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Insect Hydrocarbons Biology Biochemistry and Chemical
Ecology 1st Edition Blomquist Gary J. Digital Instant
Download
Author(s): Blomquist Gary J., Bagneres Anne-Genevieve
ISBN(s): 9780521898140, 0521898145
Edition: 1
File Details: PDF, 6.23 MB
Year: 2010
Language: english
This page intentionally left blank
 Insect hydrocarbons

A unique and critical analysis of the wealth of research conducted on the biology,
­biochemistry and chemical ecology of the rapidly growing field of insect cuticular hydro­
carbons. Authored by leading experts in their respective fields, the twenty chapters show
the complexity that has been discovered of the nature and role of hydrocarbons in entomo­
logy. Covers, in great depth, aspects of chemistry (structures, qualitative and quantitative
analysis), biochemistry (biosynthesis, molecular biology, genetics, evolution), physiology,
taxonomy, and ecology. Clearly presents to the reader the array of data, ideas, insights and
historical disagreements that have accumulated during the past half century. An emphasis
is placed on the role of insect hydrocarbons in chemical communication is shown, espe­
cially among the social insects. Includes the first review on the chemical synthesis of insect
hydrocarbons. The material presented is a major resource for current researchers and an
unending source of ideas for new researchers.

gary j. blomquist is chair of the Department of Biochemistry and Molecular Biology

at the University of Nevada, Reno. He has published over 200 original research papers,
reviews, chapters and books, including co-editing the books Pheromone Biochemistry and
Molecular Biology (G. J. Blomquist and R. G. Vogt, 2003) and Pheromone Biochemistry
(G. D. Prestwich and G. J. Blomquist, 1987). His work has been cited over 4500 times
(ISI). Much of Blomquist’s research career has involved the study of insect hydrocarbons,
with an emphasis on their biosynthesis, endocrine regulation and chemical analysis. He
published his first paper on insect hydrocarbons 40 years ago (1969) and has remained
active in the field, collaborating with many of the early leaders including Larry Jackson,
Dennis Nelson, Ralph Howard, Coby Schal and Anne-Geneviève Bagnères.

anne-geneviève bagnères is Director of Research at the CNRS and team leader at the
Institut de Recherche sur la Biologie de l’Insecte (IRBI) in Tours, France. She completed
a PhD on the role of cuticular hydrocarbons in social insects at the University of Paris
6 in 1989, and received the Chancellerie of the Universities of Paris Prize for her Ph.D.
work. She spent a year of postdoctoral studies in David Morgan’s laboratory in 1990, and
a sabbatical in 1996–97 in the laboratories of Gary Blomquist and Coby Schal. AGB is
primarily interested in the chemical ecology of social insects, where she continues to be a
leading contributor and proponent of the concept of chemical signature. While her primary
research focuses on termites, she participates in several collaborative studies involving
the chemistry of other insects. She is an active member of the International Society of
Chemical Ecology (ISCE) and the bureau of the French section of the International Union
for the Study of Social Insects. She has published nearly 100 original research papers,
reviews and chapters.
 Inse c t H ydro c a r bo n s
Biolo gy, Bi o c h emis t ry, a n d
C he mi c al E c o l o g y

Gary J. Blom qui s t

University of Nevada

ANNE-GENEVIÈVE BAGNÈRES
Centre National de la Recherche Scientifique,
Université de Tours
CAMBRIDGE UNIVERSITY PRESS
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accurate or appropriate.
 Contents

List of contributorspage vii

Foreword ix
Ralph W. Howard
Acknowledgments xi
Part I Chemistry, Biochemistry, and Physiology
1 Introduction: history and overview of insect hydrocarbons 3
Gary J. Blomquist and Anne-Geneviève Bagnères
2 Structure and analysis of insect hydrocarbons 19
Gary J. Blomquist
3 Biosynthesis of cuticular hydrocarbons 35
Gary J. Blomquist
4 Molecular biology and genetics of hydrocarbon production 53
Claude Wicker-Thomas and Thomas Chertemps
5 Site of synthesis, mechanism of transport and selective
  deposition of hydrocarbons 75
Anne-Geneviève Bagnères and Gary J. Blomquist
6 Cuticular lipids and water balance 100
Allen G. Gibbs and Subhash Rajpurohit
7 Chemical taxonomy with hydrocarbons 121
Anne-Geneviève Bagnères and Claude Wicker-Thomas
8 Chemical synthesis of insect cuticular hydrocarbons 163
Jocelyn G. Millar
9 Oxygenated derivatives of hydrocarbons 187
James S. Buckner
Part II Chemical Communication
10 Perception and olfaction of cuticular compounds 207
Mamiko Ozaki and Ayako Wada-Katsumata
11 Nestmate recognition in social insects and the role of hydrocarbons 222
Jelle S. van Zweden and Patrizia d’Ettorre

v
vi Contents

12 Cuticular hydrocarbon cues in the formation and maintenance

  of insect social groups 244
Michael Greene
13 Hydrocarbon profiles indicate fertility and dominance status
  in ant, bee, and wasp colonies 254
Jürgen Liebig
14 Chemical deception/mimicry using cuticular hydrocarbons 282
Anne-Geneviève Bagnères and M. Cristina Lorenzi
15 Behavioral and evolutionary roles of cuticular hydrocarbons in Diptera 325
Jean-François Ferveur and Matthew Cobb
16 Contact recognition pheromones in spiders and scorpions 344
Marie Trabalon and Anne-Geneviève Bagnères
17 Hydrocarbons as contact pheromones of longhorned beetles (Coleoptera:
  Cerambycidae) 375
Matthew D. Ginzel
18 Polyene hydrocarbons, epoxides, and related compounds as components
  of lepidopteran pheromone blends 390
Jocelyn Millar
19 Volatile hydrocarbon pheromones from beetles 448
Robert J. Bartelt
20 Future directions in hydrocarbon research 477
Abraham Hefetz, Claude Wicker-Thomas and Anne-Geneviève Bagnères
Index 486
 Contributors

Bagnères, Anne-Geneviève, CNRS UMR 6035, I.R.B.I., University of Tours, Parc de

Grandmont, 37200 Tours, France
Bartelt, Robert J., ARS, Crop Bioprotection Research, 1815 N University St Peoria, IL
61604–3902, USA
Blomquist, Gary J., Department of Biochemistry and Molecular Biology, University of
Nevada, Reno, NV 89557–0014, USA
Buckner, James S., Red River Valley Agricultural Research Center, USDA-ARS,
University Station, Fargo, ND 58105, USA
Chertemps, Thomas, University of Paris 6, UMR INRA 1272 Physiologie de l’Insecte
Signalisation et Communication, 75252 Paris Cedex 05, France
Cobb, Matthew, Faculty of Life Sciences, University of Manchester, Oxford Road,
Manchester, M13 9PT, UK
d’Ettorre, Patrizia, Centre for Social Evolution, Department of Biology, University of
Copenhagen, Universitetsparken 15, 2100, Copenhagen, Denmark
Ferveur, Jean-François, UMR CNRS 5548, Développement-Communication Chimique,
Université de Bourgogne, Faculté des Sciences, 6 bd. Gabriel, 21000 Dijon, France
Ginzel, Matthew D., Department of Entomology, Purdue University, 901 West State
Street, West Lafayette, IN 47907–2089, USA
Gibbs, Allen G., Department of Biological Sciences, 4505 Maryland Parkway, University
of Nevada, Las Vegas, Las Vegas, NV 89154, USA
Greene, Michael, Department of Integrative Biology, University of Colorado Denver,
Denver, CO 80217, USA
Hefetz, Abraham, George S. Wise Faculty of Life Sciences, Department of Zoology, Tel
Aviv University, Ramat Aviv 69978, Israel
Howard, Ralph W., 701 Pine Street, Wamego, Kansas 66547, USA
Liebig, Jürgen, School of Life Sciences, Arizona State University, Tempe, AZ 85287–
4501, USA
Lorenzi, M., Cristina. Department of Animal and Human Biology, University of Turin,
10123 Torino, Italy
Millar, Jocelyn G., Department of Entomology, University of California, Riverside, CA
92521, USA

vii
viii List of contributors

Ozaki, Mamiko, Department of Biology, Kobe University, Rokkodai, Nada-ku, Kobe,

657–8501, Japan
Rajpurohit, Subhash, Department of Biological Sciences, 4505 Maryland Parkway,
University of Nevada, Las Vegas, NV 89154, USA
Trabalon, Marie, IPHC-DEPE, Physiologie du Comportement, UMR 7178 CNRS, Faculté
des Sciences et des Techniques, B.P. 70239, 54 506 Vandoeuvre-Les-Nancy, France
van Zweden, Jelle S., Centre for Social Evolution, Department of Biology, University of
Copenhagen, Universitetsparken 15, 2100, Copenhagen, Denmark
Wada-Katsumata, Ayako, Section of Insect Physiology, Kyoto University, Oiwake-cho,
Kitashirakawa, Sakyo-ku, Kyoto 606–8502, Japan
Wicker-Thomas, Claude, CNRS UPR 9034. Laboratoire Evolution, Génomes et
Spéciation, Avenue de la Terrasse, Bâtiment 13, Boite Postale 1, 91198 Gif sur Yvette,
France
 Foreword

As every young chemist once realized or was told, hydrocarbons are simple (even
“boring”) molecules of limited practical importance, and even less scientific interest.
Certainly this was the situation in the early 1960s when the field of chemical ecology was
established. Although it was known that insects and other arthropods seemed to have very
high molecular weight hydrocarbons on their cuticle, nothing else was known of their
chemistry or of their biological importance, other than that they were possibly involved in
water retention. As in many cases in science, progress is sometimes stymied by lack of a
particular tool or technique. The invention of gas chromatographs and their coupling with
mass spectrometers in this same time period formed the critical impetus for the birth of a
major field of science, and the realization that those “simple” hydrocarbon molecules are
the end product of eons of evolution, that they are far from simple, and that they are criti­
cal elements in an amazing variety of biological roles not only in arthropods, but also in
microorganisms, plants, and numerous animal phyla.
By 1980, enough progress had been made in the elucidation of the chemistry and eco­
logical roles of arthropod hydrocarbons to warrant a comprehensive review in the Annual
Review of Entomology (130 papers constituted the entire literature at that time). Progress
continued unabated in the ensuing years, and in 2005 a second limited review of the progress
since 1980 of our knowledge of the chemistry and biological roles of hydrocarbons was
published in the Annual Review of Entomology. During this same interval, several other
specialist reviews were also published. Despite the enormous amount of insect hydrocar­
bon research conducted worldwide during these years (literally thousands of publications
and numerous symposiums and presentations), no one had stopped long enough to write a
monograph on the field. This book, therefore, meets a long-felt need by numerous scientists
to stop and see where the field has been and where it is going.
The first generation of insect hydrocarbon scientists are now all either retired or near
retirement, and subsequent younger generations have taken their place, using new technol­
ogies and building on the solid foundation they inherited to explore ever-expanding facets
of these fascinating molecules. Clearly, this discipline has matured to a point where no sin­
gle scientist is capable of covering all that has been discovered, nor is it possible to cover,
even superficially, the entire field in a few book chapters. The authors of this monograph

ix
x Foreword

are all leading experts in their respective fields, and even a cursory examination of the Table
of Contents will reveal the complexity that has been discovered of the nature and role of
hydrocarbons in the science of life. The twenty chapters cover in great depth aspects of
chemistry (structures, qualitative and quantitative analysis), biochemistry (biosynthesis,
molecular biology, genetics, evolution), physiology, taxonomy, and ecology. They clearly
present to the reader the enormous wealth of data, ideas, insights and historical disagree­
ments that have accumulated during the past half century. The material so presented is a
major resource for current researchers, an unending source of ideas for new researchers,
and serves as an exemplar for surveys of similar rapidly developing areas of research in
other biological disciplines.
I am privileged to have been one of the early researchers in this marvelous field of
scientific endeavor and to have made some small contributions to its development over
a period of roughly thirty years. The comradeship I have experienced on a worldwide
basis with numerous very gifted scientists (including many of the authors in this book) has
brought me many moments of great pleasure and a strong sense of accomplishment that I
might never have known otherwise. The editors of this book, Gary Blomquist and Anne-
Geneviève Bagnères, have traversed this journey with me almost from the first. I thank
them for their many kindnesses over the years, and for bringing this monograph to fruition.
Their enthusiasm and scholarship have never failed to be less than outstanding, nor have
their gifts as friends and colleagues to not only me, but so many others.

Ralph W. Howard, USDA-ARS, Retired

 Acknowledgments

We thank our mentors, collaborators, co-workers, post-docs and students for making
work in this field both fun and rewarding. GJB especially thanks his Ph.D. adviser, Larry
Jackson, for sparking a lifelong interest in insect hydrocarbons, and long-time collabora­
tors Ralph Howard, Dennis Nelson and Coby Schal for making the journey more enjoyable
and productive. AGB thanks Jean-Luc Clément for his special ability to have made science
and working with people so enjoyable during their collaborative years, and for his vision­
ary impact on deciphering the role of cuticular hydrocarbons in chemical communication.
AGB also thanks Ralph Howard, David Morgan, and Coby Schal for their kindness and
serving as catalysts for her work, and her team and Institute (IRBI) for accommodating her
calendar. She thanks her colleagues of the CNRS GDR d’Ecologie Chimique for their sup­
port. We thank the authors of this work for preparing up-to-date and stimulating chapters
in a timely manner. It was a great pleasure to interact and learn from them. We thank Cheri
Blomquist and Andy Corsini for their help in editing a number of chapters. And finally
AGB wishes to thank – and dedicate this book to – her family, Frédéric for his continued
support, and to her children Lucas and Eva.
GJB acknowledges the financial support from the National Science Foundation,
the USDA-NRI and the Nevada Agriculture Experiment Station for the work of his
laboratory.

xi
 Part I
Chemistry, Biochemistry, and Physiology
 1
Introduction: history and overview of
insect hydrocarbons
Gary J. Blomquist and Anne-Geneviève Bagnères

The long-chain hydrocarbons of insects play central roles in the waterproofing of the insect
cuticle and function extensively in chemical communication where relatively non-volatile
chemicals are required. The recognition of the critical roles that hydrocarbons serve as sex
pheromones, kairomones, species and gender recognition cues, nestmate recognition, dom­
inance and fertility cues, chemical mimicry, primer pheromones and task-specific cues has
resulted in an explosion of new information in the past several decades, and, indeed, served
as the impetus for this book.
A number of reviews and chapters on specific topics related to insect hydrocarbons
have been published over the past few decades (Jackson and Blomquist, 1976a; Blomquist
and Jackson, 1979; Howard and Blomquist, 1982, 2005; Blomquist and Dillwith, 1985;
Blomquist et al., 1987; Lockey, 1988, 1991; Howard, 1993; Nelson and Blomquist, 1995;
Gibbs, 2002), and in this book we attempt to bring this information up-to-date and in one
place. At the time the first insect hydrocarbons were chemically identified in the 1960s and
early 1970s, no one could have predicted the amount of interest that they would generate.
Indeed, a pioneer in this field advised one of the authors (GJB) to go into a field other than
insect hydrocarbons as he began his independent research career in the early 1970s, as he
saw no future in this area. This prophecy proved very wrong and illustrates how difficult it
is to predict the future of any scientific field.
The ability of insects to withstand desiccation was recognized in the 1930s to be due
to the epicuticular layer of the cuticle. Wigglesworth (1933) described a complex fatty or
waxy substance in the upper layers of the cuticle which he called “cuticulin”. The ­presence
of hydrocarbons in this wax of insects was suggested by Chibnall et al. (1934) and Blount
et al. (1937), and over the next few decades the importance of hydrocarbons in the ­cuticular
wax of insects was established (Baker et al., 1963 and references therein). The first relatively
complete chemical analyses of the hydrocarbons from any insect, the American cockroach,
Periplaneta americana (Baker et al., 1963), occurred after the ­development of gas-liquid
chromatography (GLC). The three major components of the hydrocarbons of this insect,
n-pentacosane, 3-methylpentacosane and (Z,Z)-6,9-heptacosadiene, represent the three
major classes of hydrocarbons on insects, n-alkanes, methyl-branched alkanes and alkenes.
Baker and co-workers (1963) were able to identify n-pentacosane by its elution time on
GLC to a standard and its inclusion in a 5-angstrom molecular sieve. 3-Methylpentacosane

3
4 Chemistry, Biochemistry, and Physiology

was identified by its failure to be absorbed in a 5-angstrom ­molecular sieve and mass
­spectral data. Silver nitrate column chromatography separated the C27 diene from other
­hydrocarbon components, and the double-bond positions and isomeric composition were
determined by a variety of techniques including infrared analysis and oxidative cleavage
of the double bonds. The composition of the hydrocarbons of P. ­americana is unique in
their simplicity, with over 90% of the hydrocarbons comprised of only three components
(Jackson, 1972), and it was a fortunate choice of insect species for Baker et al. (1963). In
general, the hydrocarbon composition of insects is much more complex, and sometimes
consists of well over a hundred components. In an earlier analysis of the cuticular wax of
the Mormon cricket, Anabrus simplex (Baker et al., 1960), the complexity of the hydrocar­
bon mixture (Jackson and Blomquist, 1976b) made it impossible for individual components
to be characterized in the early 1960s (Baker et al., 1960). The development and applica­
tion of combined gas–liquid chromatography and mass spectrometry was key to the rapid
and efficient analysis of insect hydrocarbons. In the late 1960s and during the next few
decades, GC–MS analysis of insect hydrocarbons was established (Nelson and Sukkestad,
1970; Martin and MaConnell, 1970), and over the next several decades the hydrocarbons
of hundreds of insect species were analyzed, first on packed columns and then much more
efficiently on capillary columns. It was recognized that for many insect species, very com­
plex mixtures of normal (straight-chain), methyl-branched and unsaturated components
exist, with chain lengths ranging from 21 to 50 or so carbons. In ­retrospect, the occurrence
of extremely complex mixtures of components might have suggested that hydrocarbons
could play important roles in chemical communication, but only after the recognition of the
number and variety of roles they do play have we come to more fully appreciate the import­
ance of insect hydrocarbons in chemical communication. The variety of chain lengths and
the number and positions of the methyl branches and double bonds provide the insect with
the chemical equivalent of the visually variable colored plumage of birds. Cvačka et al.
(2006) reported hydrocarbons of up to 70 carbons in chain length using MALDI–TOF
mass spectrometry, indicating that the earlier chain-length limits of 50 or so carbons might
be a limitation of GC–MS techniques and not the ability of the insect to make longer chain
components. The structure and chemical analysis of insect hydrocarbons are covered in
Chapter 2 (Blomquist, this book).
A series of studies in the 1960s demonstrated that labeled acetate was incorporated into
insect cuticular hydrocarbons (Vroman et al., 1965; Lamb and Monroe, 1968; Nelson,
1969) establishing the de novo biosynthesis of the majority of components. Because of the
simple hydrocarbon composition and the ease of isolating 3-methylpentacosane and (Z,Z)-
6,9-heptacosadiene from the P. americana, early in vivo biosynthetic studies concentrated
on this insect (Conrad and Jackson, 1971; Blomquist et al., 1975; Major and Blomquist,
1978; Dwyer et al., 1981). Work with radiolabeled precursors established the elongation–
decarboxylation pathway for hydrocarbon biosynthesis (Major and Blomquist, 1978) and
the incorporation of propionate, as a methylmalonate unit, to form the ­methyl-branching
unit (Blomquist et al., 1975). Over the next several decades, studies evolved from in vivo
studies to the use of microsomal preparations to gain an understanding of how the major
 Introduction 5

components were made and regulated. In the final step of hydrocarbon formation, the elon­
gated acyl-CoA is reduced to aldehyde and oxidatively decarbonylated to hydrocarbon
(Reed et al., 1994, 1995). The biosynthesis of insect hydrocarbons is presented in Chapter
3 (Blomquist, this book). Only recently have the powerful tools of molecular biology been
applied to studies of insect hydrocarbon biosynthesis, and this is covered in Chapter 4
(Wicker-Thomas and Chertemps, this book).
In the mid-1960s, Locke (1965) presented an illustration of insect cuticular lipids in
which the newly synthesized hydrocarbons exited the epidermal oenocyte cells where they
were synthesized, were transported through pore canals and then formed an outer layer on
the cuticle. He pictured the polar head groups of fatty acids interacting with the cuticle,
and then the hydrocarbons layered on top of the acyl chains of fatty acids. This served
as an excellent model in which to test a number of hypotheses, many of which are still
unanswered. It is now clear that newly synthesized hydrocarbons are taken up first by
lipophorin and transported via the hemolymph (Bagnères and Blomquist, Chapter 5, this
book). How they get transferred to the surface of the insect is unknown, although a number
of species are able to selectively transport shorter chain hydrocarbon pheromones and phe­
romone precursors to the pheromone gland on the abdomen, whereas longer chain cuticular
hydrocarbons are transported to cover the entire cuticle (Schal et al., 1998; Jurenka et al.,
2003). In many cases, hydrocarbons comprise the majority of the cuticular lipids, making
the proposed role of fatty acids of lesser importance, and the arrangement of hydrocarbons
on the surface of the insect is unknown, although suggestions have been made that the
components of most importance in chemical communication may be on the outer surface
(Ginzel et al., 2003; Ginzel, Chapter 17, this book). A clear understanding of hydrocarbon
transport to the cuticle and arrangement of hydrocarbons on the surface of insects is, at this
time, unavailable.
The large surface-to-volume ratio of insects makes it important that excessive ­evaporation
be prevented. The first recognized and perhaps still primary function of insect cuticular
lipids, especially hydrocarbons, is to restrict water loss, a fact first recognized by Ramsay
(1935). Early investigations centered on the measurement of transpiration of water and the
role of cuticular lipids in preventing water loss, and these early studies were reviewed in
Barton-Brown (1964), Beament (1961, 1964), Edney (1957), Richards (1951), and Locke
(1965). It is now clear that the cuticle is permeable to water vapor, and that the cuticular
lipid layer plays a major role in reducing transpiration. Cuticles from which the surface
lipids are removed with organic solvents are relatively permeable to water. Vegetable oils,
lecithin and a series of wetting agents and detergents show widely different effects on
­permeability of the cuticle to water (Beament, 1945). The waterproofing observed with
intact insects is closely duplicated when extracted cuticular lipids are deposited on collo­
dion membranes or intact wing membranes (Beament, 1945). The transpiration rate from
an insect is found to increase rather abruptly at a temperature that closely corresponds to
the transition point or change of phase point of the lipids on the cuticle of a particular spe­
cies. This transition point is near the melting range of the ­extractable lipid (Wigglesworth,
1945; Beament, 1945). Rapid desiccation occurs as a result of scratching the outer surface
6 Chemistry, Biochemistry, and Physiology

of the epicuticle with abrasive dusts. The abrasion needs to be only deep enough to pen­
etrate the epicuticle. Adsorption of the lipids onto the dust may also play a role to a certain
extent, especially on insects where the lipids are soft (Wigglesworth, 1945). If the abraded
insect is kept in a moist atmosphere to prevent ­desiccation, the lipid layer is apparently
restored, along with the ability to resist ­desiccation. The lipid melting model achieved text-
book status (Randall et al., 1997; Chapman, 1998), although a number of researchers have
pointed out its limitation. In the last several decades, Gibbs and Rajpurohit (Chapter 6, this
book) have re-examined this phenomenon, and presents our current understanding of the
role of cuticular hydrocarbons and other lipids in restricting water loss from insects.
The cuticular hydrocarbons of many insect species are extremely complex and involve
mixtures of normal, mono-, di- and tri-unsaturated and mono-, di-, tri-, tetra- and
­pentamethyl-branched components of chain lengths between 21 and 50 carbons. The
number of known components is very large and the number of possible isomers much
­larger. Insects biosynthesize the vast majority of their hydrocarbon components, and thus
­hydrocarbon composition may be considered a part of an insect’s genotype and ­therefore
available for taxonomic use. The early studies by Jackson (1970), Jackson and Baker
(1970), Jackson and Blomquist (1976a), Lockey (1980) and Carlson and collaborators
(Carlson and Service, 1979, 1980; Carlson and Brenner, 1988; Carlson, 1988) recognized
the special role of cuticular hydrocarbons in chemical taxonomy. The importance of using
hydrocarbons in chemical taxonomy has continued to grow, with more recent efforts cen­
tered on Drosophilidae and termites. This work is covered in Chapter 7 (Bagnères and
Wicker-Thomas, this book).
In order to more fully understand and interpret mass spectra and retention ­indices of
methyl-branched and unsaturated hydrocarbons, standard known hydrocarbon ­compounds
were synthesized and analyzed (Pomonis et al., 1978, 1980). The availability of standards
with one or more methyl branches and specific double bonds allowed for ­better ­understanding
of the roles individual components and hydrocarbon classes played in determining ­critical
transition temperatures and waterproofing characteristics of ­hydrocarbons (Gibbs and
Pomonis, 1995). With the increasing recognition that many hydrocarbon ­components play
important roles in chemical communication, the importance of ­synthesizing ­hydrocarbon
standards to determine the exact role of individual ­compounds became more important. The
first review of the chemical synthesis of long-chain ­hydrocarbons is presented in Chapter 8
(Millar, this book).
While the hydrocarbon fraction of insect cuticular lipids is certainly the most studied and
has been shown to play a key role in a wide range of chemical communication, other ­lipids
are often present on the surface of insects. The most common cuticular lipids in addition to
hydrocarbons include a variety of types of esters, free fatty acids, primary and secondary alco­
hols, ketones and sterols. Triacylglycerols and the more polar phospholipids are not com­
mon components of insect cuticular lipids. In some cases, hydrocarbons are hydroxylated and
metabolized to oxygenated components, and these products include some of the short range
and contact pheromones of the housefly (Blomquist, 2003) and the German cockroach (Schal
et al., 2003). The oxygenated cuticular lipids are discussed in Chapter 9 (Buckner, this book).
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narrow band, but Ewald states correctly that this is the pancreas,
which in many of these cases can be distinctly palpated. According
to Glenard, the kink in the colon causes the constipation, while the
depression of the stomach and intestines leads to vascular
disturbance and impairment of the motor and secretory functions. In
floating kidney there are attacks (simulating gastralgia or renal colic)
characterized by severe abdominal pain, chills, nausea, vomiting,
fever, and collapse. Scarcely any mention is made of such
symptoms, which were first described by Dietl in 18G4, and a more
wide-spread knowledge of their occurrence in connection with this
condition is desirable. My attention was called to them in 1880 by
Palmer Howard in the case of a stout lady, who suffered repeatedly
with the most severe attacks of abdominal pain and vomiting, which
constantly required morphia. A
 720 DISEASES OF THE KIDNEYS. tumor was discovered a
little to the right of the navel, and the diagnosis of probable
neoplasm was concurred in by Flint (Sr.) and Gaillard Thomas. The
patient lost weight rapidly, became emaciated, and in the spring of
1881 again went to New York, where she saw Van Buren, who
diagnosed a floating kidney and said that these paroxysms were
associated with it in a gouty person. He cut off all stimulants,
reassured the lady that she had no cancer, and from that time she
rapidly recovered, and the attacks have been few and far between.
In this patient any overindulgence in eating or in drinking is still
liable to be followed by a very severe attack. These attacks may also
be mistaken for renal colic, and the operation of nephrotomy has
been performed. In other instances the attacks of pain may be
thought to be due to intestinal disease or to recurring appendicitis.
The cause of these paroxysmal attacks is not quite clear. Dietl
thought they were due to strangulation of the kidney or to twists or
kinks in the renal vessels due to the extreme mobility. During the
attacks the urine is sometimes high-colored and contains an excess
of uric acid or of the oxalates. It is stated, too, that blood or pus
may be present. The kidney may be tender, swollen, and less freely
movable. Intermittent hydronephrosis has sometimes been
associated with movable kidney. The diagnosis is rarely doubtful, as
the shape of the organ is usually distinctive and the mobility marked.
Tumors of the gall-bladder, ovarian growths, and tumors of the
bowels may in rare instances be confounded with it. Treatment. —
The kidney has been extirpated in many instances, but the operation
is not witliout risk, and there have been several fatal cases. Stitching
of the kidney — nephrorrhaphy — as recommended by Hahn, is the
most suitable procedure, and statistics recently published by Keen
show that relief is afforded in many cas3s by the procedure. It does
not, however, always succeed. The treatment by trusses and
bandages is not satisfactory, though great relief is sometimes
obtained. As a rule, bandages, with pads pressing to the right of the
navel, are not well borne, as the kidney is often sensitive. In some
instances, however, the greatest relief is experienced by this
procedure. An air-pad beneath the bandage, as recommended by
Newman, is probably the best. In other cases a broad bandage well
padded in the lower abdominal zone pushes up the intestines and
makes them act as a support. In the attacks of severe colic morphia
is required. When dependent, as seems sometimes the case, upon
an excess of uric acid or the oxalates, the diet must be carefully
regulated.
 CIRCULATORY DISTURBANCES. 721 II. CIRCULATORY
DISTURBANCES. Normally the secretion of urine is accomplished by
the maintenance of a certain blood-pressure within the glomeruli
and by the activity of the renal epithelium. Bowman's views on this
question have been generally accepted, and the watery elements are
held to be filtered from the glomeruli ; the amount depending on the
rapidity and the pressure of the blood current ; the quality, whether
normal or abnormal, depending upon the integrity of the capillary
and glomerular epithelium ; while the greater portion of the solid
ingredients are excreted by the epithelium of the convoluted tubules.
The integrity of the epithelium covering the capillary tufts within
Bowman's capsule is essential to the production of a normal urine. If
under any circumstances their nutrition fails, as when, for example,
the rapidity of the blood-current is lowered, so that they are
deprived of the necessary amount of ox3'gen, the material which
filters through is no longer normal (i. e., water), but contains serum
albumen. Cohnheim has shown that the renal epithelium is
extremely sensitive to circulatory changes, and that compression of
the renal artery for only a few minutes causes serious disturbance.
The circulation of the kidney is remarkably influenced by reflex
stimuli coming from the skin. Exposure to cold causes heightened
blood-pressure within the kidneys and increased secretion of urine.
So also in the chills of malaria, after which a large amount of pale
urine may be passed. Congestion of the Kidneys. — (1) Active
Congestion; Hypermmia. — Acute congestion of the kidney is met
with in the early stage of nephritis, whether due to cold or to the
action of poisons and severe irritants. Turpentine, cubebs,
cantharides, and copaiba are all stated to cause extreme hyperaemia
of the organ. The most typical congestion of the kidney which we
see post mortem is that in the early stage of acute Bright's disease,
when the organ may be large, soft, of a dark color, and on section
blood drips from it freely. It has been held that in all the acute fevers
the kidneys are congested, and that this explained the scanty, high-
colored, and often albuminous urine. On the other hand, by Koy's
oncometer, Walter Mendelson has shown that the kidney in acute
fever is in a state of extreme anaemia, small, pale, and bloodless;
and that this anaemia, increasing with the pyrexia and interfering
with the nutrition of the glomerular epithelium, accounts for the
scanty, dark-colored urine of fever and for the presence of albumen.
In the prolonged fevers, however, it is probable that relaxation of the
arteries again takes place. Certainly it is rare to find post mortem
such a condition of the kidney as is described by Mendelson. On the
contrary, the kidney of fever is commonly swollen, the blood-vessels
are congested, and the cortex frequently shows traces of cloudy
swelling. However, the circulatory disturbances in acute fevers are
probably less im 
 722 DISEASES OF THE KIDNEYS. portant than the irritative
effects of either the specific agents of the disease or the products
produced in their growth, or in the altered metabolism of the tissues.
The urine is diminished in amount, and may contain albumen and
tube-casts. (2) Passive Congestion ; Meclia^iical Hypermmia. — This
is found in cases of chronic disease of the heart or lung, with
impeded circulation, and as a result of pressure upon the renal veins
by tumors, the pregnant uterus, or ascitic fluid In the cardiac kidney,
as it is called, the cyanotic induration associated with chronic heart-
disease, the organs are enlarged and firm, the capsule strips off, as
a rule, readily, the cortex is of a deep red color, and the pyramids of
a purple red. The section is coarse-looking, the substance is very
firm, and resists cutting and tearing. The interstitial tissue is
increased, and there is a small celled infiltration between the
tubules. Here and there the Malpighian tufts have become sclerosed.
The blood-vessels are usually thickened, and there may be more or
less granular, fatty, or hyaline changes in the epithelium of the
tubules. The condition is indeed a diffuse nephritis. The urine is
usually reduced, is of high specific gravity, and contains more or less
albumen. Hyaline tube-casts and blood-corpuscles are not
uncommon. In uncomplicated cases of the cyanotic induration
uraemia is rare. On the other hand, in the cardiac cases with
extensive arterio-sclerosis, the kidneys are more involved and the
renal function is likely to be disturbed. III. ANOMALIES OF THE
URINARY SECRETION. 1. HiEMATURIA. The following division may
be made of the causes of haematuria : (1) General Diseases. — The
malignant forms of the acute specific fevers, such as small-pox,
malaria, yellow fever, etc. ; scurvy, purpura, and haemophilia.
Occasionally in leukaemia haematuria occurs. (2) Renal Causes. —
Acute congestion and inflammation, as in Bright's disease, or the
effect of toxic agents, such as turpentine, carbolic acid, and
cantharides. When the carbolic spray was in use many surgeons
suffered from haematuria in consequence of this poison. Renal
infarction, as in ulcerative endocarditis. New growths, in which tlie
bleeding is usually profuse. Tubercle rarely causes haematuria,
though at the onset, when the papillae are involved, there may be
bleeding. Stone in the kidney is a frequent cause. Parasites: T\\q
filar ia sanguinis liominis and i\\Q Bilharzia cause a form of
haematuria met with in the tropics. The echinococcus is rarely
associated with haemorrhage. (3) Affections of the Urinary Passages.
— Stone in the ureter, malignant disease or ulceration of the bladder,
the presence of a calculus, parasites, and, very rarely, ruptured veins
in the bladder. Bleeding from the
 ANOMALIES OF THE URINARY SECRETION. 723 urethra
occasionally occurs in gonorrhoea and as a result of the lodgment of
a calculus. (4) Traumatism. — Injuries may produce bleeding from
any part of the urinary passages. By a fall or blow on the back the
kidney may be ruptured, and this may be followed by very free
bleeding ; less commonly the blood comes from injury of the bladder
or of the prostate. Blood from the urethra is frequently due to injury
by the passage of a catheter, or sometimes to falls or blows. And,
lastly, there are cases in which haematuria occurs for a long time
without discoverable cause, particularly in young persons. The health
may not be seriously impaired. Gull has characterized, in a happy
way, a case of this kind as one of renal epistaxis. Of special interest
is the malarial haematuria which prevails in certain districts and has
already been considered in the section on paludism. The diagnosis of
haematuria is usually easy. The color of the urine varies from a light
smoky to a bright red, or it may have a dark porter color. Examined
with the microscope, the blood-corjDuscles are readily recognized,
either plainly visible and retaining their color, in which case they are
usually crenated, or simply as shadows. In ammoniacal urine or
urines of low specific gravity the haemoglobin is rapidly dissolved
from the corpuscles, but in normal urine they remain for many hours
unchanged. Other tests are rarely necessary. The guaiacum test
consists of the addition to the urine, in a test-tube, of a drop or two
of the tincture of guaiacum and two minims of ozonic ether. A blue
color forms at the line of contact of the two fluids and diffuses itself
through the ether. The spectroscopical examination of the urine may
show either the single band of reduced haemoglobin or the double
band of oxyhaemoglobin between the lines D and E. It is important
to distinguish between blood coming from the bladder and from the
kidneys, though this is not always easy. From the bladder the blood
may be found only with the last portions of urine, or only at the
termination of micturition. In haemorrhage from the kidneys, the
blood and urine are intimately mixed. Clots are more commonly
found in the blood from the kidneys, and may form moulds of the
pelvis or of the ureter. Wlien the seat of the bleeding is in the
bladder, on washing out this organ, the water is more or less blood-
tinged ; but if the source of the bleeding is higher, the water comes
away clear. In many instances it is difficult to settle the question by
the examination of the urine alone, and the symptoms and the
physical signs must also be taken into account. 2.
HEMOGLOBINURIA. This condition is characterized ])y the presence
of blood-pigment in the urine. The blood-cells are either absent or in
insignificant numbers. The coloring matter is not haematin, as
indicated by the old name, hwma46
 724: DISEASES OF THE KIDNEYS. tinuria^ nor in reality
always haemoglobin, but it is most frequently methaemoglobin. The
urine has a red or brownish-red, sometimes quite black color, and
usually deposits a very heavy brownish sediment. When the
haemoglobin occurs only in small quantities, it may give a lake or
smoky color to the urine. Microscopical examination shows the
presence of granular pigment, sometimes fragments of blood-disks,
epithelium, and very often darkly pigmented urates. The urine is also
albuminous. The number of red blood-corpuscles bears no
proportion whatever to the intensity of the color of the urine.
Examined spectroscopically, there are either the two absorption
bands of oxyhaemoglobin, which is rare, or, more commonly, there
are the three absorption bands of methaemoglobin, of which the one
in the red near C is characteristic. Two clinical groups may be
distinguished. (1) Toxic HsBinoglobinuria. — This is caused by
poisons which produce rapid dissolution of the blood-corpuscles,
such as chlorate of potash in large doses, pyrogallic acid, carbolic
acid, arseniuretted hydrogen, carbon dioxide, naphthol, and
muscarine ; also the poisons of scarlet fever, yellow fever, typhoid
fever, malaria, and syphilis. It has also followed severe burns.
Exposure to excessive cold and violent muscular exertion are stated
to produce haemoglobinuria. A most remarkable toxic form occurs in
horses, coming on with great suddenness and associated with
paresis of the hind legs. Death may occur in a few hours or a few
days. Horses are attacked only after being stalled for some days and
then taken out and driven, particularly in cold weather. The affection
is common in horses in this country. The form of haemoglobinuria
from cold and exertion is extremely rare No instance of it, even in
association with frost-bites, came under my observation in Canada.
Blood transfused from one mammal into another causes dissolution
of the corpuscles with the production of haemoglobinuria ; and,
lastly, there is the epidemic hcamoglohinuria of the new-born,
associated with jaundice, cyanosis, and nervous symptoms. (2)
Paroxysmal Hsemoglobinuria. — This rare disease is characterized by
the occasional passage of bloody urine, in which the coloring matter
only is present. It is more frequent in males than in females, and
occurs chiefly in adults. It seems specially associated with cold and
exertion, and has often been brought on, in a susceptible person, by
the use of a cold foot-bath. Paroxysmal haemoglobinuria has been
found, too, in persons subject to the various forms of Raynaud's
disease. Many regard the relation between these two affections as
extremely close ; some hold that they are manifestations of one and
the same disorder. Druitt, the author of the well-known Surgical
Vade-mecum, has given a graphic description of his sufferings, which
lasted for many years, and were accompanied with local asphyxia
and local syncope. The connection, however, is not very common. In
only one of the cases of Raynaud's disease which I have seen was
paroxysmal haemoglobinuria present, and in it epileptic attacks
occurred at the same time. The relation of the disease to
 ANOMALIES OF THE URINARY SECRETION. 725 malaria is
not so close as has been thought by many writers. No doubt it has
been frequently confounded with a malarial haimaturia. The attacks
may come on suddenly after exposure to cold or as a result of
mental or bodily exhaustion. They may be preceded by chills and
pyrexia. In other instances the temperature is subnoi*mal. There
may be vomiting and diarrhea. Pain in the lumbar region is not
uncommon. The haemoglobinuria rarely persists for more than a day
or two — sometimes, indeed, not for a day. There are instances in
which, even in the course of a single day, there have been two or
three paroxysms, and in the intervals clear urine has been passed.
Jaundice has been present in a number of cases. According to Ealfe,
paroxysmal hsemoglobinuria may alternate with general symptoms
of the same character, but associated only with the passage of
albumen and an increased quantity of urea in the urine. In such
cases he supposes that the toxic agent, whatever its nature, has
destroyed only a limited number of the corpuscles, the coloring
matter of which is readily dealt with by the spleen and liver, while
the globulin is excreted in the urine. The cases are rarely if ever
fatal. The essential pathology of the disease is unknown, and it is
difficult to form a theory which will meet all the facts — particularly
the relation with Eaynaud's disease, which is rightly regarded as a
vaso-motor disorder. Increased haemolysis and dissolution of the
haemoglobin in the blood-serum (haemoglobinaemia) precedes, in
each instance, the appearance of the coloring matter in the urine ;
but, as Ponfick has shown, the amount of free haemoglobin must
reach a certain grade before it is excreted. Treatment. — In all forms
of haematuria rest is essential. In that produced by renal calculi the
recumbent posture may suffice to check the bleeding. Full doses of
acetate of lead and opium should be tried, then ergot, gallic and
tannic acid, and the dilute sulphuric acid. The oil of turpentine,
which is sometimes recommended, is a risky remedy in haematuria.
Extr. hamamelis virgin, and extr. hydrastis canad. are also
recommended. Cold may be applied to the loins or dry cups in the
lumbar region. The treatment of haemoglobinuria is unsatisfactory.
Nothing seems to check the occurrence of the attacks. During the
paroxysm the patient should be kept warm and given hot drinks.
Quinine is recommended in large doses, on the supposition — as yet
unwarranted — that the disease is specially connected with malaria.
If there is a syphilitic history iodide of potassium, in full doses, may
be tried. In a warm climate the attacks are much less frequent. III.
Albumtkuria. The presence of albumen in the urine, formerly
regarded as indicative of Bright's disease, is now recognized as
occurring under many circumstances without the existence of
serious organic change in the kidney.
 726 DISEASES OF THE KIDNEYS. Two groups of cases may
be recognized — those in which the kidneys show no coarse lesions,
and those in which there are evident anatomical changes.
Albuminuria without Coarse Renal Lesions.— («) Functional, Socalled
Physiological, Albuminuria. — In a normal condition of the kidney
only the water and the salts are allowed to pass from the blood.
When albuminous substances transude there is probably disturbance
in the nutrition of the epithelium of the capillaries of the tuft, or of
the cells surrounding the glomerulus. This statement is still,
however, in dispute, and Senator, Grainger Stewart, and others hold
that there is a physiological albuminuria which may follow muscular
work, the ingestion of food rich in albumen, violent emotions, cold
bathing, and dyspepsia. The differences of opinion on this point are
striking, and observers of equal thoroughness and reliability have
arrived at directly opposite conclusions. The presence of albumen in
the urine, in any form and under any circumstance, may be regarded
as indicative of change in the renal or glomerular epithelium, a
change, however, which may be transient, slight, and unimportant,
depending upon variations in the circulation or upon the irritating
effects of substances taken with the food or temporarily present, as
in febrile states. Much attention has been given of late years to the
albuminuria of adolescence, or cyclic albuminuria, which is also
believed to be a functional disorder. A majority of the cases occur in
young persons — boys more commonly than girls — and the
condition is often discovered accidentally. The urine, as a rule,
contains only a very small quantity of albumen, but in some
instances large quantities are present. The most striking feature is
the variability. It may be absent in the morning and only present
after exertion, or it may be greatly increased after taking food,
particularly proteids. The quantity of urine may be but little if at all
increased, the specific gravity is usually normal, and the color may
be high. Occasionally, hyaline casts may be found, and in some
instances there has been transient glycosuria. As a rule, the pulse is
not of high tension and the second aortic sound is not accentuated.
Various forms of this affection have been recognized by writers, such
as neurotic, dietetic, cyclic, intermittent, and paroxysmal — names
which indicate the characters of the different varieties. A large
proportion of the cases get well after the condition has persisted for
a variable period. This in itself is an evidence that the changes,
whatever their nature, were transient and slight. In these instances
tlie albumen exists in small quantity, tube-casts are not present, and
the arterial tension is not increased. In a second group the albumen
is more persistent, the amount is larger, though it may vary from day
to day, and the pulse tension is increased. In such instances the
persistent albuminuria probably indicates actual organic change in
the kidney. {b) Febrile Albuminuria. — Pyrexia, by whatever cause
produced,
 ANOMALIES OF THE URINARY SECRETION. 727 may cause
slight albuminuria. The presence of the albumen is due to slight
changes in the glomeruli induced by the fever, such as cloudy
swelling, which cannot be regarded as an organic lesion. It is
extremely common, occurring in pneumonia, diphtheria, typhoid
fever, and even in the fever of acute tonsillitis. The amount of
albumen is slight, .and it usually disappears from the urine with the
cessation of the fever. (c) Hcemic Changes. — Purpura, scurvy,
chronic poisoning by lead or mercury, syphilis, leukaemia, and
profound anaemia may be associated with slight albuminuria.
Abnormal ingredients in the blood, such as bilepigment and sugar,
may cause the passage of small amounts of albumen. The transient
albuminuria of pregnancy may belong to this haemic group, although
in a majority of such cases there are changes in the renal tissue.
Albumen may be found sometimes after the inhalation of ether or
chloroform. {d) Albuminuria occurs in certain affections of the
nervous system. This so-called neurotic albuminuria is seen after an
epileptic seizure and in apoplexy, tetanus, exophthalmic goitre, and
injuries of the head. Albuminuria with Definite Lesions of the Urinary
Organs. — {a) Congestion of the kidney, either active, such as
follows exposure to cold and is associated with the early stages of
nephritis, or passive, due to obstructed outflow in disease of the
heart or lungs, or to pressure on the renal veins by the pregnant
uterus or tumors. {b) Organic disease of the kidneys — acute and
chronic Bright's disease, amyloid and fatty degeneration, suppurative
nephritis, and tumors. (c) Affections of the pelvis, ureters, and
bladder, when associated with the formation of pus. Tests for
Albumen. — Both morning and evening urine should be examined,
and in doubtful cases at least three specimens. If turbid, the urine
should be filtered, though turbidity from the urates is of no moment,
since it disappears at once on the application of heat. Heat and
Nitric-acid Test. — The urine is boiled in a test-tube over a spirit-
lamp, and a drop of nitric acid is then added. If a cloudiness occurs
on boiling, it maybe due to phosphates, which are dissolved on the
addition of an acid. Persistence of the cloudiness indicates albumen.
Heller'' s Test. — A small quantity of faming nitric acid is poured into
the test-tube, and with a pipette the urine is allowed to flow gently
down the side upon the acid. At the line of junction of the two fluids,
if albumen is present, a white ring is formed. This contact method is
trustworthy, and, for the routine clinical work, is probably the most
satisfactory. A diffused haze, due to mucin, is sometimes seen just
above the white ring of albumen. A colored ring at the junction of
the acid and the urine is due to the oxidation of the coloring matters
in the urine. Sir William Roberts strongly recommends the
magnesium-nitric test. One volume of strong nitric acid is mixed with
five volumes of the saturated
 728 DISEASES OF THE KIDNEYS. solution of sulphate of
magnesium. This is used in the same way as the nitric acid in
Heller's test. Picric acid, introduced by George Johnson, is a delicate
and useful test for albumen. A saturated solution is used and
employed as in the contact method. It has been urged against this
test that it throws down the mucin, peptones, and certain vegetable
alkaloids, but these are dissolved by heat. For minute traces of
albumen the trichloracetic acid may be used, or Millard's fluid, which
is extremely delicate and consists of glacial carbolic acid (ninety-five
per cent), 2 drachms ; pure acetic acid, 7 drachms ; liquor potassae,
2 ounces 6 drachms. A quantitative estimate of the albumen can be
made by means of Esbach's tube, but the rough method of heating
and boiling a certain quantity of acidulated urine in a test-tube and
allowing it to stand, is often employed. The depth of deposit can
then be compared with the whole amount of urine, and the
proportion is expressed as a mere trace, almost solid, one fourth,
one half, and so on. This, of course, does not give an accurate
indication of the proportion of albumen in the total quantity of urine.
For the more elaborate methods the reader is referred to the works
on urinalysis. The above tests refer entirely to serum albumen. Other
albuminous substances occur, such as serum globulin, peptones, and
hemialbumose. By saturating the urine with magnesium sulphate,
the globulin is precipitated, coagulated, and then readily separated
from the serum albumen. Traces of peptones are found in the urine
in many acute diseases and in chronic suppuration. They are not
precipitated by heat or nitric acid, but are thrown down by picric acid
and dissolved by heat. If the urine contains peptones, a rose or
pinkish tint is formed at the junction of the two fluids when urine is
allowed to flow gently into a test-tube containing Fehling's solution.
Peptonuria has no clinical significance. Propepton, or hemialbumose,
is not of any practical importance. It was found by Bence- Jones in
the urine in osteomalacia, and occurs occasionally in other
affections. Prognosis. — This depends, of course, entirely upon the
cause. Febrile albuminuria is transient, and in a majority of the cases
depending upon ha3mic causes the condition disappears and leaves
the kidneys intact. An occasional trace of albumen in a man over
forty, with or without a few hyaline casts, and with increased tension
and thick vessel walls, usually indicates changes in the kidneys. The
persistence of a slight amount of albumen in young men without
increased arterial tension is less serious as even after continuing for
years it may disappear. I have already spoken of the outlook in the
so-called cyclic albuminuria. Practically in all cases the presence of
albumen indicates a change of some sort in the glomeruli, the
nature, extent, and gravity of Avhich it is difficult to estimate, so that
other considerations, such as the presence of
 ANOMALIES OF TFIE URINARY SECRETION. 729 tube-
casts, the existence of increased tension, the general condition of
the patient, and the influence of digestion upon the albumen, must
be carefully considered. The physician is daily consulted as to the
relation of albuminuria and life assurance. As liis function is to
protect the interests of the company, he should reject all cases in
which albumen occurs in the urine. It is even doubtful if an
exception should be made in young persons with transient
albuminuria. Naturally, companies lay great stress upon the presence
or absence of albumen, but in the most serious and fatal malady
with which they have to deal, chronic interstitial nephritis, the
albumen is often absent or transient, even when the disease is well
developed. After the fortieth year, from a standpoint of life
insurance, the state of the arteries is far more important than the
condition of the urine. IV. Pyukia [Pus in the Urine). Causes. — (1)
Pyelitis and Pyelonephritis. — In large abscesses of the kidney,
pyonephrosis, the pus may be intermittent, and for days or even
weeks the urine is free. In calculous and tuberculous pyelitis the
pyuria is usually continuous, though varying in intensity. In these
cases, as a rule, the pus is mixed with the urine, which is acid in
reaction. In the early stages of pyelitis the transitional epithelium
may be abundant, but is not in any way distinctive. In the pyelitis
and pyelonephritis following cystitis the urine is usually alkaline, and
contains more mucus ; micturition is usually more frequent, and the
history points to a previous bladder affection. (2) Cystitis. — The
urine is alkaline, often fetid, the pus ropy, and the amount of urine
greatly increased. The ropy, thick mucus usually comes with the last
portions of the urine. Triple phosphate crystals may be present in
the freshly passed urine. (3) Urethritis^ particularly gonorrhoea. The
pus appears first, is in small quantities, and there are signs of local
inflammation. (4) In leucorrhoea the quantity of pus is usually small,
and large flakes of vaginal epithelium are numerous. In doubtful
cases, when leucorrhoea is present, the urine should be withdrawn
by a catheter. (5) Rupture of Abscesses into the Urinary Passages. —
In such cases as pelvic or perityphlitic abscess there have been
previous symptoms of pus formation. A large amount is passed
within a short time, then the discharge stops abruptly or rapidly
diminishes within a few days. Pus gives to the urine a white or
yellowish-white appearance. On settling ther(! is a heavy grayish
sediment, and the supernatant fluid is usually turbid, '^^riie
sediment is often tenacious and ropy. The reaction is generally
alkaline, and the odor may be ammoniacal even when passed.
Examination with the microscope reveals the presence of a large
number
 730 DISEASES OF THE KIDNEYS. der, well formed ; the
protoplasm is granular, and often shows many translucent processes.
The only sediment likely to be confounded with pus is that of the
phospluites ; but it is whiter and less dense, and is distinguished
immediately by microscopical exjimination. With the pus there is
always more or less epithelium from the bladder and pelvis, but
since in these situations the forms of cells are practically identical,
they afford no information as to the locality from which the pus has
come. The treatment of pus in the urine is considered under the
conditions in which it occurs. V. Chyluria — Non-parasitic. This is a
rare affection, occurring in temperate regions and unassociated with
the filaria Jiominis sanguinis. The urine is of an opaque white color ;
it resembles milk closely, is occasionally mixed with blood
(haematochyluria), and sometimes coagulates into a firm, jelly-like
mass. In other instances there is at the bottom of the vessel a loose
clot which may be distinctly blood-tinged. Under the microscope the
turbidity seems to be caused by numerous minute granules — more
rarely oil droplets similar to those of milk. Traces of albumen are
usually present. The amount of urine passed is generally increased,
and the chylous condition is intermittent. It may persist for years
without deterioration of health or evidence of serious disease. Since
the discovery of the Jilaria homiiiis sanguinis it has been incorrectly
held by some that all of the cases of chyluria are of this parasitic
nature. I had an opportunity in Montreal of making a careful study of
a French-Canadian woman, a patient of J. B. McConnell's, who had
had chyluria for more than thirteen years. The urine was quite milky
in color and occasionally mixed with blood. Neither ova nor embryos
were found in the urine or in the blood examined at night. After her
death I was enabled to make a thorough dissection of the abdominal
lymph vessels, which were found perfectly normal. The thoracic duct
was not enlarged, the renal lymphatics were not distended ; the
kidneys were increased in size, but showed no special changes. The
most careful examination of the lymph glands and vessels failed to
reveal the presence of parasites. The pathology of the condition is
unknown. No known remedies have any influence upon the chyluria.
(For parasitic chyluria see Filariasis.) VI. liTTFiriiiA (Lithwmia; Lithic-
acid Diathesis). The amount of uric acid excreted daily depends
greatly upon the diet, ranging from lialf a gramme on a vegetable to
as high, even, as two
 ANOMALIES OF THE URINARY SECRETION. Y31 grammes
on an animal diet. In the urine of herbivora it occurs only in traces.
In that of carnivora it may be absent altogether. On the other hand,
in the urine of birds and reptiles it is the chief nitrogenous
ingredient. As Sir William Roberts remarks, its presence in the
human urine is somewhat of an anomaly, as its place is very much
better taken by urea, which is easily soluble and better adapted to
the mammalian plan of a liquid urine. He regards it as a sort of
vestigial remnant. Place and Mode of Formation of the Uric Acid. —
It is now very generally conceded that uric acid is formed in the
tissues and excreted by the kidneys. It may occur in traces in the
blood even in health. Von Jaksch, who has recently examined the
blood of 109 individuals, found no trace in 9 healthy persons, nor
was it present in cases of typhoid fever or in nervous affections or in
diseases of the liver and gastro-intestinal canal, except when
anaemia coexisted. On the other hand, it was present in connection
with all those diseased processes in ^vhich oxidation was disturbed,
either directly, as in affections of the lungs, such as pneumonia, or
indirectly, as in anaemia, in which the oxygen-carriers are deficient.
According to Haig, the amount in the blood rises and falls with the
degree of alkalinity (as more is held in solution), and all
circumstances which increase this are associated with an increase in
the amount of uric acid. As to the place of formation, the
experimental evidence points strongly to the liver, and, according to
Minkow^ski, it is formed there by the synthesis of ammonia and
lactic acid. The views, however, as to its place of production and the
antecedents are by no means harmonious. Garrod still holds that the
kidneys are concerned not only with its excretion, but with its
formation. On the other hand, Ebstein thinks that it is chiefly
produced in the muscles and in the bone marrow. Nor is it yet
settled whether uric acid is only an intermediate step in the
formation of urea or whether it has an independent origin. Mode of
Elimination. — Uric acid is extremely insoluble, a gramme requiring
for its solution, at ordinary temperature, fourteen litres of water, and
about half that amount at body temperature. In the 1,500 to 2,000
0. c. of urine passed in the day the uric acid could not be dissolved,
but it is eliminated in combination as soluble salts, chiefly as urates
of ammonium and sodium. The power in the blood of holding the
uric acid in solution depends upon the degree of alkalinity ; thus it
lias been long known that the excretion of uric acid some hours after
breakfast is high. This is in what Sir William Roberts calls " the
alkaline tide." Ilaig has shown that this excretion can be increased or
diminished by increasing or diminishing the alkalinity of the blood ;
thus, under salicylate of soda, given in fifteen grain doses tliree
times a day, the excretion of the uric acid is increased on the first
and second days, and subsequently falls to the normal amount. He
explains this by supposing that the salicylate finds a considerable
quantity of uric acid stored in the liver, spleen, and other tissues,
gets this into solution, and the greater part of it is passed in the
urine. His obser 
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