds; of these plants [p. 295] without their seeds; and
of the seeds themselves: nos. 39, 40, and 41.
The amount of earthy phosphate that the seed alone contains in greater proportion
than the stem is not sufficient to explain the increase in earthy phosphate obtained by
the incineration of the entire, seed-bearing plant, without adding the earthy phosphate
that comes partly from the decomposition of the phosphate of potash contained in
these same seeds.
The ash of bark contains a much smaller proportion of earthy phosphate than does
the ash of sapwood. See, in the table of analyses, nos. 6, 7, 14, 15, 20, 21, 23, 24, 27
and 28. The reason for this is the same as for the alkaline salts (Sect. 9.4).
The ash of the sapwood contains more earthy phosphate than does the ash of the
wood. This result is consistent with the reduction of ash in the wood (see the table of
incinerations), but is contrary to the presence of alkaline salts in this same wood. I
tried (Sect. 9.4) to give an explanation for this anomaly. I do not consider the stated
reason as fully satisfactory, however. It is possible that the parts that stop growing
lose, [p. 296] or allow the decomposition of, their phosphoric acid and retain only
the phosphate earth.
The discovery of a large amount of phosphorus in seeds is very old. It dates to
the time of Pott, and Marggraf confirmed it.5 Vauquelin recognized that the stems
of wheat and oats contained less phosphate than did the seeds. These results were
confirmed by my observations on several other, very different, plants. This famous
chemist, jointly with his colleague Fourcroy, withdrew magnesium ammonium phos-
phate [“magnesian ammoniacal phosphate”] from some seeds before burning them.
The action of potash on this phosphate probably led to the formation of the phos-
phate of potash that my analyses revealed so abundantly in several ashes. Phosphate
of potash is surely also produced by the action of potash on simple phosphates of
lime and magnesia, as will be seen in detail in Sect. 9.11.1.

9.6 Of Free or Carbonated Lime in Ash [p. 297]

When I washed leaves with water (table of analyses, nos. 16 and 17), the proportion
of carbonated lime in their ash increased. This result occurred only because of a
larger withdrawal of the other fixed substances, namely, the alkaline salts and the
phosphates. For this reason, it is understandable why, in all my analyses nos. 1, 2,
12, 13, 14, 16, 18, 35, 36, etc. [no. 14 does not seem to belong in this list], I found
that the proportion of carbonate of lime increased as the plant grew. It is because the
plant lost its salts and phosphates in much greater measure than the lime.
We only rarely find plants that, through a particular disposition, retain their salts
in very nearly the same proportion during their growth. In these plants, carbonate of
lime does not increase appreciably. This is the case in, for example, fava plants.
In herbaceous green plants all of whose parts are [p. 298] growing, carbonate of
lime is present only in very small amounts. These plants contain barely more than
ten or twelve hundredths of it, and I believe that ash composed of nearly equal parts

5
Opuscules chimiques of Marggraf, Vol. 1, p. 68.
112 9 Observations on the Ash of Plants

carbonate of lime and alkaline salts would be an impossible product for one of these
plants, at least on a natural site. This is because the alkaline salts that are present in
a green plant are composed largely of alkaline phosphates, which are changed into
earthy phosphates by incineration.
We will therefore not conclude from the fact that certain plants (such as the grasses
before they are fully developed) yield no free or carbonated lime through incineration
that the substance that yields this earth in other plants is not present in those with
which we are concerned. It is very likely that it is.
The ash of barks contains a large amount of carbonate of lime, much more than the
sapwood, because barks are depleted of their alkaline salts and earthy phosphates.
See nos. 6, 7, 14, 15, 23, 24, 27 and 28.
The ash of well formed woods contains [p. 299] more carbonate of lime than does
the ash of the sapwood because the formed woods are depleted of their phosphates.
The ash of most seeds, and even of all those that I examined, contains no carbonate
of lime because the seeds are rich in phosphate of potash. It is possible that, due to
the presence of this salt, plants that are heavily laden with seeds yield ash that is
richer in carbonate of lime at the time of flowering than at fruit maturity.
The seeds of some Lithospermum produce a slight effervescence with acids. They
contain carbonate of lime or magnesia before combustion, but it is possible that their
ash does not produce these earths in a free or carbonated form.

9.7 Of Silica in Ash

Silica is never present in large amounts in ash unless the plants are depleted of
their salts and phosphates. Large amonts of alkaline salts and of [p. 300] silica are
incompatible in the composition of the ash of a young, herbaceous green plant, all
of whose parts are growing.
When a fresh plant is washed with water, the proportion of silica in the ash
increases. See table of analyses, nos. 16 and 17.
The ash of young plants, and of leaves emerging from their buds, contains very
little silica. But the proportion of this earth increases as the plant develops and
is depleted of its alkaline salts. The purest plant molds, exposed to atmospheric
influences, are, for this reason, very rich in this earth.
The proportion of silica does not increase appreciably in plants, such as fava, that
retain the same proportion of alkaline salts throughout their growth.
I sowed seeds of maize [Zea mays] and wheat [“froment”: Triticum sativum]
at the same time, in the same soil. When I examined these plants a month after
germination, or a month before flowering, I found that all the visible parts of the
maize were growing. Its ash [p. 301] then contained 8/100 silica and 80/100 alkaline
salts. The radical leaves of the wheat at this time, however, had already become dry
or yellow, although the plant was thriving. Its ash then contained 12/100 silica and
65/100 alkaline salts. At flowering time, or a month after the preceding observations,
all parts of the maize were growing as before and it still yielded ash containing 8/100
silica and 80/100 alkaline salts. But the wheat, although vigorous, had an increased
number of dried or yellowed leaves. Its ash then contained 32/100 silica and 54/100
alkaline salts. These observations show how, through very basic circumstances that
9.9 Influence of the Atmosphere on the Ash of Plants 113

vary according to the growth of the different plants, they must contain dissimilar
ashes, even when the plants are assumed to absorb identical nourishment. But it is
very likely that the nourishment is not the same in the same soil, and that the roots
modify it when it enters, based on the width of their pore openings.
Most grasses are distinguished from other plants by a larger silica content. We
may conclude from this that [p. 302] these grasses take up and lose much more. We
must believe that they absorb a more abundant nourishment and that they deplete
themselves of part of it. The plants that are richest in silica should, other things being
equal, be the most exhausting [of the soil].
I will not say at present that these grasses differ by having root pores that are
more open. It is possible, on the contrary, that their pores are narrower, since these
plants, when young, contain less carbonate of lime and often more alkaline salts than
other plants. It must be acknowledged only that they have a much greater suction
force and much more abundant excretions. Indeed we see in many of these plants
an appreciable, very unusual, transpiration (Physiologie végétale of Senebier, Vol. 4,
p. 87).
Seeds lacking their external coats contain less silica than the leafy stem bearing
them.
I do not pretend, however, to be able to explain everything. In the distribution of
silica in trees, there are effects whose cause I do not understand. The trunks of trees
[p. 303], their bark, sapwood, and well formed wood often contain almost no silica,
whereas the leaves are rich in it, especially in autumn. It is by the periodic dropping
of leaves that trees cast off this earth. I examined barks of five different trees—those
of poplar, young and old oak, hazelnut tree, mulberry, and hornbeam—and I found
a substantial amount of silica only in the bark of mulberry. I found almost none in
the well formed wood of any these trees. Their leaves contained a notable amount,
four or five times more than the wood or bark.

9.8 Of Metallic Oxides in Ash

The proportion of oxides of iron and manganese in ash increases as growth advances.
Tree leaves yield ash that is richer in these substances in autumn than in springtime. It
is the same with annual plants. Seeds contain metals in smaller proportion than does
the stem that bears them. When a plant is washed with water, [p. 304] the proportion
of its metallic oxides increases. The purest humuses are always very rich in metallic
oxides, much more so than the plants that formed them.

9.9 Influence of the Atmosphere on the Ash of Plants

We can determine whether the atmosphere introduces earths and salts into plants by
germinating seeds with the aid of distilled water and seeing whether the weight of
the ash of the plants that are developed in this way exceeds the weight of the ash that
a preliminary experiment indicates in the seed.
Forty-one fava beans (Vicia faba), also known as marsh beans, weighing, together,
79.135 grams (2-1/2 ounces + 51 grains), were placed in the flared necks of 41 vials,
114 9 Observations on the Ash of Plants

each filled with 594 cubic centimeters (30 cubic inches) of distilled water. I exposed
them to the open air, in sunlight, on an exterior window sill protected from rain. These
plants reached a height of a foot and even more. [p. 305] Their stems were weak and
could not support themselves without several fastenings. They flowered, but these
flowers were small, imperfect, and almost unrecognizable. I ended the experiment
after 2-1/2 months, during or immediately after flowering, because at this stage the
plants had reached their greatest development and because their growth had become
weak. The leaves and stem tips collapsed under their own weight.6 They had absorbed
1.7835 decaliters (900 cubic inches) of distilled water. After the experiment, the vials
contained 3.963 liters (200 cubic inches) of water rich in conferva [a green alga].
The liquid, after evaporation, left a dry residue weighing 5.8 decigrams (11 grains),
which produced by combustion 1-1/2 grains of ash. This ash [p. 306] seemed to be
composed of 3 parts lime, 1 part silica, and an imponderable amount of alkaline salts.

Weight of fava seeds before the 79.135 grams (2-1/2 ounces +

experiment 51 grains)
Approximate weight of the green 611 (20 oz.)
plants that arose from the
preceding seeds
Dry weight of the same plants 84.337 (2-3/4 oz. + 5 gr.)
Weight of the ash of the fava seeds 2.601 (49 gr.)
before they developped
Weight of the ash of the plants that 3.025 (58 gr.)
arose from the seeds
Difference, or weight of the ash 424 milligrams (9 gr.)
deposited by the atmosphere
The 49 grains of ash of the seeds, The 58 grains of ash 58 grains of ash of
before the experiment, contained of the fava plants flowering fava
that developed in plants, grown in
distilled water plant mold,
contained contained
Potash 10.9 13 33.2
Potassium phosphate 21.5 19.25 0
[“phosphate of potash”]
Chlorides [“muriates”] 1.4 2.5 7
and alkaline sulfates
Earthy phosphates 13.68 17.5 8.7
Earthy carbonates 0 0 3
Silica 0 Imponderable amount 1.2
Metallic oxides 0.25 0.25 0.25

6
I would have been able to obtain much more vigorous growth, no different from that in open
ground, at least up to the fruiting stage, if I had sown these seeds in pots full of pure sand or gravel,
but I do not know whether the juices of the roots would have attacked the stones themselves. The
erosion that lichens sometimes seem to cause on rocks suggests this. Moreover, the prodigious
number of insects that, in open country, leave their remains on leaves, and the debris of all kinds
that the winds transport there, would cast doubt on the results of the experiment.
9.10 Details on the Procedures used for Incineration 115

[p. 307] From the foregoing, we see that the plants gained 424 milligrams
(9 grains) [conversion is a little off: 424 milligrams equal 8 Paris grains; 9 grains
equal 477 milligrams] of ash and that the composition of the ash of the entire plant
that developed in distilled water differs very little from the products of the seed.
The atmosphere, I believe, deposited on these plants all the common constituents
of ash, but principally lime. This earth decomposed a part of the phosphate of potash
and formed phosphate of lime. During growth, a small amount of alkali was evap-
orated. The difference between the ash of the fava plants that developed in distilled
water and the ash from plants grown in humus shows us how great is the influence
of the soil on their composition.
I do not believe that we can conclude, from the small gain that we have just seen
in plants grown in distilled water, that they themselves formed, with gases and water,
these additional substances. If we consider how quickly a body of any kind exposed
to the open air is covered with dust due to the immense number of corpuscles floating
in our atmosphere, and if we note that the 41 fava plants [p. 308] presented to these
corpuscles, for about 2 months, a stopping place of more than a square foot of surface
area, we should be less surprised that this gain occurred than if it had not occurred.
Its amount suggests nothing supernatural here. We note further that distilled water
itself is never pure. It contains foreign substances7 whose weight we cannot estimate
because they escape with the water during evaporation.

9.10 Details on the Procedures used for Incineration

Annual plants were collected whole, including any dead leaves that they might have.
I cut away only those roots [p. 309] that were still impregnated with soil. The lower
parts of the stem were carefully freed of soil splatterings. When I collected annual
plants having mature seeds, I did so at the precise moment of this maturity and not
after the death of the plant. Tree leaves, even those of autumn, were always chosen
green and in a perfectly healthy condition. Care was taken at this season, however,
not to take the leaves of the most recent shoots, but of the oldest.
All these plants were sheltered from the rain immediately after they were collected.
When they seemed dry, they were placed for several weeks in an oven heated to 20◦
Réaumur. Their green or soft parts mostly became friable and brittle there, and in
this dry state the plants were weighed. Despite this precaution, however, the plants
were not all brought to the same degree of dryness. The dryness that they can reach
at the same temperature varies not only according to their species, but also within
the same plant, depending on the stage of growth. It seemed to me that, in general,
green plants collected [p. 310] when young could be dried more completely than

7
Karsten and Westrumb noted that water that is distilled with the greatest care and, if recently
prepared, is not at all modified by reagents, undergoes, after exposure for a fortnight to light or
4 weeks to darkness, changes that have all the characteristics of fermentation products. It then
somewhat clouds solutions of lead and silver. It gives slight indications of ammonia and, at other
times, of acid. (Kleine Physikalisch-chemische Abhandlungen von Westrumb. Zweites Heft [spelling
of title corrected by translator]).
116 9 Observations on the Ash of Plants

plants harvested at a more advanced age. The latter often retain, at the temperature
indicated above, a semi-flexibility, which is a certain sign of the presence of water
in substances, such as those that I am concerned with, which become stiff or brittle
at a higher temperature. The impossibility of estimating the amount of this water,
liquid and external to the essential composition of the plant, must always put a little
uncertainty into the products of plant analysis.
The plants dried by this method were burned on a large iron slab. The residue of
this operation was incinerated anew in a crucible heated to a dark red, until no parts
of the charcoal were susceptible to burning. There are plant substances, such as the
straw and seed of wheat [Triticum sativum], that I could not incinerate completely
or reduce to gray or white ash. But sugar, starch, gum, maize plants and their seeds,
likewise those of fava, barley, and oats, could be reduced to white ash.
[p. 311] The more time devoted to the operation, and the gentler the heat of the
furnace, the more complete the incineration. Care must be taken, especially in the
always very difficult combustion of seeds, not to stir the ashes often. Such action, by
crushing the salts and charcoal, by condensing it while it is somewhat fluid, prevents
its combustion and sometimes makes it impossible. The yields of my incinerations
are, in general, lower than those of the investigators who preceded me. But most of
them, having had in mind only the large-scale extraction of salts, did not complete,
in a crucible, the incineration of the residue of the first combustion in the open air.
According to Perthuis, plants yield more ash if they are burned when green than
if they are burned when dry. But he probably reported this result so that plants would
not be exposed, after harvest, to the effects of water [presumably because water
would deplete them of salts, which Perthuis sought to maximize], for, green or dry,
they provided me the same amount of ash.
All the products of my incinerations were weighed hot, upon removal from the
crucible, and analyzed similarly.

9.11 Details on the Procedures used for Analyzing Ash [p. 312]

The goal I set myself was to understand the order in the composition of the ash,
not only in different plants, but also in their different parts and according to their
developmental stage. My lifetime would have been insufficient for this work had
I obliged myself to make very detailed analyses. The procedures that I followed
seemed precise enough for the general research that I had in mind. They consist of
the following operations:
A. The ash, dried at a red heat and pulverized if it was agglutinated, was boiled
with 20,000 times its weight of distilled water. The filtered lye was evaporated to
dryness at a heat up to red hot. The weight of its residue indicated to me the weight
of the water-soluble salts that the ash contained. It is this result that was recorded in
the first column of the table, when a new solution of this residue, in a small amount
of water, left no [p. 313] insoluble material. But, in the contrary case, the solution
was filtered and evaporated anew, until the salts were completely dissolved. The part
that became insoluble by evaporation is the carbonate of lime in ash, such as that of
barks, which contains a large amount of this earth. It is formed by a phosphate of
9.11 Details on the Procedures used for Analyzing Ash 117

potash and lime or magnesia [de Saussure’s term was “phosphate potassé de chaux
ou de magnésie”] in the ash of grasses and seeds that contain no free or carbonated
lime, but, on the other hand, a great deal of earthy phosphate and potash. Silica
occurs only in insignificant amounts, at least if one uses, as I always tried to do,
only the lowest degree of fire possible to produce incineration. My yields of alkaline
salts are, in general, much larger than those obtained from the same plants by my
predecessors. I can attribute this difference only to my more complete incinerations
and to the care that I took in drying my plants at harvest. The water is far from
removing all the alkaline salts from the ash, but the amount obtained is always very
nearly proportional to that which the ash contains, if there is no phosphate of potash.
[p. 314] When this salt was encountered in the ash, I almost always made a more
detailed analysis, which I will discuss in the following [see G, below].
B. The washed, water-insoluble ash was boiled with seven or eight times its weight
of rectified nitric acid in a porcelain capsule. The residue was treated, at a red heat,
with six times its weight of powdered soda in a platinum crucible. The [resulting]
glass [presumably water glass, sodium silicate] was dissolved in water and then
mixed with acid. The solution was evaporated to dryness. The residue was put in
digestion with acid and filtered. The insoluble product of this operation was, after
being dried at a red heat, recorded under the term silica.
C. The nitric solution of ash was precipitated by potassium ferrocyanide [“prus-
siate of potash”] and filtered. The solution was then concentrated by evaporation
and diluted with a small amount of water and acid. It was precipitated a second time
by potassium ferrocyanide. The separation of the precipitates was accomplished by
a double filter, and the difference between the incineration products of each single
filter indicated the amount of metallic oxide contained in the ash, after subtraction
of the weight of the metallic oxide [p. 315] in the potassium ferrocyanide.
D. The nitric solution, freed of metallic oxides and concentrated by evaporation,
was precipitated by ammonia. The precipitate, dried at a red heat, indicated very
nearly, after deducting the alumina from it, the weight of the earthy phosphates. By
this term is meant the phosphates of lime and magnesia.
E. I dissolved these phosphates in nitric acid; they were precipitated by potash
in excess; and the mixture was boiled. The filtered liquor was saturated by acid and
precipitated by ammonia, which revealed the alumina. This earth was rectified by a
new solution and precipitation. After being dried at a red heat and pulverized, it was
put in digestion in vinegar to remove the earths that were combined with it. But the
amount [of alumina] was extremely small, and often none, and in more than 40 ashes
in which I looked for it, I found that it did not exceed 1/100 of their weight. I believed
that I had found a large amount of it in the analyses I published several years ago
in the Journal de Physique [i.e., de Saussure (1800); see Translator’s Introduction],
on the incineration of some plants [p. 316] grown in different soils. But I have since
recognized that my error arose from the impurity of the ashes and the solubility of
the earthy phosphates in potash. The same kinds of plants were burned again and
analyzed more carefully.
F. The nitric solution D, freed of metallic oxides and earthy phosphates, was
precipitated by crystallized carbonate of soda. The mixture was boiled for a long
time and filtered after cooling. This operation separates the earthy carbonates, that
118 9 Observations on the Ash of Plants

is, those of lime and magnesia. I note the latter earth so as to omit nothing, for it
seemed to me that pure or carbonated magnesia was never present in ash in more
than a negligible amount. It is not the same with phosphate of magnesia, which was
discovered in ash by Fourcroy and Vauquelin.
G. The column headed deficit in the table indicates the difference between the
sum of the separate constituents and the amount placed in the experiment. This
difference is very large and, at first glance, must cast an unfavorable light on the
results. But I am satisfied that this deficit should not be [p. 317] apportioned among
all the products of analysis. It concerns only the water-soluble alkaline salts, which,
during incineration, formed with the earths and earthy phosphates a combination that
could not be completely destroyed by decoction in water. This deficit must therefore
be added to the salts recorded, deducting from it four or five hundredths for the
unavoidable loss during manipulations, in order to find very nearly the absolute sum
of the alkaline salts that the ash contains. I always found these lost salts when, after
treating with nitric acid the ash washed in water, and after precipitating the acid
solution by ammonia and carbonate of ammonia, I subjected the filtered liquor to
evaporation at the highest heat in a platinum crucible. This evaporation requires,
from the moment the salts begin to congeal, a very graduated fire and around 7 or
8 hours, so that the salts are not lost by explosions of boiling. If the fire has not
been intense enough, the salts retain a little nitrous [sic] acid, but it can be reliably
eliminated by adding a little charcoal dust while they are in fusion.
I followed these procedures [he is referring to the following procedures rather
than the preceding ones] in those analyses [p. 318] recorded as more precise [i.e., in
the observations column of his table, in the boxes headed “more precise analysis of
the same ash”]. I did not always treat the ash with boiling water [presumably he is
referring to the initial treatment with boiling water, described in A] but I dissolved
it, with its salts, in nitric acid. The latter procedure is quicker and much more exact
because, if the lyes contain phosphate of potash and lime, filtration or even precise
decanting is almost impossible because of the viscosity8 of the liquor. It holds in
solution a large amount of lime or rather phosphate of lime, whose amount can be
estimated only by dissolving the salts in an acid and precipitating them by ammonia.
I will give here the procedures that I followed in analyzing salts rich in phosphate of
potash and depleted of earthy phosphates. It is unnecessary to look for phosphate of
potash in ash containing [p. 319] more than one or two hundredths pure or carbonated
lime. Phosphate of potash is present in ash containing a larger proportion of lime,
but in such small amount that it can be ignored.
a. I mix the salts with vinegar and evaporate almost to dryness. The residue is mixed
with alcohol, which, in dissolving the acetate, becomes enriched with the greater
part of the excess potash contained in the ash. This product is set aside.9

8
Viscosity of the lye of ash, and its reduction to gel by evaporation, is an almost certain indication
of the presence of phosphate of potash and lime or magnesia. However, this salt may be present
without this characteristic, if there is a large excess of potash. Phosphate of potash is not viscous
or gelatinous by itself, whatever the proportions of its components. It has this property only when
it contains lime or magnesia without excess potash.
9
Perhaps it would be more exact to eliminate this first operation (a), which separates, by vinegar
and alcohol, only a part of the excess potash and removes a little phosphate of potash. I used this
9.11 Details on the Procedures used for Analyzing Ash 119

b. The alcohol-insoluble salts, which are phosphate of potash, the chlorides, the
sulfates, and a little free potash, are dissolved in water and mixed with acetate
of lime in excess. The latter decomposes the phosphate of potash. The mixture
is boiled strongly and evaporated almost to dryness. The magma is diluted with
a large amount of water, separated by a filter, and dried at a high heat.10 [p. 320]
It is mixed with vinegar (c) until this acid removes nothing more from it. The
insoluble residue of this operation is pure phosphate of lime, resulting from the
decomposition of the phosphate of potash. The pure phosphate of lime, augmented
in the ratio of 100–129, gives the weight of the alkaline phosphate (d) contained
in the ash without excess potash.
All the acetous solutions (a) and (c), including the wash water, are combined, evap-
orated at a red heat, and filtered. They are then evaporated again at a red heat. This
residue (e) is weighed. It contains all the potash in the ash, including that which is a
component of the potash of phosphate, and also the chlorides and alkaline sulfates.
All of these salts are dissolved in nitric acid rather than in water because the potash
contains a little carbonic acid, whose quantity is estimated from the weight loss of
the mixture. This nitric solution is precipitated successively by barium nitrate and
silver nitrate. The weight of each of these precipitates gives, by known calculations,
[p. 321] the chlorides and alkaline sulfates. The sum of these last is subtracted from
the weight (e), or from that comprising the potash, chlorides, and sulfates. The dif-
ference gives the weight of all the potash contained in the phosphate of potash and
of the potash that was in excess. Now, we know the weight of the potash that is a
component of the potash of phosphate, already given in (d). One hundred parts of
this salt contain 65 parts of potash [as explained in the first paragraph of the note
below]. Therefore, to this salt is attributed the potash belonging to it. The rest is the
potash in excess that is contained in the ash.

9.11.1 Note on the Combination of Phosphate of Potash

with Lime

I formed phosphate of potash by slowly combining, by the wet method [i.e., using
solvents, as opposed to the dry method, which uses fire], phosphoric acid with potash,
until the mixture was just neutralized and did not cause any change in the plant dyes
used as reagents. I dried this compound at a red heat and weighed it. [p. 322] It was
dissolved in water and decomposed by calcium chloride [“calcareous muriate”].
The mixture was evaporated to dryness, and, after being dissolved in water, was
filtered. I deduced from the principles recognized by Klaproth for phosphate of lime

procedure only because it makes the rest of the analysis less cumbersome and saves much acetic
acid and calcium acetate.
10
Vinegar dissolves phosphate of lime a little, if it has not been dried after its precipitation.
120 9 Observations on the Ash of Plants

that 100 parts of dry phosphate of potash contain, at the degree of saturation indicated
above:

Potash 65
Phosphoric acid 35

It is a phosphate of potash thus constituted that is referred to in the analyses of ash. It

is such that 129 parts of this salt are capable of forming only 100 parts of phosphate
of lime. [Note: See (c) and (d) under G., in Sect. 9.11, above].
Aqueous solutions of phosphate of potash are not, as has been said, viscous and
gelatinous. Even concentrated, they pass freely through the finest filters.
If this solution is mixed with 20 or 30 times its volume of limewater, the mixture,
which forms a soluble phosphate of potash and lime, retains all its transparency.
It undergoes no apparent change other than becoming viscous and passing only
extremely slowly through ordinary filters. [p. 323] If more limewater is added to it, a
precipitate begins to form. This is not phosphate of lime, it is a phosphate of potash
and lime that has become insoluble in water, through an excess of earth. When we
dissolve it in nitric acid, we withdraw from it only a quarter or a half of its weight
as phosphate of lime. Very nearly the same effects occur with magnesia, but the
compounds of this seemed less viscous to me.
The same, but less pronounced, results can be obtained by substituting a solution
of calcium acetate [“calcareous acetate”] for the limewater. If the solution is dilute,
the first drops do not decompose the phosphate of potash, or at least the precipitate
that is formed is redissolved by stirring the liquor. This observation shows that the
calcium acetate found so abundantly by Vauquelin in plant saps can exist there with
the phosphate of potash, which is completely decomposed by calcareous salts of any
kind only by evaporating the mixture to dryness. I say the same thing for magnesia.
The aqueous solution of phosphate of potash and lime is not clouded by potash,
soda, ammonia, [p. 324] or phosphoric acid, but it is partially decomposed by all the
alkaline carbonates, which precipitate carbonate of lime from it. It is also partially
decomposed by oxalic acid. Finally, it is partially decomposed by simple drying at a
red heat. In this residue is found phosphate of lime, which has become insoluble in
water through an excess of earth, and phosphate of potash and lime, which is soluble
in water but which contains less lime than in the first solution. This effect occurs
more or less in all the lyes of ash that are evaporated to dryness and redissolved in
water.
Action of Potash on Phosphate of Lime Fourcroy and Vauquelin saw clearly (An-
nales de Chimie,Year 11 [i.e., 1803]) that when liquid potash is boiled with phosphate
of lime, a very small amount of lime is deposited. But this decomposition, which is
contrary to the old order of affinities, of which Berthollet provided so many examples
in other, analogous compounds, is not the only noteworthy result in this experiment.
I observed that a very large part of the phosphate of lime is completely dissolved
in the potash, and that a phosphated potash of lime [de Saussure’s term is “potasse
9.11 Details on the Procedures used for Analyzing Ash 121

phosphatée de chaux”] then forms. [p. 325] I boiled, for an hour, 300 parts of potash
(dissolved in about twice its weight of water) with phosphate of lime in paste form
and recently precipitated by ammonia. Before the experiment, the phosphate, dried
at a red heat, weighed 25 parts. The decoction was filtered. The residue that was
undissolved by the potash weighed no more than 9 parts. It was dissolved in nitric
acid and precipitated by ammonia. The precipitate, dried at a red heat, weighed only
6 parts. The filtered solution was mixed with carbonate of soda or ammonia, which
separated from it 1/2 part of carbonate of lime. In this experiment, the potash thus
dissolved 3/4 of the phosphate of lime, and really decomposed only 1/50 of it.
I obtained almost exactly the same results by treating, with a melting fire in a
platinum crucible, dry phosphate of lime with dry potash. The phosphate weighed
20 parts, and the potash 80. The opaque glass that resulted from this mixture was
dissolved in water and filtered. The undissolved residue in this fluid weighed 12 parts,
which were dissolved in nitric acid and precipitated by ammonia. The ammonia
separated [p. 326] only 6-1/2 parts of phosphate of lime from it. The filtered solution
was precipitated by carbonate of soda, which separated 2-1/2 parts of carbonate of
lime from it.
The compounds of potash in excess with phosphate of lime that were obtained by
the procedures I have just described differ in some properties from the compound
obtained by mixing limewater with phosphate of potash.
These properties show that the potash in excess is not inessential and acts on the
lime by a very strong affinity. I have said that the aqueous solution of phosphate of
potash and lime was decomposed by oxalates. But the phosphated potash of lime is
not clouded by these reagents. They reveal lime there only if the potash that appears
in excess has been saturated very precisely with acid. It still requires that the oxalic
acid precipitate all the lime, even in this circumstance. A quadruple combination is
probably formed here [i.e., lime, potash, phosphate, and oxalate].
Liquid phosphate of potash and lime, or a mixture of limewater with phosphate
of potash, is partly decomposed by drying, but liquid phosphated potash of lime is
not significantly changed [p. 327] by this operation. The phosphate of potash and
lime forms, with water, a viscous or gelatinous solution, which does not pass easily
through a filter. The phosphated potash of lime passes very freely through a filter and
does not form a gel by concentration.
Table 9.1 Tables of Incinerations and Analyses
122

Table of Incinerations
Incineration Name of plants, time of their Ash from Ash from Water of Observations
number collection 1000 parts of 1000 parts of vegetation in
the plant green the plant dry 1000 parts of
the plant
green
1 Leaves of oak (Quercus robur), 13 53 745 In a forest; soil gravelly and
May 10 nearly sterile

2 The same, Sept. 27 24 55 549 Ibidem.

3 Debarked stems or branches of 4 Ibidem. These stems or branches

young oak, May 10 were about 1 centimeter
(5-6 lines) in diameter

4 Bark of the preceding branches 60 This bark includes the bast and
epidermis

5 Oak wood, separated from 2 This was part of a trunk that was
sapwood about 2 dm (8 inches) in
diameter
6 Sapwood of the preceding oak 4
wood

7 Bark of the preceding oak trunks 60 This bark includes the bast and
epidermis
9 Observations on the Ash of Plants
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding salts phosphates carbonates Oxides
to incineration [alkaline [phosphates [carbonates
No. salts] of lime or of lime or
magnesia] magnesia]
1. Oak leaves, 47 24 0.12 3 Less than 0.64 25.24 In this analysis and the
May 1/100 of the following ones, the deficit
weight of is attributable almost
the ash entirely to water-soluble
salts
2. The same, 17 18.25 23 14.5 1.75 25.5
Sept.
3. Debarked 26 28.5 12.25 0.12 1 32.58
stems or
branches of
9.11 Details on the Procedures used for Analyzing Ash

young oaks,
May
4. Bark of the 7 4.5 63.25 0.25 1.75 22.75
preceding
branches
5. Oak wood, 38.6 4.5 32 2 2.25 20.65
separated
from sapwood
6. Sapwood of 32 24 11 7.5 2 23.5 The silica may have been
the same oak accidental in this sapwood,
wood for I did not find it in
young branches of oak
7. Bark of the 7 3 66 1.5 2 21.5
preceding oak
123

trunks
 124

Table 9.1 (continued)

Table of Incinerations
Incineration Name of plants, time of their Ash from Ash from Water of Observations
number collection 1000 parts of 1000 parts of vegetation in
the plant green the plant dry 1000 parts of
the plant
green
8 Bast of the preceding bark 73

9 Extract of the wood of preceding 61

oak

10 Humus of oak wood 41

11 Extract of the preceding humus of 111

oak wood

12 Leaves of poplar (Populus nigra), 23 66 652 In a meadow; clay soil

May 26

13 The same, Sept. 12 41 93 565 On the same branch as the

9 Observations on the Ash of Plants

preceding leaves
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding to salts [alkaline phosphates carbonates Oxides
incineration No. salts] [phosphates [carbonates
of lime or of lime or
magnesia] magnesia]
8. Bast of the 7 3.75 65 0.5 1 22.75
preceding
bark
9. Extract of the 51 Less than Sawdust of oak, boiled for a
wood of the 1/100 of the half-hour in distilled water:
preceding weight of the filtered decoction was
oak the ash evaporated to dryness at a
gentle heat
10. Humus of the 24 10.5 10 32 1 14 8.5 This nearly-black humus was
9.11 Details on the Procedures used for Analyzing Ash

preceding taken several feet above the

wood ground, from the trunk of a
growing oak; it contained
small, white lumps of silica
11. Extract of the 66 The humus was boiled for a
humus of oak half-hour in distilled water:
wood the filtered decoction was
evaporated to dryness, at a
gentle heat
12. Poplar leaves, 36 13 29 5 1.25 15.75 In this analysis and the
May following ones, the deficit
is attributable almost
entirely to water-soluble
salts
125

13. The same, 26 7 36 11.5 1.5 18

Sept.
Table 9.1 (continued)
Table of Incinerations
126

Incineration Name of plants, time of their Ash from Ash from Water of Observations
number collection 1000 parts of 1000 parts of vegetation in
the plant green the plant dry 1000 parts of
the plant
green
14 Debarked trunks of the preceding 8 In the same soil. Trunk 2 dm (8
poplars, Sept. 12 inches) in diameter

15 Bark of the preceding trunks 72 This bark includes the bast and
epidermis

16 Leaves of hazel (Corylas 61 On the uncultivated edge of a

avellana), May 1 woods. Soil gravelly and
almost sterile

17 The same, washed with cold, 57

distilled water

18 Hazel leaves, June 22 28 62 655 Ibidem.

19 The same, Sept. 20 31 70 557 Ibidem.

20 Debarked branches of the 5 These branches were 1 centimeter.

preceding hazel, May 1 (4 lines) at their greatest
diameter
21 Bark of the preceding branches 62
9 Observations on the Ash of Plants
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding salts [alkaline phosphates carbonates Oxides
to incineration salts] [phosphates [carbonates
No. of lime or of lime or
magnesia] magnesia]
14. Debarked 26 16.75 27 3.3 1.5 24.5
poplar
trunks
15. Bark of the 6 5.3 60 4 1.5 23.2
preceding
trunks. May
16. Hazel 26 23.3 22 2.5 Less than 1.5 24.7 In this analysis and the
leaves, May 1/100 of the following, the deficit is
weight of attributable almost entirely
the ash to water-soluble salts
9.11 Details on the Procedures used for Analyzing Ash

17. The same, 8.2 19.5 44.1 4 2 22.2 These fresh leaves were
washed immersed eight times in
cold, distilled water; they
remained under water
1/4 hour at each immersion
18. Hazel 22.7 14 29 11.3 1.5 21.5
leaves, June
19. The same, 11 12 36 22 2 17
Sept.
20. Debarked 24.5 35 8 0.25 0.12 32.2
hazel
branches
21. Bark of the 12.5 5.5 54 0.25 1.75 26
preceding
127

branches
Table 9.1 (continued)
128

Table of Incinerations
Incineration Name of plants, time of their Ash from Ash from Water of Observations
number collection 1000 parts of 1000 parts of vegetation in
the plant green the plant dry 1000 parts of
the plant
green
22 Wood of so-called Spanish 7 In a vegetable garden, clay soil.
mulberry (Morus nigra) Trunk 2 dm (8 inches) in
separated from sapwood, diameter
November

23 Sapwood of the preceding 13

mulberry

24 Bark of the preceding mulberry 89 This bark included the bast and
epidermis

25 Bast of preceding bark 88

26 Wood of hornbeam (Carpinus 4 6 346 In a meadow, clay soil. Trunk 1.6

betulus) separated from dm (6 inches) in diameter
sapwood, November

27 Sapwood of the preceding 4 7 390 In this tree, the sapwood was very
hornbeam indistinct from the wood

28 Bark of the preceding hornbeam 88 134 346 This bark included the bast and
epidermis
9 Observations on the Ash of Plants
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding to salts [alkaline phosphates carbonates Oxides
incineration No. salts] [phosphates [carbonates
of lime or of lime or
magnesia] magnesia]
22. Mulberry 21 2.25 56 0.12 0.25 20.38
wood,
separated from
sapwood
23. Sapwood of the 26 27.25 24 1 0.25 21.5
preceding
mulberry
24. Bark of the 7 8.5 45 15.25 1.12 23.13
preceding
mulberry
9.11 Details on the Procedures used for Analyzing Ash

25. Bast of 10 16.5 48 0.12 Less than 1 24.38 In this and the following
preceding bark 1/100 of the analyses, the deficit is
weight of attributable almost entirely
the ash to water-soluble salts
26. Wood of 22 23 26 0.12 2.25 26.63
hornbeam,
separated from
the sapwood
27. Sapwood of the 18 36 15 1 1 29
preceding
hornbeam
28. Bark of the 4.5 4.5 59 1.5 0.12 30.38
preceding
hornbeam
129
 130

Table 9.1 (continued)

Table of Incinerations
Incineration Name of plants, time of their Ash from Ash from Water of Observations
number collection 1000 parts of 1000 parts of vegetation in
the plant green the plant dry 1000 parts of
the plant
green
29 Trunks and leafless branches of 35 In a meadow, fertile soil
horse chestnut (Aesculus
hypocastanum), May 10

30 Leaves of horse chestnut, May 10 16 72 782 Ibidem.

31 The same, July 23 29 84 652 On the same branch

32 The same, Sept. 27 31 86 636 On the same branch

33 Flowers of the preceding horse 9 71 873 On the same branch

chestnut May 10
9 Observations on the Ash of Plants
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding salts [alkaline phosphates carbonates Oxides
to incineration salts] [phosphates [carbonates
No. of lime or of lime or
magnesia] magnesia]
29. Trunks and 9.5
leafless
branches of
horse
chestnut
30. Leaves of 50
horse
chestnut,
May
31. Leaves of 24
9.11 Details on the Procedures used for Analyzing Ash

horse
chestnut,
July
32. The same, 13.5
September.
33. Flowers of 50 Less than
horse 1/100 of the
chestnut, weight of
May the ash
131
 132

Table 9.1 (continued)

Table of Incinerations
Incineration Name of plants, time of their Ash from Ash from Water of Observations
number collection 1000 parts of 1000 parts of vegetation in
the plant green the plant dry 1000 parts of
the plant
green
34 Ripe fruits of the same horse 12 34 647 On the same branch
chestnut, Oct. 5

35 Pea (Pisum sativum) plants in 95 In a vegetable garden, clay soil

flower
36 The same, bearing mature seed 81 In the same garden bed as the
preceding

37 Fava (marsh) bean (Vicia faba) 16 150 895 In a vegetable garden, clay soil
plants before flowering, May
23
38 The same, during flowering, June 20 122 876 In the same garden bed as the
23 preceding
9 Observations on the Ash of Plants
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding salts [alkaline phosphates carbonates Oxides
to incineration salts] [phosphates [carbonates
No. of lime or of lime or
magnesia] magnesia]
34. Ripe fruits 75 10.5 0.75 0.5 13.25 More precise analysis of the same ash:
of the same Potash 51
horse Phosphate of potash 28
chestnut Chlorides & alk. sulfate 3
Earthy phosphates 12
Earthy carbonates 0
Silica 0.5
Metallic oxides 0.25
Loss 5.25
9.11 Details on the Procedures used for Analyzing Ash

35. Pea plants in 49.8 17.25 6 2.3 1 24.65 100

flower
36. Pea plants 34.25 22 14 11 2.5 17.25
bearing
mature seeds
37. Fava plants, 55.5 14.5 3.5 1.5 0.5 24.50
before
flowering
38. The same, 55.5 13.5 4.12 1.5 Less than 0.5 24.38 More precise analysis of the same ash:
during 1/100 of the Incompletely carbonated potash 57.25
flowering weight of Chloride and sulfate, alkaline 12
the ash Earthy phosphates 15
Earthy carbonates 5
Silica 2
Metallic oxides 0.5
133

Loss 8.25
100
 134

Table 9.1 (continued)

Table of Incinerations
Incineration Name of plants, time of their Ash from Ash from Water of vegetation in Observations
number collection 1000 parts of 1000 parts of 1000 parts of the plant
the plant green the plant dry green
39 The same, bearing mature seeds, 66 Ibidem.
July 23

40 The same, separated from the 115 Ibidem.

mature seeds

41 Seeds of the preceding plants 33 Ibidem.

9 Observations on the Ash of Plants
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding salts [alkaline phosphates carbonates Oxides
to incineration salts] [phosphates [carbonates
No. of lime or of lime or
magnesia] magnesia]
39. The same, 50 17.75 4 1.75 0.5 26 The water-soluble salts do
bearing not contain phosphate of
mature seeds potash
40. The same, 42 5.75 36 1.75 1 12.9 More precise analysis of the
separated same ash:
from the Incompletely carbonized potash 31
mature seeds Chloride of potash 14
Sulfate of potash 2
Earthy phosphates 6
Earthy carbonates 37.5
Silica 2.75
9.11 Details on the Procedures used for Analyzing Ash

Metallic oxides 0.75

Loss 6
100
41. Seeds of the Less than Analysis:
preceding 1/100 of the Potash 22.45
plants weight of Phosphate of potash 43.93
the ash Chloride of potash 0.9
Sulfate of potash 2
Earthy phosphates 27.92
Earthy carbonates 0
Silica 0
Metallic oxides 0.50
Loss 2.30
100
135
 136

Table 9.1 (continued)

Table of Incinerations
Incineration Name of plants, time of their Ash from 1000 Ash from 1000 Water of vegetation Observations
number collection parts of the plant parts of the plant in 1000 parts of the
green dry plant green
42 Blooming fava plants grown in 39
distilled water and arising from
the preceding seeds

43 Goldenrod (Solydago vulgaris), 92 On the uncultivated edge of a

before flowering, May 1 woods; gravelly soil
44 The same, ready to flower, July 15 57 Ibidem.

45 The same, bearing mature seeds, 50 Ibidem.

Sept. 20
9 Observations on the Ash of Plants
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding salts [alkaline phosphates carbonates Oxides
to incineration salts] [phosphates [carbonates
No. of lime or of lime or
magnesia] magnesia]
42. Blooming 100 parts of ash contain:
fava plants
grown in Potash 22.4
distilled Phosphate of potash 33.4
water and Chlorides and alkaline sulfates 4.3
arising from Earthy phosphates 30
the Earthy carbonates 0
preceding Silica inapprec.
seeds quant
Metallic oxides 0.5
9.11 Details on the Procedures used for Analyzing Ash

Loss 9.4
100
43. Goldenrod, 67.5 10.75 1.25 1.5 0.75 18.25 These plants had only their
in May radicle leaves
44. The same, in 59 8.5 9.25 1.5 Less than 0.75 21 In this and the following
July 1/100 of analyses, the deficit is
the weight attributable almost entirely
of the ash to water-soluble salts
45. The same, 48 11 17.25 3.5 1.5 18.75
bearing
mature seeds
137
 138

Table 9.1 (continued)

Table of Incinerations
Incineration Name of plants, time of their Ash from 1000 Ash from 1000 Water of vegetation Observations
number collection parts of the plant parts of the plant in 1000 parts of the
green dry plant green
46 Sunflower (Helianthus annuus) 147 In a vegetable garden, clay soil
plants, June 23, a month before
flowering

47 The same, beginning to flower, 13 137 877 Ibidem.

July 23

48 The same, Sept. 20, bearing 23 93 753 Ibidem.

mature seeds

49 Blooming wheat plants In a fertile field; gravelly soil

50 The same, bearing mature seeds Ibidem.

51 Wheat (Triticum sativum) plants, 79 Ibidem.

May 1, a month before
flowering

52 The same, in flower, June 14 16 54 699 Ibidem.

53 The same, July 28, bearing 33 Ibidem.

mature seeds
9 Observations on the Ash of Plants
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding to salts [alkaline phosphates carbonates Oxides
incineration No. salts] [phosphates [carbonates
of lime or of lime or
magnesia] magnesia]
46. Sunflower 63 6.7 11.56 1.5 0.12 16.67
plants, a month
before
flowering
47. The same, 61 6 12.5 1.5 0.12 18.78
beginning to
flower
48. The same, 51.5 22.5 4 3.75 0.5 17.75
bearing mature
seeds
9.11 Details on the Procedures used for Analyzing Ash

49. Blooming 43.25 12.75 0.25 32 0.5 12.25 Year 1802

wheat plants
50. The same, 11 15 0.25 54 1 18.75 The same year, seeds
bearing mature abundant and of the finest
seeds quality
51. Wheat plants a 60 11.5 0.25 12.5 Less than 0.25 15.5 Year 1803, in the same soil as
month before 1/100 of the the preceding
flowering weight of
the ash
52. The same, in 41 10.75 0.25 26 0.5 21.5 Ibidem
flower
53. The same, 10 11.75 0.25 51 0.75 23 Seed compressed and not
bearing mature abundant
seeds
139
 140

Table 9.1 (continued)

Table of Incinerations
Incineration Name of plants, time of their Ash from Ash from Water of Observations
number collection 1000 parts of 1000 parts of vegetation in
the plant green the plant dry 1000 parts of the
plant green
54 Straw of the preceding wheat, 43
separated from the seeds

55 Wheat seeds selected from the 13

preceding
9 Observations on the Ash of Plants
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding salts [alkaline phosphates carbonates Oxides
to incineration salts] [phosphates [carbonates
No. of lime or of lime or
magnesia] magnesia]
54. Straw of the 9 5 1 61.5 1 22.5 More precise analysis of the same ash:
preceding Potash 12.5
wheat, Phosphate of potash 5
separated Chloride of potash 3
from the Sulfate of potash 2
seeds Earthy phosphates 6.2
Earthy carbonates 1
Metallic oxides r.
Loss 7.8
100
55. Wheat seed, 21 38 0 0.5 Less than 0.25 40.25 More precise analysis of the
9.11 Details on the Procedures used for Analyzing Ash

selected 1/100 of same ash:

from that the weight Potash 15
borne by the of the Phosphate of potash 32
preceding ash Chloride of potash 0.16
straw Sulfate of potash..impond. cloud
Earthy phosphates 44.5
Earthy carbonates 0
Silica 0.5
Metallic oxides 0.25
Loss 7.59
100
141
 142

Table 9.1 (continued)

Table of Incinerations
Incineration Name of plants, time of their Ash from 1000 Ash from 1000 Water of vegetation Observations
number collection parts of the plant parts of the plant in 1000 parts of the
green dry plant green
56 Bran 52

57 Maize (Zea mays) plants, June 23, 122 In a vegetable garden;

a month before flowering clay soil

58 The same, in flower, July 23 81 Ibidem.

59 The same, bearing mature seeds 46 Ibidem.

9 Observations on the Ash of Plants
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding to salts [alkaline phosphates carbonates Oxides
incineration No. salts] [phosphates [carbonates
of lime or of lime or
magnesia] magnesia]
56. Bran Potash 14
Phosphate of potash 30
Chloride of potash 0.16
Sulfate of potash 0
Earthy phosphates 46.5
Earthy carbonates 0
Silica 0.5
Metallic oxides 0.25
Loss 8.59
100
9.11 Details on the Procedures used for Analyzing Ash

57. Maize plants, 69 5.75 0.25 7.5 0.25 17.25

before
flowering
58. The same, in 69 6 0.25 7.5 0.25 17
flower
59. The same, Less than
bearing mature 1/100 of the
seeds weight of
the ash
143
 144

Table 9.1 (continued)

Table of Incinerations
Incineration Name of plants, time of their Ash from Ash from Water of vegetation Observations
number collection 1000 parts of 1000 parts of in 1000 parts of the
the plant green the plant dry plant green
60 Stems of the preceding maize 84
plants,separated from their ripe
ears

61 Ears of the preceding stems 16

62 Seeds of the preceding maize 10

9 Observations on the Ash of Plants
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding to salts [alkaline phosphates carbonates Oxides
incineration No. salts] [phosphates [carbonates
of lime or of lime or
magnesia] magnesia]
60. Stems of the 56 5 1 18 0.5 19.5 More precise analysis of the
preceding maize same ash:
plants, separated Potash 59
from their ripe Phosphate of potash 9.7
ears Chloride of potash 2.5
Sulfate of potash 1.25
Earthy phosphates 5
Earthy carbonates 1
Silica 18
Metallic oxides 0.5
9.11 Details on the Procedures used for Analyzing Ash

Loss 3.05
100
61. Ears of the
preceding stems
62. Seeds of the 24 34 0 1 0.12 40.88 More precise analysis of the same ash:
preceding maize Potash 14
Phosphate of potash 47.5
Chloride of potash 0.25
Sulfate of potash 0.25
Earthy phosphates 36
Earthy carbonates 0
Silica 1
Metallic oxides 0.12
Loss 0.88
145

100
Table 9.1 (continued)
146

Table of Incinerations
Incineration Name of plants, time of their Ash from 1000 Ash from 1000 Water of vegetation Observations
number collection parts of the plant parts of the plant in 1000 parts of the
green dry plant green
63 Straw of barley (Hordeum 42 In a field; calcareous soil
vulgare), separated from its
mature seeds

64 Barley seeds of the preceding 18 This seed was such as is used for
straw sowing, i.e., provided with its
inner husk

65 Barley seeds
9 Observations on the Ash of Plants
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding to salts [alkaline phosphates carbonates Oxides
incineration No. salts] [phosphates [carbonates
of lime or of lime or
magnesia] magnesia]
63. Straw of barley, 14 7 12.5 57 Less than 0.5 9 Potash 16
separated from 1/100 of the Sulfate of potash 3.5
its mature seeds weight of Chloride of potash 0.5
the ash Earthy phosphates 7.75
Earthy carbonates 12.5
Silica 57
Metallic oxides 0.5
Loss 2.25
100
64. Barley seeds of 7 31 0 36 0.25 25.75 More precise analysis of the same ash:
the preceding Potash 18
straw Phosphate of potash 9.2
Sulfate of potash 1.5
9.11 Details on the Procedures used for Analyzing Ash

Chloride of potash 0.25

Earthy phosphates 32.5
Earthy carbonates 0
Silica 35.5
Metallic oxides 0.25
Loss 2.8
100
65. Barley seeds 22 22 0 21 0.12 29.88 These seeds, although capable of
germinating, were collected a fortnight
before full maturity. I found, in looking
for the [water-soluble] salts remaining
in the acid solution, that they, along
with the 22 parts that are listed, weighed
47 parts. Both in this analysis and the
preceding one, a large part of the silica
147

must be attributed to the husk, from

which the seed was not entirely freed.
 148

Table 9.1 (continued)

Table of Incinerations
Incineration Name of plants, time of their Ash from 1000 Ash from 1000 Water of vegetation Observations
number collection parts of the plant parts of the plant in 1000 parts of the
green dry plant green
66 Oats 31 This seed was provided with its
husk

67 Leaves of rosage (Rhododendron 30

ferrugineum), grown on the
Jura, calcareous mountain.
June 20
68 The same, grown on the Bréven, 25
granitic mountain. June 27

69 Stems and branches of rosage 8 These stems, and the following

grown on the Jura. June 20 ones, were stripped of leaves

70 Stems of rosage grown on the

Bréven. June 27
9 Observations on the Ash of Plants
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding to salts [alkaline phosphates carbonates Oxides
incineration No. salts] [phosphates [carbonates
of lime or of lime or
magnesia] magnesia]
66. Oats 1 24 0 60 0.25 14.75 By a more precise analysis,
I found, besides the same
products, 10 parts of
potash and 5 parts of
chloride and alkaline
sulfate in the same ash
67. Rosage leaves, 23 14 43.25 0.75 0.12 3.25 15.63 In this and subsequent
calcareous. analyses, the deficit is
June 20 attributable almost entirely
to water-soluble salts
9.11 Details on the Procedures used for Analyzing Ash

68. Rosage leaves, 21.1 16.75 16.75 2 0.12 5.75 31.53 Growth was more retarded in
siliceous. June the siliceous than in the
27 calcareous soil. This
observation also applies to
all the following harvests
69. Rosage stems, 22.5 10 39 0.5 0.12 5.4 22.48
calcareous.
No. 67. June
70. Rosage stems, 24 11.5 29 1 11 24.5
siliceous. June
149
 150

Table 9.1 (continued)

Table of Incinerations
Incineration Name of plants, time of their Ash from Ash from Water of vegetation Observations
number collection 1000 parts of 1000 parts of in 1000 parts of the
the plant green the plant dry plant green
71 Needles of spruce (Pinus abies) 29
grown on the Jura. June 20
72 The same, grown on the Bréven, 29
June 27

73 Spruce branches stripped of 15

needles, June 20

74 Bilberry (Vaccinium myrtillas) 26

grown on the Jura. August 29

75 The same, grown on the Bréven. 22

August 20
9 Observations on the Ash of Plants
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding to salts [alkaline phosphates carbonates Oxides
incineration No. salts] [phosphates [carbonates
of lime or of lime or
magnesia] magnesia]
71. Spruce needles, 16 12.27 43.5 2.5 1.6 24.13
calcareous. June
72. Spruce needles, 15 12 29 19 5.5 19.5 In this analysis and the
siliceous. June following ones, the deficit
is attributable almost
entirely to water-soluble
salts
73. Spruce branches 15
stripped of
needles
9.11 Details on the Procedures used for Analyzing Ash

74. Bilberry, 17 18 42 0.5 3.12 19.38

calcareous.
August 29
75. Bilberry, 24 22 22 5 9.5 17.5
siliceous.
August 20
151
 152

Table 9.1 (continued)

Table of Incinerations
Incineration Name of plants, time of their Ash from 1000 Ash from 1000 parts Water of vegetation Observations
number collection parts of the plant of the plant dry in 1000 parts of the
green plant green
76 Humus of rosage, calcareous, of 65 This black humus was taken from
Nos. 67 and 69 a rock of pure carbonate of
lime, which livestock could not
reach; it was exposed in the
open air to all atmospheric
influences
77 Extract of the preceding humus 140 This dry extract, black and
semi-transparent, was prepared
by repeated decoctions of the
humus in distilled water
renewed at each decoction. The
decoctions were cloudy and
could be filtered only by a
straining-bag, which did not
clarify them

78 Humus of rosage, siliceous, of 620 The large amount of ash

Nos. 68 and 70 contained in this humus was
caused by its mixture with the
sand or detritus of the gneiss
that underlay it
79 Extract of the preceding humus 142
9 Observations on the Ash of Plants
Table 9.1 (continued)
Table of Analyses
Constituents of 100 parts of the ash
Analysis No., Water-soluble Earthy Earthy Silica Alumina Metallic Deficit Observations
corresponding to salts [alkaline phosphates carbonates Oxides
incineration No. salts] [phosphates [carbonates
of lime or of lime or
magnesia] magnesia]
76. Humus of 0.5 6 29 28 3 18 15.5 I found 12 parts of alkaline
rosage, salts in the acid solution of
calcareous, of the ash freed of earths and
Nos. 67 and 69 oxides. The deficit is thus
only 3 parts. This humus did
not effervesce with acids

77. Extract of the 27 16.75 21 3 0.12 3 29.13 More precise analysis of the
preceding same ash:
humus Carbonated potash 14
Chloride of potash 23
9.11 Details on the Procedures used for Analyzing Ash

Sulfate of potash 16
Earthy phosphates 17.25
Earthy carbonates 21.5
Silica 3.25
Alumina 0.12
Metallic oxides 3
Loss 1.88
100
78. Humus of 0
rosage, siliceous

79. Extract of the 24 13 17 14 0.12 10 21.88

153

preceding
humus
Chapter 10
Chapter Summaries

By the Translator

Chapter 1. Influence of oxygen gas on germination. Seeds cannot germinate

without water and oxygen gas. Oxygen does not unite directly with the substance of
the seed. All germinating seeds give off carbon dioxide, and the volume emitted is
nearly the same as the volume of oxygen absorbed; thus oxygen’s influence seems
to be limited to removing a part of the carbon that the seed acquired when it was
formed. Seeds will not germinate in atmospheres of pure nitrogen, hydrogen, or
carbon dioxide. The carbon dioxide produced during germination inhibits continued
germination more than does a similar amount of nitrogen or hydrogen gas. For a given
species of plant, the amount of oxygen required for germination varies in proportion
to the size of the seeds and is, overall, very small. Putrefying seeds give off carbon
dioxide, methane, and nitrogen gases. During germination, seeds produce sugar, but
not as a result of the direct combination of oxygen gas with a substance in the seed.
Seeds have a lower dry weight after germination, even when corrected for the carbon
lost as carbon dioxide. This additional weight loss is attributable to the formation and
release of water from the seeds’dry substance. Light is not detrimental to germination.
Chapter 2. Influence of carbon dioxide gas on plant growth. Green plants ex-
posed to sunlight must break down carbon dioxide in order to grow. Germination and
the stages of growth immediately following it, however, are inhibited by this gas.
Extra carbon dioxide added to a green plant’s environment stimulates plant growth,
up to a point. Plants grown in sunlight, with their roots immersed in water that was
lightly impregnated with carbon dioxide, thrived and gained weight. Plants grown
with their roots in distilled water and with atmospheres containing added carbon
dioxide gas showed various responses. If they were exposed to the sun, in an at-
mosphere that was 8 percent carbon dioxide, they thrived, but at higher levels of
added carbon dioxide, their growth declined, and, at levels of 50 percent and above,
they died. For plants grown in the shade, growth was inhibited even in atmospheres
containing only 8 percent carbon dioxide. For plants grown in sunlight, if quicklime
was present to absorb the carbon dioxide of the air and water, the plants soon died
and vitiated their atmosphere.
Plants grown with their shoots in open air, in the light, and their roots in distilled
water assimilated carbon, proving that these plants must be obtaining their carbon
solely from the atmosphere, miniscule though that source is, because they could not

J. F. Hill, Chemical Research on Plant Growth, 155

DOI 10.1007/978-1-4614-4136-6_10, © Springer Science+Business Media New York 2013
156 10 Chapter Summaries

be drawing it through their roots. Plants of the same weight, grown in darkness, lost
carbon. The amount of carbon dioxide decomposed by the green parts of plants is
approximately equal to the amount of oxygen they liberate, but a small percentage of
the oxygen is retained by the plant, as is all of the carbon. Plants release nitrogen gas.
Chapter 3. Influence of oxygen gas on developed plants. The most evident effect
of oxygen gas on developed plants is to elicit the production of carbon dioxide from
the combination of this oxygen with the plants’own carbon. At night, leaves “inspire”
atmospheric oxygen and partly replace it by “expiring” carbon dioxide, whereas
during the daytime the reverse occurs, and overall the atmosphere in a closed space
remains about constant in volume. If the leaves are very thick, however, as in cactus,
they absorb the oxygen without releasing carbon dioxide. The production of carbon
dioxide gives back to plants a substance that they can decompose and thereby regain
the carbon that they have lost. The formation of carbon dioxide from atmospheric
oxygen and plant carbon was confirmed for leaves of 57 species.
Oxygen is essential to plant growth and development. For example, germinating
seeds require it; roots need it and die if put in nitrogen, hydrogen, or, especially,
carbon dioxide, even if the shoot is exposed to air and light; and shoots survive in
oxygen-free environments only if their green parts are exposed to light, allowing
them to break down carbon dioxide that they have formed from their own internal
oxygen and carbon, thereby producing and releasing enough oxygen to sustain them-
selves. Plant parts that are growing or otherwise physiologically active require more
oxygen than do less active parts. The oxygen inspired cannot be recovered by an
air pump. In non-green plant parts, the carbon dioxide produced by the combina-
tion of atmospheric oxygen with plant carbon probably ascends with the sap to be
decomposed by the leaves rather than being fixed directly in the absorbing organ.
Chapter 4. Influence of oxygen gas on some plant-derived substances. In plants
that are in the early stages of fermentation, or have just died, oxygen gas is not fixed
in the tissues nor does it combine with their hydrogen; it only removes a part of their
carbon, forming carbon dioxide. This effect is evident in the precipitation of plant ex-
tracts, the formation of acetic acid from wine, changes in wood, coagulation of plant
albumin, oxidation of turpentine oil, and putrefaction. Olive oil, in contrast, releases
no carbon dioxide. As carbon dioxide is released from substances, some of the water
that had previously been fixed in the plant is also removed. As fermentation of plant
substances proceeds, more water is disengaged, and the oxygen absorbed does not
appear entirely in the carbon dioxide produced but probably combines directly with
the hydrogen in the plant to form water. Thus, the proportion of carbon in the residue
is not reduced after carbon is withdrawn through its combination with oxygen.
Chapter 5. Of plant humus. Humus is the black substance into which dead plants
are converted by the combined action of oxygen and water. Oxygen gas penetrates
humus and combines with its carbon to produce carbon dioxide gas, which is released,
thereby becoming a nutrient for plants. Carbonization of 47 plants, plant parts, or
plant substances showed that humus contains more carbon per unit weight than the
plants that gave rise to it. The proportion of carbon contained by a humus, however,
10 Chapter Summaries 157

does not seem to be increased significantly by the continued action of the causes that
formed it. In a finished humus, although continuous exposure to air and rain leads to
continued weight loss, there is little change in the proportions of carbon, hydrogen,
and oxygen because the hydrogen and oxygen are removed, in the form of water,
in proportion to the removal of carbon as carbon dioxide. After multiple extractions
of humus, the humus still yields a small amount of extract. Acids do not dissolve
the organic matter of humus, but potash and soda do. Humus is entirely destructible
at room temperature due to the joint action of oxygen and water. As a humus loses
carbon and water, a small amount is also being lost as extract to the soil drainage
water. These changes explain why there is little accumulation of humus, even in
places where vegetation has long been established. Incineration of humus yields a
semi-vitreous ash which is not very soluble in water. The ash of the extractive juices
of a humus, however, contains all the substances of plant ash. Thus, the extractive
juices contribute to the fertility of the humus. Humus contains a larger proportion
of nitrogen than do undecomposed plants. Humus has an antiseptic action, which
retards fermentation and putrefaction.
Chapter 6. Of plant growth in environments lacking oxygen gas. Plants can sur-
vive without oxygen in their atmospheres only if they have enough area of green
tissue to produce free oxygen through the decomposition of the carbon dioxide that
they form entirely from their own tissues. Oxygen seems to aid photosynthetic as-
similation of carbon dioxide. Thus, in environments lacking oxygen, the elaboration
of a certain amount of oxygen appears necessary for the decomposition of a certain
amount of carbon dioxide. If the oxygen is removed as it is formed, plant growth is ar-
rested. Seeds will not germinate in oxygen-free environments, and newly germinated
seeds, before they have developed green parts, will die in these environments. Plants
do not assimilate nitrogen, hydrogen, or carbon monoxide gases. They survive in
these gases as they do in a vacuum, by means of the oxygen released by their leaves,
and they can do so only if they are not in direct sunlight. The amount of oxygen
needed to sustain a plant’s life is small; more is needed for growth. Carbon dioxide
always becomes harmful to plants if it is present in amounts too large for them to
decompose. Excess carbon dioxide is more harmful to plants in an atmosphere of
nitrogen than in one of ordinary air. In pure hydrogen gas, the carbon dioxide formed
by the leaf from its own substance is decomposed by the hydrogen, with production
of water and carbon monoxide gas. In an atmosphere of carbon monoxide, green
plants, in the sun, do not decompose this gas but do add oxygen.
Chapter 7. Of the fixation and decomposition of water by plants. Plants do
not decompose water directly, appropriating its hydrogen and immediately releasing
its other element in the form of oxygen gas. Instead, they exhale oxygen gas only
through the decomposition of carbon dioxide gas. They retain a small amount of
oxygen from this decomposition, but it does not account for all the oxygen that
plants contain. Rather, the assimilation of the hydrogen and oxygen of water is the
source of most of the oxygen and all of the hydrogen in plant dry substance. The
amounts of hydrogen and oxygen in plants cannot be increased significantly without
correspondingly increasing the amount of their carbon.
158 10 Chapter Summaries

Dead plants in a heap, decaying in the absence of oxygen, form carbon dioxide gas
entirely from their own substance. The oxygen component of this carbon dioxide is
the oxygen of the water that was previously fixed during growth. Growing plants, in
decomposing this newly formed carbon dioxide, eliminate oxygen that had originally
been part of the water.
Plants with thick leaves, such as cactus, liberate oxygen in air lacking carbon
dioxide. They do this by breaking down carbon dioxide that they form from their
own substance. If the carbon dioxide is removed as it is formed, by alkali, no free
oxygen is produced. The ability of these plants to form carbon dioxide from their own
substance is a result of the low porosity of their epidermis and their large volume,
which protects much of their substance from exposure to the outside atmosphere.
Although a plant adds small amounts of oxygen to atmospheres initially lacking
both oxygen and carbon dioxide, this does not prove that the plant has decomposed
water. Rather, the plant has decomposed carbon dioxide that it forms entirely from
the carbon and oxygen it already contains.
In contrast, in atmospheres that contain oxygen, plants do not form carbon dioxide
entirely from their own substance, but only from a combination of their carbon with
the atmospheric oxygen. The decomposition of this carbon dioxide by the plant
would then simply return to the atmosphere the free oxygen that the atmosphere had
previously contained. This process should result in no net increase in atmospheric
oxygen. Since experiments show no increase in oxygen content, there is no direct
decomposition of water by plants.
Chapter 8. Of the absorption of solutions by plant roots. In addition to water and
gases, developing plants need minerals from the soil, even though these substances
occur in very small amounts in plants. From ten test solutions, each containing
the same amount of a different mineral constituent, roots took up varying amounts
of minerals, but absorbed none of them in greater proportion than the water of
solution. Variations in assimilation also occurred for solutions containing several
mineral solutes. Roots absorbed toxic as well as beneficial substances. The minerals
that were absorbed varied with the plant species. The dry weight of the extract that
the most fertile soil can produce is much less than the dry weight of a plant that
develops there, so the plant clearly can be taking only a very small percentage of its
dry matter from that source, even though such substances are essential for growth.
Copper sulfate is toxic, killing the roots and thus destroying their selective power
of absorption. Sluggish growth, and the cutting off of roots, also promote the
unrestricted absorption of solutes.
Variations in assimilation are attributable largely to differences in viscosity. The
less viscous a substance, the more easily it penetrates the root. Gum, which yields a
very viscous solution, is assimilated to a much lesser extent than is sugar.
Most of a plant’s substance is derived from carbon dioxide and water. The roots
of green plants absorb only a very small amount of salts and other constituents from
the extractive and saline solutions of the humus, yet these constituents have a very
pronounced effect on plant growth.
10 Chapter Summaries 159

Chapter 9. Observations on the Ash of Plants. The ash left after plants or plant
derivatives are incinerated consists of the minerals that the plants had contained.
Measurements of ash content and mineral analyses of the ash were made on 79 dif-
ferent plants or plant parts, plant products, and humuses or extracts, including plants
of different ages or ones grown in different soils. Results are presented in tables of
incineration and mineral analyses. The table of mineral analyses lists the percent-
ages of water-soluble salts, insoluble phosphates and carbonates, silica, alumina,
and metallic oxides in the ash.
In general, herbaceous plants contain more ash than do woody plants. Ash is
more abundant where transpiration is profuse. Leaves yield more ash than stems
or fruits, and bark more than wood. The leaves of deciduous trees, which have
greater transpiration than the leaves of evergreen trees, yield more ash. The ash of
herbaceous annuals increases at first, but declines as leaves gradually die and are shed
or minerals are lost to leaching by rain. A decayed plant that has not been exposed
to heavy leaching yields more ash than a living plant of the same kind.
Mineral amounts are presented as percentages. Thus, after the leaching or other
loss of certain minerals, the remaining, less easily leached minerals occur in appar-
ently greater abundance, although they are not so in absolute terms. For example,
the alkaline salts are easily leached, so, as plants age and are exposed to weathering,
the relative proportions of other minerals increase.
The proportions of the elements in plant ash are almost always related to the
soils in which the plant grew. Thus, plants growing in a soil rich in silica contain
more silica and less lime than the same kinds of plants growing in a calcareous soil.
Plants grown in the open air, with their roots immersed in distilled water, develop
abnormally because they can obtain minerals only from the dust that settles from
the atmosphere. The ash of these same kinds of plants, grown in humus, contains
significantly more saline and earthy materials, including minerals that could not have
come from the seed.
Minerals are essential to plants. Certain minerals that are taken in might indeed
be inessential, but this has not been demonstrated for those minerals that are always
present in plants of the same species.
The text concludes with a lengthy description of the chemical methods used to
analyze the mineral composition of ash.
Appendices

Appendix 1: Conversions of Units of Measurement

Before the French Revolution, many systems of measurement were in use in different
regions of France, but the Paris system was generally used by Parisian scientists such
as Antoine Lavoisier and by other scientists writing in French, including Théodore de
Saussure. During the French Revolution, the metric system supplanted the Parisian
system. Although the units of measurement in the Paris system, when translated into
English, sound familiar (pouce is inch, livre is pound), the quantities represented
by these units are not quite the same in the two measurement systems: The Paris
pound was slightly smaller than the English avoirdupois pound, and the Paris inch
was slightly longer than the English inch. De Saussure wrote Recherches chimiques
sur la Végétation during the transition to the metric system and reported most of his
experimental findings in units of both the metric and the older, Parisian systems.
(He did not use any English units, but I have provided conversions, for general
information.)
Weight measurements, Paris and English systems:
1 Paris pound (livre) = 16 Paris ounces (onces) = 9216 Paris grains (grains)
1 Paris ounce (once) = 8 gros (gros is an obsolete French unit that has no equivalent
term in English.)
1 gros = 72 Paris grains (grains)
1 Paris pound = 0.926 English avoirdupois pound
1 Paris grain = 0.820 English Troy grain
Length and Volume Measurements, Paris and English Systems:
1 Paris inch (pouce) = 12 lines (lignes) = 1.066 English inches
1 Paris cubic inch (pouce cube) = 1.211 English cubic inches
Metric Equivalents for Paris Units of Weight, Length, and Volume:
1 Paris pound = 489.506 g
1 Paris ounce = 30.594 g
1 gros = 3.824 g
1 Paris grain = 0.053 g

J. F. Hill, Chemical Research on Plant Growth, 161

DOI 10.1007/978-1-4614-4136-6, © Springer Science+Business Media New York 2013
162 Appendices

1 Paris inch = 2.7 cm

4.4 Paris lines = 1 cm
1 Paris cubic inch = 19.84 cm3
Réaumur Temperature Scale:
In his book, de Saussure exclusively used the Réaumur (abbreviated “R”) temperature
scale, in which the freezing point of water is 0◦ and the boiling point 80◦ , assuming
a standard, 1 atmosphere of pressure. The Réaumur scale was named for French
scientist René Antoine Ferchault de Réaumur. To convert from the Réaumur to the
Celsius scale, multiply the Réaumur degrees by 5/4. To convert from the Réaumur
to the Fahrenheit scale, multiply the Réaumur degrees by 9, divide the product by 4,
and add 32. (1◦ Réaumur = 2.25 ◦ F).
Amount of Carbon and Oxygen in Carbon Dioxide:
De Saussure stated in his book that he used Lavoisier’s figures for determining the
proportions of the weights of carbon and oxygen in carbon dioxide: 28 parts of
carbon and 72 parts of oxygen. To determine the weight of carbon and oxygen in a
given volume of carbon dioxide gas (at set readings of temperature and pressure),
de Saussure used conversion factors of approximately 0.19 Paris grain of carbon and
0.49 Paris grain of oxygen, per Paris cubic inch.
Appendix 2: Glossary of Terms 163

Appendix 2: Glossary of Terms1

Absorption (absorption) de Saussure often used this term to refer to the assimilation
of a gas into a plant’s dry substance; distinguished from “consumption” (q.v.)
Acetic (acétique) relating to, or containing, acetic acid, or vinegar (C2 H4 O2 )
Acetous (acéteux) tasting or smelling like vinegar; acetous acid (acide acéteux),
which is impure acetic acid, is an older name for acetic acid (acide acétique); although
de Saussure did not use the term acetous acid, he did mention acetous solutions
Aeriform (aériforme) gaseous
Alkaline salts (sels alcalins) salts of potash (potassium, K) or soda (sodium, Na),
but potassium and sodium had not been isolated in de Saussure’s day; the salts in-
cluded carbonates, chlorides (e.g., NaCl, KCl), sulfates, and phosphates; de Saussure
also referred to these as “the water-soluble salts”
Alkaline sulfates (sulfates alcalins) Na2 SO4 , K2 SO4 , but sodium and potassium
had not yet been isolated
Alumina (alumine) aluminum oxide (Al2 O3 )
Ammonium aluminum sulfate (sulfate ammoniacal d’alumine) also called am-
monium alum
Antiseptic (antiseptique) a substance that arrests the process of putrefaction or
putrid fermentation
Baryta (Baryte) barium hydroxide (Ba(OH)2 ); de Saussure used acetate of baryta
(Ba(C2 H3 O2 )2 ), nitrate of baryta (Ba(NO3 )2 ), and water of baryta (that is, fully
dissolved in water, Ba+2 + 2OH− ) as reagents for precipitation; the element barium
was not isolated from baryta until 1808, by Humphrey Davy2
Calcareous spar (spath calcaire) calc-spar, or calcite
Caloric (calorique) also called “matter of heat”; a precursor to the concept of en-
ergy; chemists, impressed by the enormous amount of heat liberated during the
combustion of inflammable bodies in oxygen, assumed that oxygen gas was a com-
bination of a very large amount of caloric with a “base” of oxygen or an “oxygenous
principle”; all gases were considered to consist of a combination of caloric with a

1
This glossary is organized based on the literal translations of terms used by de Saussure. His
original French terms are given in parentheses, then modern equivalents are provided and, where
appropriate, expanded explanations of the terms, including modern chemical formulae, although
these formulae had not been developed in his day. (For an explanation of the handling of chemical
terminology in the text of the translation, see the Note on the Translation.)
2
Davy, H. 1808. Electro-chemical researches, on the decomposition of the earths; with observations
on the metals obtained from the alkaline earths, and on the amalgam procured from ammonia. Phil.
Trans. Roy. Soc. London 98:333–370.
164 Appendices

principle or base of the gas; de Saussure thought that the caloric released by the
combination of oxygen gas with the carbon of the plant was useful to the plant
Carbonate of lime (carbonate de chaux, carbonate calcaire) calcium carbonate
(CaCO3 ), but the element calcium was not isolated until 1808, by Humphrey Davy3
Carbonic acid gas (gaz acide carbonique) carbon dioxide (CO2 )
Carburetted hydrogen gas (gaz hydrogène carburé) methane (CH4 )
Closed (fermé) sealed off or separated from the external environment; de Saussure
typically applied this term to inverted receptacles (vessels) that were partially filled
with a liquid, either water or mercury, with the mouth of the vessel resting in a saucer
or basin of water or mercury; such a system is “closed” with respect to the external
air
Consumption (consumption) de Saussure often used this term to refer to a plant’s
uptake of a gas, as distinct from absorption, or assimilation of a gas into a plant’s
dry matter
Decoction (décoction) extraction of substances by more or less prolonged contact
with a solvent or by boiling
Earth (terre) when used in a chemical sense, this usually refers to a carbonate,
oxide, or hydroxide; may be calcareous, magnesian, or siliceous
Earthy carbonates (carbonates terreux) a collective term for calcium carbonate
(CaCO3 ), which de Saussure referred to as “carbonate of lime”, and magnesium
carbonate (MgCO3 ), which he referred to as “carbonate of magnesia”
Earthy phosphates (phosphates terreux) calcium or magnesium combined with
phosphate in various forms; de Saussure called these compounds “phosphate of
lime” or “phosphate of magnesia”
Elaboration (élaboration) decomposition, as of carbon dioxide by green plant
tissue in the presence of light
Elastic (élastique) in a gaseous, as distinct from a fixed or solid, state
Element, principle (élément, principe) the concept of “element” was not well
defined in de Sausure’s day, although Lavoisier4 had held, correctly, that elements
were substances that chemical analysis failed to break down into simpler substances;
de Saussure sometimes used the older term “principle” in the sense of “element” or
“substance”, and, in this translation, depending on the context, one of these latter
terms is used instead of “principle”

3
Davy 1808. Electro-chemical researches.
4
Lavoisier, A. 1789. Traité Élémentaire de Chimie, présenté dans un ordre nouveau et d’après les
découvertes modernes. 2 vols. Paris: Cuchet. (1790 English translation by Robert Kerr, as “Ele-
ments of Chemistry”. Edinburgh: Printed for William Creech; reprinted 1965, Dover Publications,
New York).
Appendix 2: Glossary of Terms 165

Empyreumatic oils (huiles empyreumatiques) liquid oils obtained by distilling

various organic substances at high temperatures; these oils are acidic and soluble;
they do not retain the taste and odor of the substance from which they were obtained,
but instead have a taste and/or odor of burnt plant or animal matter5
Eudiometer (eudiomètre) a kind of gas burette used to measure the “goodness”
of air, that is, its oxygen content; there were various eudiometric techniques, using
different chemicals—for example, nitric oxide (a method called the “nitrous air
test” by Priestley, who first developed it); phosphorus; and potassium sulfide (K2 S,
de Saussure’s “hydrosulfide of potash”); de Saussure considered the phosphorus and
potassium sulfide methods superior to the nitrous air test; another technique involved
sparking a mixture of hydrogen and the air being tested
Fava bean (fève) Vicia faba; an Old World bean also known as faba bean or broad
bean; compare with “haricot”
Fermentation (fermentation) a process not well understood in de Saussure’s day,
as yeasts and bacteria were not yet known; chemists generally referred to three
kinds of fermentation—spiritous (which produced alcohol); acid (which produced
vinegar, or acetic acid); and putrid (which produced ammonia and was also known
as putrefaction)
Flowers of sulfur (fleurs de soufre) sublimed sulfur in flower-like crystalline form;
flakes or powder
Haricot (haricot) the seed or unripe pod of the kidney bean (Phaseolus vulgaris),
a NewWorld, climbing string bean; compare with “fava bean”
Humus, plant mold (terreau, humus, terre végétale) de Saussure used all these
terms, without clear distinctions, but he mostly used “terreau”, which, along with
“humus”, is translated here as “humus”, whereas “terre végétale” is translated as
“plant mold”; Feller and Manlay6 maintain that de Saussure ascribed a broad meaning
to the word “humus” (i.e., the whole vegetative cover undergoing decomposition)
and a narrower one to the word “mold” (i.e., the black substance in which plants are
embedded); “terre végétale” can also be translated as topsoil
Hydrogen (gaz hydrogène) H2 ; see also “inflammable gas”
Hydrosulfide (hydrosulfure) older term for sulfide; in some of his eudiometric
tests, de Saussure used “hydrosulfide of potash” (potassium sulfide, K2 S)

5
Eklund, J. 1975. The incompleat chemist: being an essay on the eighteenth-century chemist in his
laboratory, with a dictionary of obsolete chemical terms of the period. Smithsonian Stud Hist Tech
33. Washington, DC: Smithsonian Institution Press.
6
Feller, C., and Manlay, R. 2001. Humus, fertility and sustainability of cropping systems: concepts
over the past three centuries. Publication No. 147. Université Laval, Faculté de foresterie et de
géomatique, Département des sciences du bois et de la forêt, Groupe de coordination sur les bois
raméaux. Quebec, Canada.
166 Appendices

Inflammable gas (gaz inflammable) older term for hydrogen gas, but was also used
for a variety of hydrocarbons
Lime, slaked (chaux éteinte à l’eau) calcium hydroxide (Ca(OH)2 ); formed by
mixing unslaked lime (CaO), also called quicklime, with water
Limewater (eau de chaux) a solution of calcium carbonate (CaCO3 ); de Saussure
used limewater to test for the presence of carbon dioxide, which was detected by the
formation of an insoluble, white precipitate of calcium carbonate; he also used this
technique to remove carbon dioxide from gas mixtures
Lute (lute) to seal or cement, as with clay, to make an impervious fitting
Magnesia (magnésie) magnesium carbonate (MgCO3 )7 ; modern magnesia is mag-
nesium oxide (MgO); magnesium was not isolated until 1808, by Humphrey
Davy8
Metallic oxides (oxydes métalliques) de Saussure mentions oxides of iron, man-
ganese, and lead
Muriates (muriates) chlorides; de Saussure referred to sodium chloride (NaCl) as
“muriate of soda”, and potassium chloride (KCl) as “muriate of potash”
Muriatic acid (acide muriatique) hydrochloric acid (HCl)
Niter (nitre) potassium nitrate (KNO3 ), also called “saltpeter”
Nitrogen (azote) in French, the term “azote” is still used for nitrogen
Nitrous air test (gaz nitreux) a test developed by Priestley for the presence of
oxygen gas; actually uses nitric oxide (see Translator’s Introduction, footnote 9)
Oxalate (oxalate) the anion C2 O4 2−
Oxide of carbon gas (gaz oxyde de carbone) carbon monoxide (CO); in de Saus-
sure’s day, hydrogen was often confused with a number of other inflammable gases,
and the possible presence of hydrogen in oxide of carbon gas was still a subject
of debate; thus, the chemist Berthollet called oxide of carbon gas “oxycarburetted
hydrogen”, as mentioned by de Saussure on p. 208 of his 1804 book; de Saussure
set forth his own views on the matter in footnote 7 on this same page; oxycarbu-
retted hydrogen is now known to be a water gas mixture, that is, hydrogen, carbon
monoxide, and carbon dioxide9
Oxygenated muriatic acid (acide muriatique oxygené) chlorine (Cl2 ); in
Lavoisier’s chemical nomenclature, there was some confusion: hydrochloric acid
was called “muriatic acid”, and chlorine was called “oxygenated muriatic acid”
Pellicle (pellicule) a thin saline crust that forms on a solution

7
Eklund 1975.
8
Davy 1808. Electro-chemical researches.
9
Eklund 1975.
Appendix 2: Glossary of Terms 167

Phosphate of potash and lime or magnesia (phosphate potassé de chaux ou

de magnésie) salts that de Saussure obtained while trying to isolate phosphates;
Boussingault10 noted this phrase in de Saussure’s book as indicating that de Saussure
recognized that some crystalline substances dissolve as if they were two salts (i.e.,
in modern notation, Ca K(PO4 ) · K Mg(PO4 ))
Phosphated potash of lime (potasse phosphatée de chaux) Ca K(PO4 ); a salt that
de Saussure obtained while trying to isolate phosphates
Plant growth (végétation) see “vegetation”
Plant organization (organisation végétale) older term for plant structure
Platina (platine) platinum (Pt); the term platina is still sometimes used today to
refer to the natural form of platinum, with its impurities
Potash (potasse) usually, potassium carbonate (K2 CO3 ), but sometimes other com-
pounds of potassium; the element potassium was not discovered until 1808, when
Humphrey Davy11 isolated it from potash, but the “basis of potash” was suspected
before the discovery of potassium, as is evident in terms such as caustic potash
(KOH), vitriol of potash (K2 SO4 ), and muriate of potash (KCl)12 ; in this translation,
“potassium” is substituted for “potash” in cases where de Saussure is using the term
as part of the name of a compound that is still a valid compound today, or where the
general context does not make substitution confusing for the reader
Powdered or effloresced (effleuri) describes a substance that is changed from a
crystalline to a powdery form through loss of the water of crystallization when
exposed to air, or that develops a powdery crust as a result of evaporation or chemical
change; de Saussure mentioned powdered soda and powdered sulfate of soda
Proximate constituents of plants (principes immédiats des végétaux) refers to
plant components obtained through chemical analysis; these components are com-
pounds, such as acids, oils, camphor, and gum, that are presumed to be simpler than
the original substance13
Prussiate of potash (prussiate de potasse) potassium ferrocyanide (K4 [Fe(CN)6 ] ·
3H2 O), a reagent used by de Saussure in analyzing for metallic oxide in ash

10
Boussingault, J.-B. 1843. Économie rurale considérée dans ses rapports avec la chimie, la
physique et la météorologie. Vol. 1, p. 102. Paris: Béchet jeune. (Translated into English in 1845 by
George Law, as Rural Economy in its Relations with Chemistry, Physics, and Meteorology. London:
H. Bailliere).
11
Davy H. 1808. The Bakerian Lecture, on some new phenomena of chemical changes produced
by electricity, particularly the decomposition of the fixed alkalies, and the exhibition of the new
substances which constitute their bases; and of the general nature of alkaline bodies. Phil Trans Roy
Soc London 98:1–44.
12
Ihde, A. J. 1964. The Development of Modern Chemistry. P. 94, footnote 2. New York, Evanston,
and London: Harper & Row.
13
Eklund 1975.
168 Appendices

Putrefaction (putrefaction) also known as putrid fermentation; produces ammonia

Pyroligneous acid (acide pyroligneux) crude acetic acid from wood (HC2 H3 O2 )14 ;
de Saussure obtained this acid as a product of wood distillation
Quicklime (chaux vive) calcium oxide (CaO); also called unslaked lime; mixing it
with water yields calcium hydroxide (Ca(OH)2 ), or slaked lime (see “lime, slaked”)
Recipient (récipient) a vessel or receptacle that was an important piece of equip-
ment used by de Saussure and other plant physiologists of his day for studying the
relationship between gases and plant life; so-called because chemists used these ves-
sels to “receive” substances produced by distillation; de Saussure’s recipients were
generally of glass and cylindrical in shape; the experimental plant was introduced
from below, into the inverted recipient, which contained either ordinary air or an ex-
perimental atmosphere; water or mercury (or both) in the lower part of the recipient
and in a saucer beneath the mouth of the recipient served to seal the apparatus from
the external air; the gases that remained at the end of the experiment were analyzed
by means of eudiometric or other techniques; examples of recipients are shown in
de Saussure’s plate of figures
Red heat (chaleur rouge) refers to a high temperature used for carrying out
chemical reactions
Saltpeter (salpêtre) potassium nitrate (KNO3 ), also called “niter”
Sap, juice (sève, suc) sap or juice of a plant
Sapwood (aubier) de Saussure considered sapwood to be formed by a tree’s sap;
distinguished from older wood (see “wood, fully formed or well formed”)
Sea salt (sel marin) sodium chloride (NaCl)
Secretion (secrétion) de Saussure used this term both in the sense of absorption of
a substance by a plant, as through the roots (see his pp. 253, 256), and of excretion
of a substance by a plant (see his p. 155)
Septic (septique) causing or resulting from putrefaction
Silica (silice) silicon dioxide (SiO2 )
Soda (soude) this term, like the term “potash”, was not well defined in de Saussure’s
day; it could refer to sodium oxide (Na2 O), sodium bicarbonate (NaHCO3 ), sodium
carbonate (Na2 CO3 ), or sodium hydroxide (NaOH); the element sodium was not
discovered until 1808, when Humphrey Davy15 isolated it from soda, but, as with
potash, the “basis of soda” was suspected before the isolation of sodium; in this
translation, “sodium” is substituted for “soda” in cases where de Saussure is using
the term as part of the name of a compound that was known at the time and is still valid

14
Eklund 1975.
15
Davy 1808. The Bakerian Lecture.
Appendix 2: Glossary of Terms 169

today, or in cases where the general context does not make substitution confusing
for the reader
Spirit of wine (esprit du vin) ethyl alcohol
Sulfate of lime (sulfate de chaux) gypsum, or calcium sulfate (CaSO4 ), but calcium
was not isolated until 1808, by Humphrey Davy16
Vegetation (végétation) in the 18th century, this term referred to the active processes
of plant growth and development (in contrast to today’s connotation of inactivity);
the term included such processes as germination, flowering, and fruiting; in this
translation, the term “plant growth” is generally substituted for “vegetation”
Vitiated air (air vicié) corrupted air, that is, air that is depleted of, or reduced in,
oxygen content
Wet, or moist, method (voie humide) in chemical analysis, a method in which
solvents are used; contrasted with the dry method, in which high temperature is
applied instead
Wheat, grain (blé) there is some ambiguity in this French term; it may refer to wheat
(known as Triticum sativum in de Saussure’s day) or more generically to grain; de
Saussure specifically refers to froment as Triticum sativum, however
Wood, fully formed or well formed (bois tout formé, bois bien formé) wood that
is older than sapwood (q.v.)

16
Davy 1808. Electro-chemical researches.
170 Appendices

Appendix 3: Bibliography of References Cited by de Saussure

(as expanded by the translator)

Some of the references in this list are only plausible guesses or provide only a brief mention of the
finding or claim de Saussure attributed to the author. Where he listed only an author’s name, it is
often impossible to know which work he is citing. If the work was published in both English and
French and de Saussure did not explicitly cite the French version, the English one is given.
Page no. Footnote no. References
in Recherches
chimiques sur
la Végétation
v Footnote Berthollet, Claude-Louis. 1803. Essai de statique
chimique. Vol. 1, pp. 513–514. Paris: Chez Firmin
Didot
v Footnote • Giobert, Jean-Antoine. 1793. Des eaux sul-
phureuses et thermales de Vaudier, avec des obser-
vations physiques, économiques et chimiques sur
la vallée de Gesse et des remarques sur l’analyze
des eaux sulphureuses en général. Chapter 4, §. 3,
pp. 64–68. Turin: De l’Imprimerie de Jaques Fea
2 Footnote 1 • Valmont de Bomare, Jacques-Christophe. 1791.
Dictionnaire raisonné universel de l’histoire na-
turelle. Vol. 2, p. 549, footnote (a). Lyon: Chez
Bruyset Frères
4 von Humboldt, Alexander. 1799. Versuche über
die chemische Zerlegung des Luftkreises und über
einige andere Gegenstände der Naturlehre. Chap-
ter 10 (pp. 236–249). Braunschweig: Friedrich
Vieweg
5 Footnote 2 • Sprengel, Kurt. 1802. Anleitung zur Kenntniss
der gewachse, in briefen. Vol. 1, p. 396. Halle:
Karl August Kümmel
7 Footnote 3 • Guyton de Morveau, Louis-Bernard. 1798. Ex-
périences et observations sur le sucre; extrait de
l’ouvrage du docteur Rollo, sur le diabète sucré.
Annales de chimie 25:37–50
8 • de Saussure, Théodore. 1799. Recherches sur
l’influence du gaz oxygène sur la germination des
graines. Journal de physique, de chimie, d’histoire
naturelle et des arts 49:92–98
9 Footnote 4 • Lavoisier, Antoine-Laurent. 1781. Mémoire sur
la formation de l’acide, nommé air fixe ou acide
crayeux, & que je désignerai désormis sous le
nom d’acide du charbon. Mémoires de l’Académie
royale des sciences, 1781, pp. 448–467. (Also in
Oeuvres de Lavoisier (1862) 2: 403–422. Paris:
Imprimerie impériale)

• Indicates de Saussure provided some bibliographic information in addition to an author’s name,

even if sketchy and often with no page numbers, although for some authors (e.g., van Helmont and
Hales), there was only one relevant publication (but two French translations of Hales) to which de
Saussure could have been referring
Appendix 3: Bibliography of References Cited by de Saussure 171

Page no. Footnote no. References

in Recherches
chimiques sur
la Végétation
10 • Guyton de Morveau, Louis-Bernard. 1798. Ex-
périences et observations sur le sucre; extrait de
l’ouvrage du docteur Rollo, sur le diabète sucré.
Annales de chimie 25:37–50
16 • Guyton de Morveau, Louis-Bernard. 1798. Ex-
périences et observations sur le sucre; extrait de
l’ouvrage du docteur Rollo, sur le diabète sucré.
Annales de chimie 25:37–50
21 Senebier, Jean. 1782. Mémoires physico-
chymiques, sur l’influence de la lumière solaire
pour modifier les êtres des trois règnes de la na-
ture, & sur-tout ceux du règne végétal. Vol. 3,
pp. 330–344 (see especially p. 342). Geneva: Chez
Barthelemi Chirol
22 Ingen-Housz, Jan. 1789. Expériences sur les
végétaux, spécialement sur la propriété qu’ils pos-
sèdent à un haut degré, soit d’améliorer l’air quand
ils sont au soleil, soit de le corrompre la nuit, ou
lorsqu’ils sont à l’ombre; auxquelles on a joint une
méthode nouvelle de juger du degré de salubrité
de l’atmosphère. Revised edition. Translated from
English by the author. Vol. 2, p. 447. Paris: Chez
Théophile Barrois le jeune
22 Footnote 7 • Thompson, Benjamin (Comte de Rumford).
1799. Essais politiques, économiques, et
philosophiques. Vol. 2, p. 273. Translated from
English by L. M. D. C. Geneva: Chez G. J. Manger
24 Footnote 9 • Lefebure, E. A. 1801. Expériences sur la ger-
mination des plantes, p. 136. Paris and Stras-
bourg: Chez Levrault, Frères; Strasbourg: De
l’Imprimerie de Louis Eck
28 Footnote 2 • Rückert, Georg Christian Albrecht. 1788.
Beiträge zur Beantwortung der Preisfrage über
den Einfluss der kunstlichen Luftarten auf die
Vegetation. Chemische Annalen 2:394–405 (see
especially p. 399)
28 Senebier, Jean. 1782. Mémoires physico-
chymiques sur l’influence de la lumière solaire
pour modifier des êtres des trois règnes de la na-
ture, & sur-tout ceux du règne végétale. Vol. 1,
p. 112. Geneva: Chez Barthelemi Chirol
29 • Percival, Thomas. 1789. Speculations on the
perceptive power of vegetables. Memoirs of the
Literary and Philosophical Society of Manchester.
Second edition. Vol 2, pp. 130–146
39 Priestley, Joseph. 1772. Observations on differ-
ent kinds of air. Philosophical Transactions of the
Royal Society of London 62:147–264
172 Appendices

Page no. Footnote no. References

in Recherches
chimiques sur
la Végétation
40 Senebier, Jean. This is a general finding, reported
in many of his works, including: 1782. Mémoires
physico-chymiques sur l’influence de la lumière
solaire pour modifier les êtres des trois règnes de
la nature, & sur-tout ceux du règne végétal. 3 vols.
Geneva: Chez Barthelemi Chirol
50 • Hassenfratz, Jean-Henri. 1792. Sur la nutri-
tion des végétaux, Premier Mémoire. Annales de
chimie 13:178–192
53 Footnote 8 • Ingen-Housz, Jan. 1787. Expériences sur les
végétaux, spécialement sur la propriété qu’ils pos-
sèdent à un haut degré, soit d’améliorer l’air quand
ils sont au soleil, soit de le corrompre la nuit, ou
lorsqu’ils sont à l’ombre; auxquelles on a joint une
méthode nouvelle de juger du degré de salubrité
de l’atmosphère. Revised edition. Translated from
English by the author. Vol. 1, pp. 39–43 [not p. 36,
as cited by de Saussure]. Paris: Chez Théophile
Barrois le jeune
53 Footnote 8 • Ingen-Housz, Jan. 1789. Expériences sur les
végétaux, spécialement sur la propriété qu’ils pos-
sèdent à un haut degré, soit d’améliorer l’air quand
ils sont au soleil, soit de le corrompre la nuit, ou
lorsqu’ils sont à l’ombre; auxquelles on a joint une
méthode nouvelle de juger du degré de salubrité
de l’atmosphère. Revised edition. Translated from
English by the author. Vol. 2, pp. 167–170. Paris:
Chez Théophile Barrois le jeune
78 • Berthollet, Claude-Louis. 1802. Recherches
sur les lois de l’affinité. Mémoires de l’Institut
national des sciences et arts. Sciences mathéma-
tiques et physiques. Vol. 3, pp. 1–96
79 Footnote 4 (Plausibly) Dalton, John. 1802. New theory of the consti-
tution of mixed aeriform fluids, and particularly
of the atmosphere. A journal of natural philoso-
phy, chemistry and the arts, by William Nicholson
5:241–244
80–81 Hales, Stephen. 1727. Vegetable staticks: Or, an
account of some statical experiments on the sap in
vegetables: being an essay towards a natural his-
tory of vegetation. (See Chapter 6) London: W. and
J. Innys (Reprinted, 1961; London: Macdonald)
91 Footnote 6 • Ingen-Housz, Jan. 1789. Expériences sur les
végétaux, spécialement sur la propriété qu’ils pos-
sèdent à un haut degré, soit d’améliorer l’air quand
ils sont au soleil, soit de le corrompre la nuit, ou
lorsqu’ils sont à l’ombre; auxquelles on a joint une
méthode nouvelle de juger du degré de salubrité
de l’atmosphere. Revised edition. Translated from
English by the author. Vol. 2, pp. 144–145. Paris:
Chez Théophile Barrois le jeune
Appendix 3: Bibliography of References Cited by de Saussure 173

Page no. Footnote no. References

in Recherches
chimiques sur
la Végétation
104 Hales, Stephen. 1727. Vegetable staticks: or, an
account of some statical experiments on the sap
in vegetables: being an essay towards a natural
history of vegetation. London: W. and J. Innys.
(Reprinted, 1961; London: Macdonald) [Presum-
ably, this refers to Hales’ general finding of air
being taken in through roots and its beneficial ef-
fect on plant growth, since Hales did not know
about individual gases.]
106 • Duhamel du Monceau, Henri-Louis. 1788. La
physique des arbres; où il est traité de l’anatomie
des plantes et de l’économie végétale. Vol. 1, p. 84.
Paris: Chez la Veuve Dèsaint
109 Woodward, John. 1699. Some thoughts and ex-
periments concerning vegetation. Philosophical
Transactions of the Royal Society of London
21:193–227
120 Footnote 9 • Hales, Stephen. 1779. La statique des végétaux,
et celle des animaux; expériences lues a la Société
Royale de Londres. New edition. Translated from
English by Compte de Buffon. Part I. La statique
des végétaux, pp. 130–131. Paris: De l’Imprimerie
de Monsieur
120 Footnote 10 • Hales, Stephen. 1779. La statique des végétaux,
et celle des animaux; expériences lues a la Société
Royale de Londres. New edition. Translated from
English by Compte de Buffon. Part I. La statique
des végétaux, pp. 76–77. Paris: De l’Imprimerie
de Monsieur
125 • Berthollet, Claude-Louis. 1791. Éléments de
l’art de la teinture. Vol. 1, p. 48. Paris: Chez Firmin
Didot
125 • Senebier, Jean. 1800. Physiologie végétale. Vol.
3, pp. 145–147. Geneva: J. J. Paschoud [De Saus-
sure cited p. 144, but discussion appears to start
on p. 145.]
136 Footnote 1 • de Fourcroy, Antoine-François. 1800. Mémoire
sur la coloration des matières végétales par l’air
vital, et sur une nouvelle préparation de couleurs
solides pour la peinture. Annales de chimie 5:80–
91
141 de Fourcroy, Antoine-François. 1789. Mémoire
sur l’existence de la matière albumineuse dans les
végétaux. Annales de chimie 3:252–262
143 Footnote 2 • Lavoisier, Antoine-Laurent. 1789. Traité élé-
mentaire de chimie, présenté dans un ordre nou-
veau et d’après les découvertes modernes. Vol. 1,
pp. 143–148. Paris: Chez Cuchet
174 Appendices

Page no. Footnote no. References

in Recherches
chimiques sur
la Végétation
144 Footnote 3 • Berthollet, Claude-Louis. 1803. Essai de sta-
tique chimique. Vol. 2, p. 525. Paris: de
l’Imprimerie de Demonville et Sœurs, Chez
Firmin Didot
144 Rozier, François (l’abbé). 1786. Cours complet
d’agriculture théorique, pratique, économique, et
de médecine rurale et vétérinaire, suivi d’une
méthode pour étudier l’agriculture par principes;
ou dictionnaire universel d’agriculture. Vol. 4,
p. 525. Paris: Rue et Hôtel Serpente
153 Footnote 6 Priestley, Joseph. 1777. Experiments and observa-
tions on different kinds of air. Vo1. 3, pp. 92–97.
London: Printed for J. Johnson
156 Footnote 7 • Berthollet, Claude-Louis. 1803. Essai de sta-
tique chimique. Vol. 2, p. 546. Paris: de
l’Imprimerie de Demonville et Sœurs, Chez
Firmin Didot
166 Footnote 1 Klaproth, Martin Heinrich. 1802. Beiträge zur
chemischen Kenntniss der Mineralkörper. Vol.
3, possibly Chapter 115 (pp. 319–325). Berlin:
Heinrich August Rottmann
167 (Probably) Vauquelin, Louis-Nicolas. 1798 or 1799. Expéri-
ences sur les sèves des végétaux. Paris: Chez
Quillau, Imprimeur de la Société des Pharmaciens
de Paris
177–179 • de Saussure, Horace-Bénédict. 1796. Voyages
dans les Alpes, précédés d’un essai sur l’histoire
naturelle des environs de Genève. Vol. 5, § 1319,
pp. 206–209. Neuchatel: Chez Louis Fauche-
Borel
189 #17, Table of Kirwan, Richard
Carbonizations
191 #32, Table of Klaproth, Martin Heinrich
Carbonizations
206 Footnote 3 • Priestley, Joseph. 1790. Experiments and ob-
servations on different kinds of air, and other
branches of natural philosophy, connected with the
subject. Vol. 3, pp. 331–333. Birmingham: Printed
by Thomas Pearson
206 Footnote 4 • Ingen-Housz, Jan. 1789. Expériences sur les
végétaux, spécialement sur la propriété qu’ils pos-
sèdent à un haut degré, soit d’améliorer l’air quand
ils sont au soleil, soit de le corrompre la nuit, ou
lorsqu’ils sont à l’ombre; auxquelles on a joint une
méthode nouvelle de juger du degré de salubrité
de l’atmosphère. Revised edition. Translated from
English by the author. Vol. 2, pp. 146–151. Paris:
Chez Théophile Barrois le jeune
Appendix 3: Bibliography of References Cited by de Saussure 175

Page no. Footnote no. References

in Recherches
chimiques sur
la Végétation
206 Footnote 5 • Priestley, Joseph. 1790. Experiments and ob-
servations on different kinds of air, and other
branches of natural philosophy, connected with
the subject. Vol. 3, p. 334. Birmingham: Printed
by Thomas Pearson
207 (Plausibly) Senebier, Jean. 1793. Mémoire sur divers
phénomènes produits par des feuilles de plantes
exposées sous l’eau à l’action de la pompe pneu-
matique. Mémoires de l’Académie royale des
sciences, 1790–1791 (Turin) 5:36–56 [In this pa-
per, Senebier speculated that nitrogen is not taken
into the plant from the air; later he stated that
Spallanzani proved this: Senebier, Jean. 1800.
Physiologie végétale. Vol. 3, p. 159. Paris: Chez
Fuchs]
207 Woodhouse, James. 1802. Experiments and ob-
servations on the vegetation of plants, which
shew that the common opinion of the amelio-
ration of the atmosphere, by vegetation in solar
light, is ill founded. A Journal of natural philoso-
phy, chemistry and the arts, by William Nicholson
2:150–162
208 Berthollet, Claude-Louis. 1803. Essai de sta-
tique chimique. Vol. 2, pp. 61–93. Paris: de
l’Imprimerie de Demonville et Soeurs, Chez
Firmin Didot
209 Huber, François, and Senebier, Jean. 1801. Mé-
moires sur l’influence de l’air et de diverses
substances gazeuses dans la germination de dif-
férentes graines, pp. 160–173 (see especially
pp. 167–168). Geneva: Chez J. J. Paschoud
212 Footnote 8 Guyton de Morveau, Louis-Bernard; Chaussier,
François; and Bertrand (abbé). 1784. Description
de l’aérostate, l’Académie de Dijon, contenant
le détail des procèdes, la théorie des opérations,
les dessins des machines & les procès-verbaux
d’expériences, pp. 41–42. Dijon: Chez Causse;
Paris: Chez Théophile Barrois
217 • Senebier, Jean. 1800. Physiologie végétale. Vol.
3, pp. 228 ff. Geneva: J. J. Paschoud
218 Senebier, Jean. 1800. Physiologie végétale. Vol.
3, pp. 388 ff. Geneva: J. J. Paschoud
176 Appendices

Page no. Footnote no. References

in Recherches
chimiques sur
la Végétation
218 Ingen-Housz, Jan. 1789. Expériences sur les
végétaux, spécialement sur la propriété qu’ils pos-
sèdent à un haut degré, soit d’améliorer l’air quand
ils sont au soleil, soit de le corrompre la nuit, ou
lorsqu’ils sont à l’ombre; auxquelles on a joint une
méthode nouvelle de juger du degré de salubrité
de l’atmosphère. Revised edition. Translated from
English by the author. Vol. 2, p. 440. Paris: Chez
Théophile Barrois le jeune
218 Footnote 1 • Spallanzani, Lazaro. 1799. Lettre du Professeur
Spallanzani, au célèbre chimiste Giobert, sur les
plantes renfermées dans des vases remplis d’eau et
d’air, et exposées à la lumière immédiate du soleil,
ou à l’ombre. Journal de physique, de chimie,
d’histoire naturelle et des arts 48:135–141
219 Senebier, Jean. 1800. Physiologie végétale. Vol.
3, pp. 227 ff. Geneva: J. J. Paschoud
219 Berthollet, Claude-Louis. 1786. De l’influence
de la lumière. Observations sur la physique, sur
l’histoire naturelle et sur les arts 29: 81–85.
234 Footnote 5 de Candolle, Augustin-Pyramus. 1801. Extrait
d’un mémoire sur les pores de l’écorce des feuilles.
Journal de physique, de chimie, d’histoire na-
turelle et des arts 52:130–134 (see especially
p. 131) [Summarizes de Candolle’s research on
stomata, including mention of his finding that suc-
culents have fewer of these pores than do other
plants, but this is not the original paper]
240–241 Footnote 1 Bonnet, Charles. 1754. Recherches sur l’usage des
feuilles dans les plantes. See especially pp. 260–
262; related discussions on various other pages.
Göttingen and Leiden: Chez Elie Luzac, fils
241 Tull, Jethro. 1731. The new horse-houghing hus-
bandry: or, an essay on the principles of tillage
and vegetation, wherein is shewn, a method of
introducing a sort of vineyard-culture into the
cornfields, in order to increase their product and di-
minish the common expence, by the use of instru-
ments lately invented. Dublin: George Grierson.
[Many later editions]
241 Text and foot- van Helmont, Jan. 1648. Ortus medicinae. Edited
note 2 by F. W. van Helmont. Amsterdam: Apud Lu-
dovicum Elzevirium. (English translation, 1662,
by J. Chandler, as Oriatrike, or, Physick refined.
The common errors therein refuted, and the whole
art reformed & rectified: being a new rise and
progress of phylosophy and medicine for the de-
struction of diseases and prolongation of life.
London: Printed for Lodowick Loyd; reprinted,
1966, Culture et Civilisation, Brussels)
Appendix 3: Bibliography of References Cited by de Saussure 177

Page no. Footnote no. References

in Recherches
chimiques sur
la Végétation
241 Footnote 2 Tillet, Mathieu. 1772. Expériences & obser-
vations sur la végétation du blé dans chacune
des matières simples dont les terres labourables
sont ordinairement composées, & dans différens
mélanges de ces matières, par lesquels on s’est rap-
proché de ceux qui constituent ces mêmes terres
à labour. Histoire de l’Académie royale des sci-
ences, 1772, avec les mémoires de mathématique
& de physique, pour la même année, pp. 99–156.
Paris: de l’Imprimerie Royale, 1775
241 Footnote 2 Bonnet, Charles. 1779. Expériences sur la végé-
tation des plantes dans d’autres matières que la
terre, & principalement dans la mousse. Oeuvres
d’histoire naturel et de philosophie de Charles
Bonnet. Vol. 3. Two memoirs, pp. 203–262.
Neuchatel: Chez Samuel Fauche
241 Footnote 2 Duhamel du Monceau, Henri-Louis. 1748. Sur
les plantes qu’on peut élever dans l’eau. Histoire
de l’Académie royale des sciences, 1748, avec
les mémoires de mathématique & de physique,
pour la même année, pp. 272–301 (see espe-
cially pp. 275–276, regarding oak). Paris: de
l’Imprimerie Royale, 1752
241 Footnote 2 Bergman, Torbern Olaf. 1788. Opuscula physica
et chemica. Vol. 5, p. 143. Lipsiae [Leipzig]: In
officina libr. Jo. Godofr. Mülleriana
241 Footnote 2 Kirwan, Richard. 1802. The manures most advan-
tageously applicable to the various sorts of soils,
and the causes of their beneficial effect in each par-
ticular instance. 5th edition. See pp. 39, 58, 66–67.
London: Printed for Vernor and Hood, Poultry
241 Footnote 2 Hassenfratz, Jean-Henri. 1792. Sur la nutrition des
végétaux, Premier Mémoire. Annales de chimie
13:178–192
245 (Possibly Giobert, Giovanni Antonio. 1790. Ricerche
but not chimiche intorno agli ingrassi ed ai terreni. Mem-
seen) orie della Reale Società Agraria di Torino, dalla
sua instituzione (1785) al 1790. Vol. 5:1–347; Vol.
6:1–216
245 Hassenfratz, Jean-Henri. 1792. Sur la nutrition des
végétaux, Premier Mémoire. Annales de chimie
13:178–192
247 Footnote 3 • Dalrymple, William. 1800. On the culture of
wheat [De la culture du blé]. Bibliothèque Britan-
nique, Agriculture Anglaise 5:485–502; 503–535.
255 Footnote 7, (Plausibly) Berthollet, Claude-Louis. 1802. Recherches sur
in table les lois de l’affinité. Mémoires de l’Institut na-
tional des sciences et arts. Sciences mathéma-
tiques et physiques 3:1–96
178 Appendices

Page no. Footnote no. References

in Recherches
chimiques sur
la Végétation
258 Footnote 11 • Leblanc, Nicolas. 1802. De la cristallotechnie,
ou essai sur les phénomènes de la cristallisation.
Journal de physique, de chimie, d’histoire na-
turelle et des arts 55:300–313. [Page numbering is
awry: pp. 301–308 are misnumbered as pp. 297–
304; and according to the Table of Contents, the
article begins on p. 308]
262 (Plausibly) Duhamel du Monceau, Henri-Louis. 1767. Ob-
servations sur les sels qu’on retire des cendres
des végétaux. Mémoires de l’Académie des sci-
ences, 1767, avec les mémoires de mathématique
et de physique, pour la même année, pp. 233–239;
Suite des expériences sur les sels qu’on peut retirer
des lessives du kali. Ibid., pp. 239–240 [Discusses
the difference in the kind of alkali found in the
ash of seaside plants when they are grown inland,
but does not address de Saussure’s statement that
seaside plants languish far from the seaside]
263 Thouvenel, Pierre
263 Cornette, Claude-Melchior
263 • de Bullion, Marquis. 1791. Mémoire sur les
moyens d’obtenir le nitre ou salpêtre des terres
qui le contiennent, de le purifier & d’en fabri-
quer facilement de la poudre à canon. Mémoires
d’agriculture, d’économie rurale et domestique,
publiées par la Société royale d’agriculture (winter
trimester), pp. 38–59
266 Footnote 13 • Haüy, René-Just. 1801. Traité de minéralogie.
Vol. 1, p. 7. Paris: de l’Imprimerie de Delance.
Chez Louis
267 Footnote 14 van Helmont, Jan. 1648. Ortus medicinae. Edited
by F. W. van Helmont. Amsterdam: Apud Lu-
dovicum Elzevirium. (English translation, 1662,
by J. Chandler, as Oriatrike, or, Physick refined.
The common errors therein refuted, and the whole
art reformed & rectified: being a new rise and
progress of phylosophy and medicine for the de-
struction of diseases and prolongation of life.
London: Printed for Lodowick Loyd; reprinted,
1966, Culture et Civilisation, Brussels)
267 Footnote 14 • Kirwan, Richard. 1802. The manures most ad-
vantageously applicable to the various sorts of
soils, and the causes of their beneficial effect in
each particular instance. 5th edition, pp. 28–29.
London: Printed for Vernor and Hood, Poultry
Appendix 3: Bibliography of References Cited by de Saussure 179

Page no. Footnote no. References

in Recherches
chimiques sur
la Végétation
268 Footnote 15 Hales, Stephen. 1727. Vegetable staticks: or, an
account of some statical experiments on the sap in
vegetables: being an essay towards a natural his-
tory of vegetation, p. 8. London: Printed for W. and
J. Innys. (p. 3, 1961 reprint edition; London: Mac-
Donald) [Hales actually said the exact opposite of
what de Saussure claimed he said: Hales said 43 of
the sunflower’s weight, not 41 , was water.]
273 Footnote 1 • Kirwan, Richard. 1802. The manures most ad-
vantageously applicable to the various sorts of
soils, and the causes of their beneficial effect
in each particular instance. 5th edition, p. 41.
London: Printed for Vernor and Hood, Poultry
273 Footnote 1 • Rückert, Georg Christian Albrecht. 1789–1790.
Der Feldbau: chemisch untersucht um ihn zu
seiner letzten Vollkommenheit zu erheben. 3 vols.
Erlangen: Johann Jacob Palm.
273 • Perthuis de Laillevault, Léon de. 1797. Mémoire
sur les moyens de multiplier la fabrication de la
potasse en France. Annales de chimie 19:157–193
275 Footnote 2 • Hales, Stephen. 1779. La statique des végétaux,
et celle des animaux; expériences lues a la Société
royale de Londres. New edition. Translated from
English by Compte de Buffon. Part I. La statique
des végétaux. Paris: De l’Imprimerie de Monsieur
[De Saussure cited pp. 3 and 43; p. 43 does not
seem to be relevant, nor does p. 43 of the 1735
French edition of this book.]
275 Footnote 3 • Bonnet, Charles. 1779. Oeuvres d’histoire na-
turelle et de philosophie. Octavo edition. Vol.
4. Recherches sur l’usage des feuilles, p. 77.
Neuchatel: Chez Samuel Fauche
276 Hales, Stephen. 1779. La statique des végétaux,
et celle des animaux; expériences lues a la So-
ciété royale de Londres. New edition. Translated
from English by Compte de Buffon. Part I. La sta-
tique des végétaux, p. 17. Paris: De l’Imprimerie
de Monsieur
277 Hales, Stephen. 1727. Vegetable staticks: or, an
account of some statical experiments on the sap
in vegetables: being an essay towards a natural
history of vegetation. See Experiment 30, pp. 92–
93. London: W. and J. Innys (p. 52, 1961 reprint
edition; London: Macdonald)
180 Appendices

Page no. Footnote no. References

in Recherches
chimiques sur
la Végétation
283 • Lampadius, Wilhelm August. 1801. Extrait d’un
Mémoire de Lampadius, Professeur de Chimie à
l’École des Mines de Freyberg, sur la formation
et la nature des terres. Journal des mines 55:525–
528 [The article is actually in No. 55, not No. 65
as cited by de Saussure; there was no paper by
Lampadius listed in No. 65]
289–291 • Vauquelin, Louis-Nicolas. 1799. Expériences
sur les excrémens des poules, comparés à la
nourriture qu’elles prennent, et réflexions sur la
formation de la coquille de l’œuf. Annales de
chimie 29:3–26 (see pp. 16–19)
296 Footnote 5 • Marggraf, Andreas Sigismund. 1762. Opuscules
chymiques. Vol. 1. (See pp. 68–70.) Paris: Chez
Vincent, Imprimeur-Libraire
296 (Possibly) Pott, Johann Heinrich. 1759. Dissertations
chymiques. 4 vols. Collected and translated from
Latin and German by J.-F. Demachy. Paris: Chez
Jean-Thomas Herissant [In the Marggraf reference
cited on this same page, de Saussure mentioned
Pott; de Saussure may have gotten this reference
directly from Marggraf]
296 Vauquelin, Louis-Nicolas
296 (Possibly) de Fourcroy, Antoine-François, and Vauquelin,
Louis-Nicolas. 1806. Mémoire pour server à
l’histoire chimique de la germination et de la
fermentation des graines et des farines. Annales
du Muséum d’histoire naturelle 7:1–18 [This is
the first publication of research undertaken by
Vauquelin and de Fourcroy starting in 1800; al-
though this paper appeared after de Saussure’s
book was published, he could have heard about
early stages of the work, especially because he
took courses from Vauquelin in 1800]
302 • Senebier, Jean. 1800. Physiologie végétale. Vol.
4, p. 87. Geneva: J. J. Paschoud
308 Footnote 7 Karsten, Diebrich Ludwig Gustav
308 Footnote 7 • Westrumb, Johann Friedrich. 1788. Kleine
Physikalisch-chemische Abhandlungen, aus den
chemischen Journalen gesammelt, und mit eini-
gen Verbesserungen und Anmerkungen. Part 2.
Leipzig: In der J.G. Müllerschen buchhandlung
311 Perthuis de Laillevault, Léon de. 1797. Mémoire
sur les moyens de multiplier la fabrication de la
potasse en France. Annales de chimie 19:157–193
(see p. 174)
Appendix 3: Bibliography of References Cited by de Saussure 181

Page no. Footnote no. References

in Recherches
chimiques sur
la Végétation
315 • de Saussure, Théodore. 1800. De l’influence du
sol sur quelques parties constituantes des végé-
taux. Journal de physique, de chimie, d’histoire
naturelle et des arts 51:9–40
316 (Possibly) de Fourcroy, Antoine-François, and Vauquelin,
Louis-Nicolas. 1806. Mémoire pour server à
l’histoire chimique de la germination et de la
fermentation des graines et des farines. Annales
du Muséum d’histoire naturelle 7:1–18 [This is
the first publication of research undertaken by
Vauquelin and de Fourcroy starting in 1800; al-
though this paper appeared after de Saussure’s
book was published, he could have heard about
early stages of the work, especially because he
took courses from Vauquelin in 1800]
322 (Possibly) Klaproth, Martin Heinrich. 1795–1802. Beiträge
zur chemischen Kenntniss der Mineralkörper.
Vols. 1–3. Posen: Decker
323 Vauquelin, Louis-Nicolas. 1798 or 1799. Expéri-
ences sur les sèves des végétaux. Paris: Chez
Quillau, Imprimeur de la Société des Pharmaciens
de Paris
324 • de Fourcroy, Antoine-François, and Vauquelin,
Louis-Nicholas. 1803. Mémoire sur la présence
d’un nouveau sel phosphorique terreux dans les
os des animaux, et sur l’analyse de ces organes
en général. Annales de chimie 47:244–261. (See
especially pp. 248–249)
324 (Plausibly) Berthollet, Claude-Louis. 1802. Recherches sur
les lois de l’affinité. Mémoires de l’Institut na-
tional des sciences et arts. Sciences mathéma-
tiques et physiques 3:1–96
Index

A Ash, xxix, 57, 58, 63–68, 71, 98, 104–108, 110,

Absorption of solutions, 91, 158 114–120, 157, 159
Academy of Geneva, xxxiii, xxxiv, xxxviii Ash of plants, x, 103, 113, 159
Acetate, 118, 119 Assimilation, 3, 56, 84n2, 86, 89, 157, 158
Acetic acid, 55, 56, 61, 63–65, 119n9, 156, Atmosphere, ix, xiii, xx, xxi, xxiii–xxvi, xxix,
163, 165, 168 xxxi, xxxv, 7, 12, 15, 17, 20, 21, 25, 27–30,
Acetification, 55 32, 33, 35, 37, 38, 41, 43–45, 51, 54, 56, 59,
Acidulated water, 12 60, 69, 75, 77, 79, 82, 85, 87n3, 89, 92, 102,
Agriculture, x, xiv, xxx, xxx(n28), 93n3 113, 115, 155, 158, 159
Agronomy, xiii, xxx, xxx(n28), xxxviii Atmospheric air, 4, 6, 11–14, 14n4, 17–19, 20,
Air, ix, xiv, xix, xxiii, 2, 2n2, 15, 21, 26, 28, 31, 22, 23, 25, 26, 28–36, 38, 41, 43–47, 50, 51,
32, 35, 37, 42, 45, 48, 60, 61, 64, 66, 69, 70, 53, 56, 57, 60, 68, 69, 76, 79, 81n8, 83–85,
74, 76, 77, 79, 79n5, 80–82, 87n3, 99, 116, 87–89, 91, 92
157 Atriplex hortensis, xxix, 22
Air bubble, 47
Air pump, 28, 29, 32, 81, 82, 156 B
Balance-sheet approach, xxii, xxix
Alcohol, xviii, xxxviii, 48, 56, 65, 67, 67n4,
Berthollet, Claude-Louis, xx, xxvii, xxxi, 32,
97, 100n12, 118, 118n9, 119
55, 60, 80, 83, 120, 170, 172–177, 181
Alcohol-insoluble salts, 119
Berzelius, Jöns Jacob, xxvii(n20), xxviii
Alkali, 22, 98, 115, 158
Bituminous empyreumatic oil, 63, 64
Alkaline salts, 67, 68, 103, 105, 107–114, 117,
Black, Joseph, xvii, xxxvi
118, 125, 129, 139, 145, 149, 151, 153, 159
Blondel-Mégrelis, Marika, xxx(n28),
Alpine expeditions, xxxiv, xxxv
xxxiv–xxxvi, xxxviii, xxxix
Alps, xxxii, xxxv, xxxvi, 106 Boiled water, 1, 2, 5, 88
Alternating day/night regime, xxiv Bonnet, Charles, xix, xxxiv, xxxvi, 91n1, 91n2,
Alumina, xxvi(n19), xxxviii, 79n6, 106, 117, 104, 176, 177, 179
159, 163 Botany, xxx, xxxiii, xxxvi
Aluminum, xxxv, xxxviii, 79n6 Boussingault, Jean-Baptiste, xiv, xxix,
Ammonia, xxvi, 55, 60, 63–65, 67, 79, 79n6, xxix(n26), xxx
107n4, 109, 115n7, 117, 118, 120, 121 Boyle’s Law, xxxv
Analyses of plant ash, xxv
Analyzing ash, 116 C
Ancien régime, xlix Cactus, xxvi, 19, 20, 22, 25–36, 55, 76, 81, 82,
Annual plants, 90, 92, 93, 101, 105, 105, 108, 87–89, 156, 158
110, 113, 115 Calcareous humus, 65
Antiseptic, 6n5, 68, 70, 157 Calcareous sand, 70, 106
Aristotelian elements, xvi, xvii Calcination, xviii
Aristotle, xiv Calcium, xxv(n18), xlx

J. F. Hill, Chemical Research on Plant Growth, 183

DOI 10.1007/978-1-4614-4136-6, © Springer Science+Business Media New York 2013
184 Index

Calcium carbonate, 27, 36, see also Carbonate D

of lime Dalton, John, xxxii, 32n4, 172
Calendar system, xlix Darkness, 8, 21, 22, 25, 26n1, 27–31, 37, 43,
Caloric, 51, 59n6, 80, 163, 164 45, 48–50, 76, 77, 84, 89, 90, 156
Candaux, Jean-Daniel, xxxiv, xxxvi, xxxviii, Davy, Humphrey, xiv, xxviii, 163n2, 164n3,
xxxix 166, 167n11
Carbohydrate, xxv, see also Sugar Daytime, 33–35, 43, 46, 156
Carbon, xiv, xx, xx(n15), xxi, xxiv, xxv(n18), de Candolle, Augustin Pyrame, xxviii, xxiv,
xxxvii, 3, 4, 6, 7, 38, 50, 56–58, 61, 67, 69, xxx(n28), xxxiv, 176
70, 74, 155–157 de Saussure, Albertine-Amélie Boissier, xxxiii
Carbonated lime, 105, 110–112, 117, 118 de Saussure, Horace-Bénédict, xxii, xxxiii,
xxxiii(n32), xxxiv–xxxvi, xxxvi(n34)
Carbonate of ammonia, 64, 67, 118
de Saussure, Nicolas, xxxiii, xxxvi
Carbonate of lime, 27, 36, 110–113, 116, 120,
de Saussure’s archives, xxxix
121
Dead wood, 57
Carbon dioxide gas, xxiii, xxiv, xxvi, 3–5, 7,
Debarked wood, 48
11, 13–16, 18–23, 27–33, 35–38, 43–61, 68,
Decay, 1, 3, 4, 42
77–83, 85–90, 92, 101, 155–158
Decayed wood, 103
Carbon dioxide metabolism, xxxvii
Deciduous tree, 38, 104, 159
Carbonic acid gas, 3, see also Carbon dioxide Decoction, 58, 66, 67, 69, 118, 121, 164
gas Decomposition, xv, xxi–xxiv, xxvi, 5, 8n8, 16,
Carbonization, 7, 17, 18, 20, 21, 57, 57n5, 20–22, 31, 33–35, 64, 68, 76, 77, 81, 83,
67n3, 70, 71, 156 86–90, 97, 111, 119, 120, 157, 158
Carbon monoxide, xxiv, 80, 80n7, 81, 82, 157 Decomposition of carbon dioxide, xxiv, 8n8,
Cavendish, Henry, xvii, xxxvi 16, 21, 22, 86, 89, 157
Charcoal, 4n4, 7, 17, 18, 20, 21, 55, 57, 58, Decomposition of water, xxi, xxiii, 5, 34, 35,
63–66, 70, 71 83, 84, 88, 90, 157
Chemical elements, xvii, xviii Dephlogisticated air, xviii, xix, xxi
Chemical equation for photosynthesis, xxvii, Desiccation, 69n6
186 Developed plant, 11, 25, 156
Chemical revolution, xv(n2), xvi Dew, 105, 108
Chemical terminology, xviii, xlx Discovery of oxygen, xviii
Chemistry, xvi–xviii, xxii, xxx, xxxiii, xxxvi, Distilled water, xvi, xxiii, 6, 12, 12n2, 16, 20,
xxxviii 21, 26n1, 31, 34, 41–43, 50, 58, 66, 68, 84,
Chlorine, xxv, 2 88, 90, 91n1, 94, 98, 105, 106, 113–115
Chlorophyll, 22 Dry method, 119, 169
Clay, 38, 69, 71 Dry plant matter, 21, 85, 86
Dry plant substance, 20, 84
Climate, 68, 104
Dry weight, xxiii, 7, 17, 57, 84, 86, 101, 155,
Closed vessel, 6, 7, 17, 20, 21, 84, 85, 88, 164
158
Col du Géant, xxxv Drying oven, 6, 7
Combustion, xviii, 4, 4n4, 16, 49, 66, 67, 98,
105, 116
E
Combustion of seeds, 116
Earths, 69, 91, 92, 106, 107, 112, 113, 117, 118
Compression, 5, 31, 32
Earthy carbonates, 117
Cork, 74, 108 Earthy phosphates, 105, 109, 110, 112, 117,
Corpuscles in the atmosphere, 92, 115 118
Corrupted air, 16 Earthy salt, 110
Cotyledons, 2–4, 13n3, 37 Energy concept, xxvii, 51
Crassulacean acid metabolism (CAM), xxvi, Energy of sunlight, xiv
23, 33, 35, 36 Epidermis, 15n4, 36, 74, 81, 88, 88n5, 90, 108,
Creative force, x, 107 158
Crude acetic acid, 63, 64 Etiolation, 8, 84
Index 185

Eudiometer, ix, xxxiv, 16, 17, 19, 20, 33, 37n7, German translation, xxviii, xxx
77, 88, 165 Germinating seeds, xxiv, 4, 11, 113, 155, 156
Eudiometry, xviii Germination, xxv, xxxvii, xxxviii, 1–9, 11, 23,
Evaporation, 65, 94n4, 104, 109, 114–118, 59, 75, 81, 83, 110, 112, 155
118n8 Glandular pores, 2
Evergreen, 38, 92 Gluten, 60, 61
Evergreen tree, 38, 104, 159 Grandeau, Louis, xxviii
Excision of roots, 97 Granitic sand, 106
Exhalation, see Expiration Grasses, 112, 113, 117
Expiration, 26, 31, 33–36, 45, 47, 50, 51 Gravel, 63, 106, 114n6
Extract, xxiii, xxv–xxvii, 6, 7, 53–55, 58, 61, Green coloring matter, 92
71, 73, 79, 94, 110, 156, 159 Green fruit, xxxviii, 28n2, 50
Extractive juices, 50, 51, 54, 55, 59, 70, 74, Green parts of plants, 16, 31, 51, 156
101n14, 157 Green state, 84, 98, 101n14, 105
Extractive liquids, 54 Grew, Nehemiah, xvi
Extractive liquor, 55 Growth force, 26, 26n1, 32, 36, 43, 44, 105,
Extractive solution, 53, 70, 102 108
Gum, 94, 94n4, 95, 97, 98, 116, 158
F
Fabri, Renée, xxxvi H
Facsimile editions, xiv Hales, Stephen, vii, xvi, 32, 41, 46, 101,
Farming operations, 93 101n15, 104, 172, 173, 179
Fermentation, xv, xxv, 6, 53–55, 59–61, 68, 80, Hard wood, 109
86, 89, 94, 115n7, 156, 157 Heat, xviii, xxiv, 7, 8, 8n8, 9, 14, 21, 28, 50,
Fern, 92 58, 60, 65, 71, 80, 86, 93, 98, 116–121
Fertilizer, xv, xxix, xxix(n24) Herbaceous plant, 26, 38, 103, 104, 107,
Field trials, xxix 110–112, 159
Filter, 91, 91n1, 98, 109, 117, 119–121 History of chemistry, xiv, xv(n3)
Fire, xiv, 17, 54, 55, 70, 71, 100n12, 117–119, Humidity, 67n4, 99, 100n12
121 Humus, ix, xiii, xv, xxiii, xxv, xxviii–xxx,
Fixation of water, 85, 86 xxxvii, 11, 12n2, 13, 41, 44, 50, 51, 63–70,
Fixed air, xvii–xxi 100, 106, 165
Fleshy plants, 22, 37 Humus extract, xxiii, 67, 94, 100, 101, 110
Flower, 48–50, 73, 78, 106, 114, 130–132, 136, Humus theory, xv, xxviii, xxx, xxxii
138, 139, 142, 143, 165 Hydrogen, xvii, xx, xxv(n18), 4, 4n4, 5, 6n5, 7,
Flowering, 21, 40, 73, 106, 110, 112, 114, 8, 28–31, 35, 45, 48–50, 54–57, 60, 69–71,
132–134, 136, 138–140, 143, 169 80, 80n7, 81, 84–86, 88, 90, 100, 155–157
Fluidity, 98 Hygrometer, xxxiv, 57, 69, 69n5, 84
Formation of water, 48, 59
Fourcroy, Antoine-François, xxxi, 53, 55, 111, I
118, 120, 173, 180, 181 Imbibition, 1, 26
French Revolution, xxvii(n22), xxxiii, xxxv, Incineration, xxv, xxxii, 69, 91, 98, 100,
xlix, l 103–105, 110–112, 115–118, 157, 159
Fresh weight, 21, 84, 101 Inflammable gas, 60
Frontenex, xxxvi Inflammable principle, xviii
Fruit, xxxviii, 22, 28n2, 50, 104, 110, 112, 132, Ingen-Housz, Jan, xix–xxi, xxiii, xxiv, xxvi,
133, 159 xxxii, xxxvii, 8, 21, 36n6, 79, 83, 171, 172,
Fully formed wood, 58, 108 174, 176
Inhalation, see Inspiration
G Inspiration, 26–31, 33–38, 43, 45, 47, 50
Gaseous state, xvi, 37, 56n4, 89 Institute of France, xxvii, xxvii(n22), xxxviii
Gassed water, 12, 12n1 Integument, 2, 109
Geneva, Switzerland, xi, xiii Iron, xxv(n18), 64, 69, 71, 78, 100, 105, 106,
Geology, xxxiii, xxxiv, xxxviii 113, 116, 166
186 Index

Iron oxide, 106 Metric system, xlix, 4, 161

Iron sulfide, 78 Middle Ages, xiv
Mineral concentrations in plants, xxv
L Mineral nutrients, xxviii
Lavoisier, Antoine-Laurent, xvii–xxii, xxxi, Mineral substances, xxv, 99, 100
xxxii, xxxviii, 4, 7, 54, 55n2, 69, 86, 170, Minerals in plant ash, xiii, xxxi
173 Mont Blanc, xxxv
Le Bréven, 106 Monte Rosa, xxxv
Leaching, xxvi, 109, 159 Moss, 92
Leaf bud, 75, 82 Mucilage, xx, 6, 7
Leaf, leaves, xix, xxiv, 15, 22, 23, 25, 27, 28, Mucilaginous seeds, 6, 59
31, 32, 38, 44, 47, 48, 51, 78, 83, 92, 93,
100n12, 104, 105, 108, 110, 115, 157–159 N
Library of Geneva, xxxix Napoleonic Empire, xxxiii(n31), xlix
Light, xix, xxiii, xxvii, xxxi, 8, 11, 13, 17, Natural philosophers, xvi, xxxiv, 8, 46
20–23, 31, 45, 48, 65, 77, 77n1, 115n7, 118, Nitric acid, 2, 37, 109, 117–121
155, 156 Nitric oxide, xvii, xvii(n9), xxxv, 77, 78, 81
Lime, xv, xxix, 14–16, 22, 51, 78, 79, 100, 105, Nitrogen dioxide, xvii(n9)
106, 110–121, 159, 166 Nitrogen gas, ix(n1), xxi, xxiii, xxiv, xxvi,
Limewater, xxxv, 14–19, 27, 31, 36, 58, 65, 67, xxxi, 4, 6, 11, 14, 14n4, 17–20, 22, 23,
83, 84, 89, 120, 121, 166 26n1, 27–29, 33–35, 37, 37n7, 41, 44–49,
Linseed oil, 59 55, 59, 59n6, 60, 64, 69, 75–82, 86, 88, 90,
Liquid, xx, 2, 4, 12, 13n3, 17, 33, 50, 53, 54, 92, 155, 156
60, 65, 70, 77, 86, 88–91, 96–98, 108, 110, Nitrous air test, xvii, xvii(n9), 77
114, 116, 120, 121 Non-green parts of plants, xix, xxxii, 166
Liquidity, 53
Nutrients, xiv, xvi, xx, xx(n15), xxv(n18),
Liquor, 55, 56, 91n1, 97n6, 109, 117, 118, 120
xxviii, 13, 92, 93, 95, 100–102
Lute, 15, 44, 45, 47, 63, 64, 67, 71, 166
Luting, 45
Lye, 67, 116, 118, 118n8, 120 O
Oil, xviii, 2, 6n5, 12, 49, 59, 59n6, 63, 64, 156
M Ordinary air, xxiii, xxxiv, 3–5, 12–19, 29,
Macaire-Prinsep, Jean-François, xxx 33–37, 37n7, 41, 43–45, 48–50, 54, 57, 60,
Maceration, 58, 66, 67 75–80, 82–87, 87n3, 89, 90, 157
Magiels, Geerdt, xxxii Oxide of manganese, 2
Magnesia, 100n12, 105, 106, 110–112, 117, Oxygen chemistry, xviii, xviii(n10)
118, 118n8, 120 Oxygen gas, ix(n1), xiv, xx, xxiii–xxv, 1, 3, 4,
Magnesium, xxv(n18), 111 4n4, 5–8, 13–23, 25–61, 68–70, 75–90, 101,
Malpighi, Marcello, xvi 155
Manganese, xxv(n18), 2, 69, 100, 105, 113, Oxygenated muriatic acid, 2, 166
166 Oxygen-free environments, 6, 57, 75, 82, 156
Manure, xv, xxix, 42, 63, 65, 65(n2), 68, 91, Overall equation for photosynthesis, xiv, xxvii
93, 100n12, 101, 106, 107
Marine plant, 99 P
Marsh plant, 14n4, 22, 23, 38, 77–80, 94 Parenchyma, xix, xxvi, 15n4, 31, 36, 48, 82,
Mature plant, xxii, 11 83, 88, 89
Mephitic air, xviii Parisian units of measurement, xxxi
Mercury, xvii, 1–3, 5, 6, 6n5, 16, 26, 29, 38, 41, Peat, 64
43–47, 53, 54, 56, 57, 60, 68, 81n8, 87, 88 Pellicle, 53–55, 166
Mesophyll, 22 Perennial, 92, 93
Metallic oxide, 2, 110, 113, 117, 123 Petal, 22, 49, 51, 76
Metals, xiv, xviii, xxxv, 91, 113 Philomatic Society, xxxviii, xxxviii(n35)
Methane, 6, 63, 64, 155 Phloem, xxxviii, 74
Methodological limitations, xxxi Phlogisticated air, xviii, xix, xxi
Index 187

Phlogiston, xv(n2), xviii, xix, xxi, xxi(n16), Putrefaction, 7, 8n6, 43, 60, 61, 68, 99, 105,
xxii 156, 157, 163, 165, 168
Phlogiston theory, xv(n2), xviii, xviii(n10), xxii Pyrolignate of ammonia, 63, 64
Phosphate of lime, 99, 115, 118–121 Pyroligneous acid, 63, 168
Phosphate of potash, 110–112, 115, 117–121
Phosphate of potash and lime or magnesia, 118, Q
118n8, 120, 121 Quicklime, 4, 22, 35, 50, 78, 155, 166, 168
Phosphated potash of lime, 120, 121
Phosphoeudiometer, 16 R
Phosphoric acid, 111, 119, 120 Radicle, 1, 2, 4, 5, 12, 75, 81, 82
Phosphorus, xxv(n18), xxvi, 111 Rain water, 65, 92, 93, 101, 101n14, 104–106,
Photosynthesis, xiii, xiv, xiv(n1), xv, xv(n3), 108, 109
xvi, xvii, xix, xix(n12), xx, xxi, xxiii, xxvii, Réaumur temperature scale, 2
xxxi, xxxiv, xxxvi, 22 Receptacle, 4, 8, 11–22, 26, 30, 32, 35, 36, 38,
Photosynthesis pioneers, xv, xxxiv, xxxvi 41, 44, 48, 49, 54, 55, 57, 69, 77, 78, 81,
Physiology, xv, xvi, xviii, xviii(n11), xxvi, xxx, 84–87, 87n3, 89
xxxi, xxxv–xxxviii Recipient, 1, 168
Pictet, Marc-Auguste, xxxiv Red heat, 80, 98, 116, 117, 119–121
Pigment, xxix, xxxi, 22 Requirement for light in photosynthesis, xix
Pine, xxxvi, 19, 38, 92, 97 Respiration, xviii, xix, xxiii, xxiv, 16, 26
Robin, Paul, xxx, xxx(n28), xxxiv–xxxvi,
Plant albumin, 55, 61, 156
xxxviii, xxxix
Plant anatomy, xvi, xxxvi(n34)
Root(s), x, xiv, xv, xx, xxi, xxv, xxxix, 13, 16,
Plant chemistry, xxxv, xxxvi, xxxviii
18, 22, 41–44, 47, 51, 77, 78, 82, 87, 89, 91,
Plant dyes, 60, 119
92, 95, 97, 100–102, 108, 156, 158
Plant macronutrients, xxv, xxv(n18)
Root suction, 94, 97, 98
Plant micronutrients, xxv(n18)
Root support, 93
Plant minerals, xxi, xxviii
Royal Society of London, xxxix
Plant mold, 15, 16, 41, 43, 44, 68, 79n5, 82, 91,
Ruben, Samuel, xxxi, xxxi(n29)
91n2, 92, 101n14, 112
Plant nutrition, xiii–xv, xv(n3), xvi, xxi, xxiv, S
xxx, xxxvii, xxxviii, 20, 50, 93 Saline solution, 1, 94n5, 98n11, 99, 100, 102,
Plant sap, 120 104, 106, 158
Plant science, xiv, xxxviii Saline substance, 67, 100, 104, 108, 109
Plant stress, xxiv Sand, 11, 38, 63, 67, 70, 92, 93, 106, 114n6
Plate of Figures, lix Sap, 58, 84, 108, 156, 168
Plumule, 5, 8 Sapwood, 51, 57, 58, 74, 105, 108, 111–113,
Pneumatic chemistry, xvii 123, 128, 129, 168
Pneumatic trough, xvii, xvii(n7) Sawdust, 57, 58, 70
Political turmoil, xxxiii Schleiden, Matthias Jakob, xiv, xxx
Polstorff, L., xxix Secretion, 26, 59, 96, 97, 102
Portrait of Théodore de Saussure, ii, xxxiii Seed maturation, 110
Potash, xxix, xxxv, 11, 14n4, 15, 22, 31, 34–36, Seeds, 1–8, 11, 20, 59, 66, 71, 82, 92, 109–116,
51, 58, 65, 67, 78, 89, 98, 107, 110–112, 155, 157, 159
115, 117–121, 133, 135, 137, 141, 143, 145, Senebier, Jean, xix–xxi, xxiv, xxxii, xxxiv,
147, 153, 157, 163, 165–168 xxxvi, 8, 12, 16, 22, 48, 79, 80, 83, 171–173,
Potassium, 98, 105 175, 176, 180
Potassium sulfide, ix, xviii(n9), 34, 78, 80 Shade, xxxvii, 9, 11, 13, 14, 26n1, 28, 37, 43,
Precipitate, 53, 58, 65, 67, 100n12, 117, 45, 47–50, 75–77, 80, 81, 87, 155
119–121 Shrubs, 39
Priestley, Joseph, ix, xvii–xix, xxi(n16), xxxv, Silica, 91n1, 105–107, 109, 110, 112–114, 117,
xxxvi, 16, 59n6, 79, 171, 174, 175 123, 159, 168
Proximate constituents of plants, 53, 70 Siliceous soil, 106
Purity of atmosphere, 13, 14, 17–19, 28, 36, 85 Slaked lime, 14, 36, 50
188 Index

Soda, l, 65, 93, 98, 99, 117, 120, 121, 157, 163, Transmutation, xiii, xv, xvii, xix, xxii, xxv,
166–168 xxviii, 91
Sodium, l, 98, 105 Transpiration, 47, 82, 100, 104, 105, 113, 159
Soft wood, 109 Tull, Jethro, xiv, 91, 176
Soil, xiii, xiv, xvi, xxi, xxiii, xxvi, xxx, 16, Turpentine, 59, 156
26n1, 38, 42, 45, 50, 65, 84, 91, 93, 99, 100,
101, 101n14, 105–110, 112, 113, 115, 126, U
158 Unslaked lime, 166, 168
Soil fertility, xv, 92
Soil science, xiii, xv, xxx V
Solar rays, 8, 9, 37, 82 Vacuum, 28, 47, 54, 59n6, 76, 81, 82, 157
Solid substance of plants, xvi, 84, 110 Vacuum pump, 81, 82, see also Air pump
Solution, xx, 53, 54, 91, 94, 96–98, 102, 106, van Helmont, Jan, xv, xxi, xxiv, 91, 101n14,
117–121, 158 176, 178
Species, x, xxii, xxv, lii, 5, 97, 103, 104, 107, van Niel, Cornelius Bernardus, xxxi
115, 155, 156, 158, 159 Vauquelin, Louis-Nicolas, xxxviii, 65, 109,
Spirit of wine, xviii, 48, 169 111, 118, 120, 174, 180, 181
Sprengel, Carl S., xxix Vinegar, xxiv, 1, 55, 56, 61, 117–119, 119n10
Spring water, 16, 22, 42, 63, 65, 92 Viscosity, 98, 108, 118, 118n8, 158
St. Théodule glacial pass, xxxv Vital force, xiii, xiv, xv, xxi, xxv, xxvii, xxx, 91
Stagnant water, 42, 64 Vitiation, xix, 169
Stahl, Georg Ernst, xv, xviii Voigt, Siegmund, xxviii, xxx
Starchy seeds, 59 von Haller, Albrecht, xxxvi(n34)
Stomata, xvi, 2, 23 von Liebig, Justus, xiv, xxix, xxx, xxxii
Succulent plant, 15n4, 38, 41, 83, 88n5 von Mayer, Julius Robert, xxvii
Succulents, xxvi, 22, 23, 30, 35, 36, 38, 51, 52, von Sachs, Julius, xiv, xv(n3), xxviii, xxx(n28)
83, 87, 90, 92 Voyages dans les Alpes, xxxvi, 68, 174
Suction, 47, 94, 96, 98, 104, 113
Sugar, xxiv, 6, 55, 91n1, 93–95, 97, 98, 116,
W
155, 158
Waksman, Selman A., xv, xxviii, xxix,
Sulfide, 37n7, 46, 78
xxx(n28)
Sulfuric acid, xxxv, 2, 65
Water, ix, xiii, xiv, xviii, xx, xxi, xxiii, xxv, 1,
Sun, xxvii, 8, 11, 12n1, 14–16, 20, 21, 23, 26,
3–5, 7–9, 12, 15, 16, 21, 34, 44, 50, 57, 58,
33, 35, 36, 45, 50, 76, 79–82, 87–90, 155,
60, 70, 75, 80, 85–90, 100, 110, 112, 115,
157
116, 119–121, 155, 157, 159, 162
Sun-exposed plant, 21, 23
Water of vegetation, xxvi, 13n3, 21, 26n1, 31,
Sunlight, xiv, xix, xxxvii, 8, 9, 12, 13, 17, 31,
32, 84, 104, 105, 122
36, 43, 45, 48, 77, 78, 82, 84, 89, 90, 114,
Water-soluble organic matter, xv
155, 157
Surface area, 22, 37, 45, 76, 88, 106, 115 Water-soluble salts, 103, 107, 116, 159
Swiss Republic, xlix Water-soluble substances, 92, 101, 102
Well formed wood, 108, 112, 113
T Wet method, 119, 169
Table of carbonizations, 71 Wiegmann, Arend Friedrich August, xxix
Tables of amounts of oxygen consumed by Wieler, Arwed, xxx
different leaves, 39 Wine, xxiv, 55–57, 61, 156
Tables of incinerations and analyses, xxxii, 122 Woodward, John, xvi, 42, 173
Tap-rooted plants, 42 Woody branch, 15, 45, 46, 51, 75
Temperature, 2, 7, 8, 17, 26–28, 55–57, 70, 71, Woody plant, 82, 103, 104, 159
109, 116 Woody stem, 45
Thaer, Albrecht Daniel, xiv, xxviii, xxix
Thick-leaved plant, 81, 89, 156, 158 Y
Thin-leaved plant, xxvi, 22, 35–37, 87, 88, 90 Yellowed leaves, 112
Topsoil, 68 Young plant, 8, 112
Index of Authors de Saussure Cited

Note: Page numbers that are not in brackets I

refer to de Saussure’s text; bracketed numbers Ingen-Housz, Jan, ix, 8, 21, 36n6, 79, 83, [171,
refer to the fuller details for the references, as 172, 174, 176]
traced by the translator in Appendix 3.
K
Karsten, 115n7, [180]
B
Kirwan, Richard, 72, 91n2, 101n14, 103, [174,
Bergman, Torbern Olaf, 91n2, [177] 177–179]
Berthollet, Claude-Louis, ix(n1), 32, 55, 60, Klaproth, Martin Heinrich, 64, 73, 119, [174,
83, 97n7, 120, [170, 172–177, 181] 181]
Bonnet, Charles, 91n1, 91n2, 104, [176, 177,
179]
L
Lampadius, Wilhelm August, 106, 107, [180]
C
Lavoisier, Antoine-Laurent, 4, 7, 54, 55, 69, 86,
Cornette, Claude-Melchior, 99, [178] [170, 173]
Leblanc, Nicolas, 98, [178]
D Lefebure, E. A., 9, [171]
Dalrymple, William, 93, [177]
Dalton, John, 32n4, [172]
M
de Bullion, Marquis, 99, [178] Marggraf, Andreas Sigismund, 111, [180]
de Candolle, Augustin-Pyramus, 88, [176]
de Fourcroy, Antoine-François, 53, 55, 111, P
118, 120, [173, 180, 181] Percival, Thomas, 12, [171]
de Saussure, Horace-Bénédict, 68, [174] Perthuis de Laillevault, Léon de,103, 116, [179,
de Saussure, Théodore, 3, 117, [170, 181] 180]
Duhamel du Monceau, Henri-Louis, 42, 91n2, Pott, Johann Heinrich, 111 [180]
99, [173, 177, 178] Priestley, Joseph, ix, 16, 59n6, 79, [171, 174,
175]
G
Giobert, Jean-Antoine, ix(n1), 92, [170, 177] R
Guyton de Morveau, Louis Bernard, 81n8, Rozier, François (l’abbé), 56, [174]
[175] Rückert, Georg Christian Albrecht, 12n2, 103,
[171, 179]
H
Hales, Stephen, 32, 41, 46, 101, 104, [172, 173, S
179] Senebier, Jean, ix, 8, 12, 16, 22, 48, 79, 80, 83,
Hassenfratz, Jean-Henri, 20, 21, 91n2, 92, 113, [171–173, 175, 176, 180]
[172, 177] Spallanzani, Lazaro, 83, [175, 176]
Haüy, René-Just, 100, [178] Sprengel, Kurt, 2n2, [170]

J. F. Hill, Chemical Research on Plant Growth, 189

DOI 10.1007/978-1-4614-4136-6, © Springer Science+Business Media New York 2013
190 Index of Authors de Saussure Cited

T van Helmont, Jan, 91, 101n14, [176, 178]

Thompson, Benjamin (Comte de Rumford), 8, Vauquelin, Louis-Nicolas, 65, 109, 111, 118,
[171] 120, [174, 180, 181]
Thouvenel, Pierre, 99, [178] von Humboldt, Alexander, 2, [170]
Tillet, Mathieu, 91n2, [177]
Tull, Jethro, 91, [176] W
Westrumb, Johann Friedrich, 115n7, [180]
V Woodhouse, James, 79, [175]
Valmont de Bomare, Jacques-Christophe, 1n1, Woodward, John, 42, [173]
[170]
About the Author and the Translator

Théodore de Saussure (1767–1845) Plant physiologist Théodore de Saussure was

born in Geneva, Switzerland, where he lived all his life. He was the son of the
famous geologist, meteorologist, and mountain explorer Horace-Bénédict de Saus-
sure. Théodore de Saussure completed the fundamental, pioneering research on
photosynthesis and presented the first concise summary of the field in his 1804 book,
Recherches chimiques sur la Végétation. His major contributions to the knowledge
of plant nutrition were his finding that water is incorporated into the dry matter of
plants; that the source of plant carbon is atmospheric carbon dioxide, not dissolved
organic matter in the soil; that minerals are absorbed by the plant from the soil, not
created in the plant by transmutation of other substances; and that minerals are essen-
tial for plant growth. His discovery of the incorporation of water enabled completion
of the basic, overall chemical equation for photosynthesis. Although he continued to
conduct research for 40 years after his book was published, the book represents his
major contribution to science.
Jane F. Hill received an undergraduate degree in English from Carleton College,
Northfield, Minnesota, USA, in 1968, and earned a master’s degree in biology and
a doctorate in botany from The George Washington University, Washington, D.C.,
USA, in 1976 and 1980, respectively. Before entering graduate school, she worked
as a writer and editor for the United States Department of Agriculture (USDA) Soil
Conservation Service, where she developed an interest in plant growth, agronomy,
and soil science. As a graduate student, she worked in dendroclimatology research
at the United States Department of the Interior, United States Geological Survey,
Reston, VA, and later was a volunteer collaborator at plant research laboratories at
the USDA Beltsville Agricultural Research Center, Beltsville, MD, USA. She has
written extensively on a variety of subjects involving many aspects of botany and
biology, including encyclopedia essays on pioneering photosynthesis research and
other 18th and 19th century history-of-science topics. She wrote a chapter on the
early photosynthesis pioneers for Volume 34 in the series Advances in Photosynthesis
and Respiration (J.J. Eaton-Rye, B.C.Tripathy, and T.D. Sharkey, eds.; Govindjee
and Sharkey, series eds., 2012, Springer).

J. F. Hill, Chemical Research on Plant Growth, 191

DOI 10.1007/978-1-4614-4136-6, © Springer Science+Business Media New York 2013
192 About the Author and the Translator

In addition, Dr. Hill has co-authored a 500-page book on structure, function, and
development of woody plants (J. A. Romberger, Z. Hejnowicz, and J. F. Hill 1993,
Springer-Verlag; re-issued 2004, Blackburn Press). Also, she has published original
research papers on dendroclimatology and wood structure in the American Journal
of Botany.