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CRC SERIES IN NUTRITION AND FOOD
Editor-in-Chief

Miloslav Rechcigl, Jr.

Handbook of Nutritive Value for Processed Food

Volume 1: Food for Human Use
Volume II: Animal Feedstuffs

Handbook of Nutritional Requirements

in a Functional Context
Volume 1: Development and Conditions of
Physiologic Stress
Volume II: Hematopoiesis, Metabolic Function, and
Resistance to Physical Stress

Handbook of Agricultural Productivity

Volume 1: Plant Productivity
Volume II: Animal Productivity
 CRC Handbook of
Nutritional
Requirements in a
Functional Context
Volume I
Development and Conditions of
Physiologic Stress

Editor

Miloslav Rechcigl, Jr.

Nutrition Adviser and Chief
Research and Methodology Division
Agency for International Development
U.S. International Development Cooperation Agency
Washington, D.C.

CRC Series in Nutrition and Food

Editor-in-Chief
Miloslav Rechcigl, Jr.

Boca Raton London New York

CRC Press is an imprint of the

Taylor & Francis Group, an informa business
First published 1981 by CRC Press
Taylor & Francis Group
6000 Broken Sound Parkway NW, Suite 300
Boca Raton, FL 33487-2742

Reissued 2018 by CRC Press

© 1981 by CRC Press, Inc.

CRC Press is an imprint of Taylor & Francis Group, an Informa business

No claim to original U.S. Government works

This book contains information obtained from authentic and highly regarded sources. Reasonable efforts have been made to publish
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Library of Congress Cataloging in Publication Data

Main entry under title:

Handbook of nutritional requirements in a functional

context.

(CRC series in nutrition and food)

Bibliography: p.
Includes index.
CONTENTS: v. 1. Development and conditions of
physiological stress.—v. 2. Hematopoiesis, metabolic
function, and resistance to physical stress.
1. Stress (Physiology)—Nutritional aspects—Hand–
books, manuals, etc. I. Rechcigl, Miloslav.
II. Series. [DNLM: 1. Nutrition. QU145 H236]
QP82.2.S8H36   599.01   80-19981
ISBN 0-8493-3956-1 (v. 1.)
ISBN 0-8493-3958-8 (v. 2)

A Library of Congress record exists under LC control number: 80019981

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 PREFACE
CRC SERIES IN NUTRITION AND FOOD

Nutrition means different things to different people, and no other field of endeavor
crosses the boundaries of so many different disciplines and abounds with such diverse
dimensions. The growth of the field of nutrition, particularly in the last 2 decades,
has been phenomenal, the nutritional data being scattered literally in thousands and
thousands of not always accessible periodicals and monographs, many of which, fur-
thermore, are not normally identified with nutrition.
To remedy this situation, we have undertaken an ambitious and monumental task
of assembling in one publication all the critical data relevant in the field of nutrition.
The CRC Series in Nutrition and Food is intended to serve as a ready reference
source of current information on experimental and applied human, animal, microbial,
and plant nutrition presented in concise tabular, graphical, or narrative form and in-
dexed for ease of use. It is hoped that this projected open-ended multivolume compen-
dium will become for the nutritionist what the CRC Handbook of Chemistry and Phys-
ics has become for the chemist and physicist.
Apart from supplying specific data, the comprehensive, interdisciplinary, and com-
parative nature of the CRC Series in Nutrition and Food will provide the user with an
easy overview of the state of the art, pinpointing the gaps in nutritional knowledge
and providing a basis for further research. In addition, the series will enable the re-
searcher to analyze the data in various living systems for commonality or basic differ-
ences. On the other hand, an applied scientist or technician will be afforded the oppor-
tunity of evaluating a given problem and its solutions from the broadest possible point
of view, including the aspects of agronomy, crop science, animal husbandry, aquacul-
ture and fisheries, veterinary medicine, clinical medicine, pathology, parasitology, tox-
icology, pharmacology, therapeutics, dietetics, food science and technology, physiol-
ogy, zoology, botany, biochemistry, developmental and cell biology, microbiology,
sanitation, pest control, economics, marketing, sociology, anthropology, natural re-
sources, ecology, environmental science, population, law politics, nutritional and food
methodology, and others.
To make more facile use of the series, the publication has been organized into sepa-
rate handbooks of one or more volumes each. In this manner the particular sections
of the series can be continuously updated by publishing additional volumes of new
data as they become available.
The Editor wishes to thank the numerous contributors many of whom have under-
taken their assignment in pioneering spirit, and the Advisory Board members for their
continuous counsel and cooperation. Last but not least, he wishes to express his sincere
appreciation to the members of the CRC editorial and production staffs, particularly
President Bernard J. Starkoff, Earl Starkoff, Sandy Pearlman, Pamela Woodcock,
Lisa Levine Eggenberger, John Hunter, and Amy G. Skallerup for their encourage-
ment and support.
We invite comments and criticism regarding format and selection of subject matter,
as well as specific suggestions for new data which might be included in subsequent
editions. We should also appreciate it if the readers would bring to the attention of
the Editor any errors or omissions that might appear in the publication.

Miloslav Rechcigl, Jr.

Editor-in-Chief
 PREFACE
HANDBOOK OF NUTRITIONAL REQUIREMENTS
IN A FUNCTIONAL CONTEXT

Nutritional requirements of animal organisms, including those of human beings,

vary depending on the physiological state of the organisms. This accounts for vulner-
ability to malnutrition among certain population groups such as pregnant and nursing
women, young infants, children, the aged, and the sick.
Adaptation and resistance to specific climate and environmental stresses is also de-
pendent on the availability of specific nutrients in appropriate amounts. There are also
subtle differences in the nutritional requirements for specific physiological processes
and body functions as there are differences in the requirements for specific nutrients
by different tissues during their development.
The purpose of this Handbook is to bring together all the available information on
the nutr"tional requirements of animal organisms for specific processes and functions.
This is believed to be the first systematic treatment of nutrition in a functional context.
Apart from furnishing specific nutritional data, this Handbook provides a useful
framework for a comparative physiologist or biochemist searching for commonality
or differences among various biological systems.
 THE EDITOR

Miloslav Rechcigl, Jr. is a Nutrition Advisor and Chief of Research and Methodol-
ogy Division in the Agency for International Development.
He has a B.S. in Biochemistry (1954), a Master of Nutritional Science degree (1955),
and a Ph.D. in nutrition, biochemistry, and physiology (1958), all from Cornell Uni-
versity. He was formerly a Research Biochemist in the National Cancer Institute, Na-
tional Institutes of Health and subsequently served as Special Assistant for Nutrition
and Health in the Health Services and Mental Health Administration, U.S. Depart-
ment of Health, Education and Welfare.
Dr. Rechcigl is a member of some 30 scientific and professional societies, including
being a Fellow of the American Association for the Advancement of Science, Fellow
of the Washington Academy of Sciences, Fellow of the American Institute of Chem-
ists, and Fellow of the International College of Applied Nutrition. He holds member-
ship in the Cosmos Club, the Honorary Society of Phi Kappa Pi, and the Society of
Sigma Xi, and is recipient of numerous honors, including an honorary membership
certificate from the International Social Science Honor Society Delta Tau Kappa. In
1969, he was a delegate to the White House Conference on Food, Nutrition, and
Health and in 1975 a delegate to the ARPAC Conference on Research to Meet U.S.
and World Food Needs. He served as President of the District of Columbia Institute
of Chemists and Councillor of the American Institute of Chemists, and currently is a
delegate to the Washington Academy of Sciences and a member of the Program Com-
mittee of the American Institute of Nutrition.
His bibliography extends over 100 publications including contributions to books,
articles in periodicals, and monographs in the fields of nutrition, biochemistry, physi-
ology, pathology, enzymology, molecular biology, agriculture, and international de-
velopment. Most recently he authored and edited Nutrition and the World Food Prob-
lem (S. Karger, Basel, 1979), World Food Problem: a Selective Bibliography of
Reviews (CRC Press, 1975), and Man, Food and Nutrition: Strategies and Technolog-
ical Measures for Alleviating the World Food Problem (CRC Press, 1973) following
his earlier pioneering treatise on Enzyme Synthesis and Degradation in Mammalian
Systems(S. Karger, Basel, 1971), and that on Microbodies and Related Particles, Mor-
phology, Biochemistry and Physiology (Academic Press, New York, 1969). Dr. Rech-
cigl also has initiated a new series on Comparative Animal Nutrition and was Associ-
ated Editor of Nutrition Reports International.
 ADVISORY BOARD MEMBERS

E. J. W. Barrington Dorothy Hollingsworth

Cornerways The British Nutrition Foundation
Alderton, Tewkesbury Alembic House
Glascow, Scotland London, England

Charles A. Black B. Connor Johnson

Department of Agronomy Department of Biochemistry and
Iowa State University of Science and Molecular Biology
Technology The University of Oklahoma Health
Ames, Iowa Science Center
Oklahoma City, Oklahoma
Ricardo Bressani
Division of Agricultural and Food 0. L. Kline
Science American Institute of Nutrition
Institute of Nutrition of Central Bethesda, Maryland
America and Panama (IN CAP)
Guatemala City, Guatemala Gilbert A. Leveille
General Foods Corporation
Sir David Cuthbertson Tarrytown, New York
Department of Pathology and
Chemistry Margaret Mead (deceased)
University of Glasgow The American Museum of Natural
Glasgow, Scotland History
New York, New York
William J. Darby
The Nutrition Foundation, Inc. EmilM. Mrak
New York, New York Department of Food Science
University of California, Davis
Emanuel Epstein Davis, California
Department of Soils and Plant
Nutrition Anthony H. Rose
University of California, Davis School of Biological Sciences
Davis, California University of Bath
Claverton Down
Leon Golberg Bath, England
Chemical Industry Institute of
Toxicology Howerde E. Sauberlick
Research Triangle Park, North Department of Nutrition
Carolina Letterman Army Institute of Research
San Francisco, California
Earl 0. Heady
Center for Agricultural and Rural NevinS. Scrimshaw
Development Department of Nutrition and Food
Iowa State University of Science and Science
Technology Massachusetts Institute of Technology
Ames, Iowa Cambridge, Massachusetts
 ADVISORY BOARD MEMBERS (Continued)

Charity Waymouth E. M. Widdowson

The Jackson Laboratory Dunn Nutritional Laboratories
Bar Harbor, Maine Cambridge, England

Dr. S. H. Wittower
Agricultural Experiment Station
Michigan State University
East Lansing, Michigan
 CONTRIBUTORS

A. Aschkenasy, M.D., Sc.D. J. W. T. Dickerson, Ph.D.

Centre National de Ia Recherche Division of Nutrition and Food Science
Scientifique Department of Biochemistry
Laboratoire d'Hematologie University of Surrey
Nutritionelle Guilford, Surrey, England
Paris, France

C. Eckhert, Ph.D.
C. H. Barrows, Sc.D. School of Public Health
Section of Comparative Nutrition Division of Environmental and
Gerontology Research Center Nutritional Sciences
National Institute on Aging University of California
Baltimore City Hospitals Los Angeles, California
Baltimore, Maryland

V. R. Edgerton, Ph.D.
T. K. Basu, B.V.Sc., M.S., Ph.D.
Brain Research Institute
Foods and Nutrition Division
Department of Kinesiology
Faculty of Home Economics
University of California
The University of Alberta
Los Angeles, California
Edmonton, Canada

U.N. Bhuyan, M.D. W. P. Faulk, M.D.

Department of Pathology Blond Mcindoe Centre for
All-India Institute of Medical Sciences Transplantation Biology
New Delhi, India East Grinstead, Sussex, England

J. G. Brand, Ph.D. S. Frankova, D.Sc.

Monell Chemical Senses Center and Institute of Psychology
Department of Biochemistry Czechoslovak Academy of Sciences
School of Dental Medicine Prague, Czechoslovakia
University of Pennsylvania
Philadelphia, Pennsyivania
G. Goldspink, Ph.D., Sc.D., F.R.M.S.
·:J. A.. Camphr;;l, Ph.D. Der,a.rt;ncn: ·.'f Zoology
U r::vcrsi~y o; H. u.ll
Protein Res~·~~~ch Lahc:--ato~y
[( ~:·~6~tOi!-:_; ~on-H1!li, Yo;-kshi!-c
Veterans Admini5traiion l\kc.!icu\
E:cgland
Center
Bronx, New York

K. Y. Guggenheim, M.D.
R. K. Chandra, M.D. Department of Nutrition
Department of Pediatrics Hebrew University-Haddasah Medical
University of Newfoundland School
St. John's, Newfoundland Jerusalem, Israel
E. S. E. Hafez, Ph.D. J. J. Jones, M.D.
Departments of Gynecology, 3 Esplanade
Obstetrics, and Physiology Liverpool, England
School of Medicine
Wayne State University J. Kaltenbach, Ph.D.
Detroit, Michigan Department of Biological Sciences
Mount Holyoke College
H. Hagedorn, Ph.D. South Hadley, Massachusetts
Department of Entomology
Cornell University M. Kare, Ph.D.
Ithaca, New York Monell Chemical Senses Center and
University of Pennsylvania
R. E. Hammer, B.Sc. Philadelphia, Pennsylvania
School of Medicine
Wayne State University H. Karunajeewa, Ph.D.
Detroit, Michigan Department of Agriculture
Animal Research Institute
F. X. Hausberger, M.D., Sc.D. Werribee, Victoria, Australia
Department of Anatomical Sciences
Temple University G. T. Keusch, M.D.
School of Dentistry Division of Geographic Medicine
Philadelphia, Pennsylvania Tufts University School of Medicine
New England Medical Center Hospital
W. J. Hayes, Jr., M.D., Ph.D. Boston, Massachusetts
Department of Biochemistry
School of Medicine G. C. Kokkonen, B.A.
Vanderbilt University Section of Comparative Nutrition
Nashville, Tennessee Gerontology Research Center
National Institute on Aging
E. Hietanen, M.D. Baltimore City Hospitals
Department of Physiology Baltimore, Maryland
University of Turku
Turku, Finland 0. Koldovsky, M.D., Ph.D.
Department of Pediatrics
0. Heroux, Ph.D. University of Arizona Health Sciences
Division of Applied Biology Center
National Research Council Tuscan, Arizona
Ottawa, Ontario, Canada
J. LeMagnen, M.D.
L. S. Hurley, Ph.D. College de France
Department of Nutrition Paris, France
University of California
Davis, California John T. Maher, Ph.D.
Altitude Research Division
D. B. Jelliffe, M.P.H. U.S. Army Research Institute of
School of Public Health Environmental Medicine
University of California Natick, Massachusetts
Los Angeles, California
M. Nairn, Ph.D.
R. E. Johnson, M.D., D.Phil. Faculty of Agriculture
Horn of the Moon Enterprises Hebrew University of Jerusalem
Montpelier, Vermont Rehovot, Israel
A. E. Needham, D.Sc. W. J. Stadelman, Ph.D.
Department of Zoology Department of Animal Sciences
Oxford University Purdue University
Oxford, England West Lafayette, Indiana

Y. Ohira, Ph.D.
Brain Research Institute N.C. Stickland, Ph.D.
Department of Kinesiology Department of Veterinary Medicine
University of California University of Edinburgh
Los Angeles, California Edinburgh, Scotland

A. Ornoy, M.D.
G. Stirling, M.D.
Department of Anatomy and
Department of Pathology
Embryology
College of Medicine and Allied Sciences
Hebrew University-Haddasah Medical
King Abdulaziz University
School
Jeddah, Saudi Arabia
Jerusalem, Israel
1. Quarterman, Ph.D., F.R.S.C.,
C.Chem. A. J. H. VanEs, M.D.
Nutritional Biochemistry Department Institute for Livestock Feeding and
Rowett Research Institute Nutrition Research
Bucksburn, Aberdeen, Scotland Lelystad, The Netherlands

R. Rajalakshmi, Ph.D.
M. Vulterinovna, M.D.
Department of Biochemistry
M.S. University of Baroda Institute of Clinical Experimental
Baroda, India Medicine
Prague, Czechoslovakia
J. A. F. Rook, Ph.D., D.Sc.
Agricultural Research Council
London, England I. Wolinsky, Ph.D.
Department of Human Development
M. L. Ryder, M.Sc., Ph.D., F.I.Biol.
and Consumer Sciences
ARC, Animal Breeding Research
University of Houston
Organisation
Houston, Texas
Edinburgh, Scotland
I. M. Sharman, Ph.D., F.R.S.C.
Dunn Nutritional Laboratory D. H. Woollam, M.D., Sc.D.
University of Cambridge and Medical Department of Anatomy
Research Council University of Cambridge
Cambridge, England Cambridge, England
 DEDICATION

To my inspiring teachers at Cornell University-Harold

H. Williams, John K. Loosli, the late Richard H.
Barnes, the late Clive M. McCay, and the late Leonard
A. Maynard. And to my supportive and beloved
family-Eva, Jack, and Karen.
HANDBOOK OF NUTRITIONAL REQUIREMENTS IN A FUNCTIONAL
CONTEXT

Volume I

Development and Conditions of Physiological Stress

Differentiation and Development

Development of Specific Tissues
Conditions of Physiological Stress

Volume II

Hematopoiesis, Metabolic Function and Resistance to Physical Stress

Hematopoiesis
Digestion and Endocrine Functions
Chemical Senses
Metabolism of Foreign Substances
Physical Performance and Behavior
Adaptation and Resistance to Environmental Stress
 TABLE OF CONTENTS

Volume I

DIFFERENTIATION AND DEVELOPMENT

Nutrition and Differentiation in Animals ...................................... 3
Effects of Nutrition on Metamorphosis ....................................... 81
Nutrition, Regeneration, and Repair ......................................... 97
Nutritional Factors in Teratology .......................................... 113
Nutrition and Cancer. .................................................... 149
Nutrition and Senescence ................................................. 169

DEVELOPMENT OF SPECIFIC TISSUES

Nutrition and Bone Formation ............................................. 209
Nutrition and Development of Connective Tissue: Effects of Protein and
Ascorbic Acid Deficiency ................................................. 257
Effects of Nutrition on the Muscles in Mature Animals ........................ 269
Nutrition and Development of Nervous Tissue ............................... 289
Effect of Nutritional Factors on Development of Adipose Tissue ................ 365

CONDITIONS OF PHYSIOLOGICAL STRESS

Nutrition and Maintenance ................................................ 385
Nutrition and Reproduction: Animals ...................................... 397
Nutrition and Reproduction: Man .......................................... 427
Nutrition and Lactation: Animals .......................................... 439
Nutrition and Lactation: Human ........................................... 495
Nutrition and Egg Production ............................................. 503
Nutrition and Wool Growth ............................................... 517

Index .................................................................. 521

Differentiation and Development
 Volume I 3

NUTRITION AND DIFFERENTIATION IN ANIMALS

A. E. Needham

As yet there is relatively little precise information on the relationship between nutri-
tion and differentiation in animals, mainly because of the difficulty in defining differ-
entiation, and so in designing critical experiments and interpreting results. In the ta-
bles, the equation

Differentiation + Growth = Development (Morphogenesis)

is adopted. This implies that differentiation includes all qualitative aspects of devel-
opment, and growth the purely quantitative aspects.
Differentiation includes all differential morphogenetic processes, from the macro-
scopic to the molecular level, including differentially distributed synthesis, production,
cell proliferation, and growth (heterauxesis). It includes all localized processes, such
as differential movements of cells and materials, segregation of constituents, and local
removal of materials, at the various levels. Where possible, the scale of magnitude of
the differentiation, whether at molecular, cellular, tissue, or organ level is specified.
Differentiation occurs at all stages. In most animals even cleavage of the egg is dif-
ferentiative, with virtually no growth. The final phase is sexual differentiation and
reproductive maturation. There are special phases, such as metamorphosis, in some
animals.
There are differentiative components of development in the various types of asexual
reproduction, and in regeneration following trauma, etc. The latter is not sharply dis-
tinguishable, ultimately, from the smaller-scale regenerative processes that maintain
differentiation in animals at all stages of the life cycle. In morphallactic regeneration
there is extensive remodeling of differentiation.
Evidence from situations involving arrest or even reversal of differentiation, e.g.,
neoplasia, is used where instructive.
Nutrients are taken to include all physiologically useful materials, inorganic, and
organic, that must be acquired from outside the animal's synthesizing systems. Also
included, where relevant, are materials that can be synthesized by the animal but are
usually supplemented by contributions from the diet. In such cases, there is no logical
justification for omitting evidence obtained from the internally synthesized quota.
Yolk and other materials stored or sequestered in an egg for the development of the
embryo constitute nutrients for the embryo. In some instances, less sharply segregated
materials also should be considered as nutrients, e.g., in insect metamorphosis, where
no external food is used. Such materials are also relevant because they are the orthodox
nutrients for other stages or for other animals. Most dietary constituents of animals
are materials synthesized and stored by other organisms.
Evidence is used from both orthodox modes of nutrition and from experimental
methods such as injection, immersion, and force feeding. Much of the evidence has
come from studies of deficiency or excess of dietary constituents, individually or col-
lectively. In addition to causing direct deficiency, the use of metabolic antagonists of
specific constituents has been exploited, though there is a risk of actions in addition
to that of the specific antagonism. Also, there is multilateral interaction between die-
tary constituents, the results of which are still not known in detail.
Axenic culture techniques have been used to distinguish or discount contributions
to an animal's diet by gut microsymbionts. As an expedient, meridic and more fully
defined diets have been used as controls in the experiments.
4 Handbook of Nutritional Requirements in a Functional Context

The following classification of types of action of dietary constituents is assumed:

1. As a source of energy (fuel).

2. As building material (fabric).
3. As a controlling agent:
a. Proximately, as constituent of an enzyme.
b. More indirectly, as hormonal type of agent:
i. Triggering a differentiative response.
ii. Mediating an intermediate step in the response.

DEFINITIONS

Achromotrichia- hair deficient in melanin.

Acrodynia- dystrophy of extremities of body.
Alopecia- deficiency or loss of hair.
Anemia- deficiency of erythrocytes (red blood cells) or of hemoglobin, or both.
Anorexia- loss of appetite.
Ascites- excessive fluid in body cavity.
Axenic diet- one completely free of foreign (usually microsymbiotic) organisms.
Blastema- bud: usually applied to epimorphic regeneration.
Cheilosis- fissures through skin at angles of mouth.
Dermatitis- any morphogenetic abnormality of the skin.
Dystrophy- defect of morphogenetic maintenance (literally "bad feeding").
Edema- excessive extracellular fluid.
Encephalomalacia- softening of brain tissues.
Epimorphosis- regeneration from a bud of undifferentiated cells.
Erythraemia- inflammation (here interpreted as due to morphogenetic defects in the
walls of the capillaries).
Holidic diet- one composed entirely of chemically defined constituents.
Homoeorrhesis- ability to return to a genetically determined path of development.
Hyperchromic, hypochromic, and normochromic anemias- anemias in which the
amount of hemoglobin per cell is respectively higher than, lower than,
and equal to, normal.
Hyperkeratosis- excessive production of keratinized epidermal cells.
Inanition- starvation.
Koilonychia- flattening and longitudinal ridging of finger nails.
Kwashiorkor- dystrophies due to deficiency of protein or of particular essential
amino acids.
Laparotomy- surgical opening of abdominal cavity.
Marasmus- wasting due to simple inanition.
Meridic diet- one composed mainly of chemically defined constituents but including
one or more crude, illdefined items.
Micromelia- abnormal shortness of limbs.
Morphallaxis- regeneration by remodeling of the remaining parts of the body.
Oligidic diet- one mainly composed of crude, natural (biological and inorganic) ma-
terials.
Omphalocephaly- indentation of head (literally "navel-headed").
Osteomalacia- softness of the bones.
Osteoporosis- excessive porosity and lack of compactness of bones.
Parakeratosis- excessive production of epidermal cells that retain nuclei and fail to
mature normally.
Paresis- loss of motor control of muscle; sensation normal.
 Volume I 5

Peresis- atrophy of neurons, with increase in glial tissues.

Perosis- deformity of tibiotarsal joint involving displacement of tendon ("slipped
tendon").
Polycytemia- excessive concentration of cells in blood.
Prophylactic- preventive, anticipatory measure to avoid disease.

INTRODUCTORY NOTES TO TABLES 7 to 39

In assessing the action of a particular dietary constituent on specific processes of

differentiation, it may be necessary to discount certain nonspecific actions on, e.g.,
food consumption, digestion, absorption, and utilization, or on metabolic rate, etc.
In many instances, a deficiency of one particular constituent causes anorexia, thus
symptoms of a general rather than specific deficiency may be predominant.

INTRODUCTORY NOTE TO TABLE 12

Kwashiorkor-A deficiency disease of weaned infants on diets lacking in the total

amount of protein and in relative amounts of essential amino acids. The deficiency
results in loss of appetite (anorexia), and therefore in symptoms of marasmus (general
inanition) as well as in symptoms specific to protein deficiency. However, protein de-
ficiency is responsible for most of the serious symptoms of marasmus. The main dif-
ferentiation abnormalities of kwashiorkor are general retardation or arrest of devel-
opment, hypotrophy of endocrine organs, hyperkeratosis, alopecia,
hypochromotrichia, inflammation of dermis with lesions of the skin, and anemia
(sometimes). A number of the organ systems, including the pancreas and the liver,
first hypertrophy and later hypotrophy; the epidermis and skin melanin also follow
this pattern. The two-phase response could be interpreted as an initial device to com-
pensate for the decrease in performanc of the system, followed by collapse due to
exhaustion. Experimentally, kwashiorkor-symptoms have been induced in rats starved
of only one of a number of the essential amino acids: Trp, ""· " 9 Phe, 159 His, Thr,
Met, 160 Val, or Lys. 161 The symptoms are more pronounced if the animals are force-
fed than when allowed to feed ad lib and so able to refuse gross excess of the other
amino acids (AA).
6 Handbook of Nutritional Requirements in a Functional Context

Table 1
RESPONSE OF DIFFERENTIATION TO VARlO US LEVELS OF FOOD INTAKE

Species Nature of test Result Ref.

High Level of Intake

Man High level during development Accelerated reproductive matura- I, 2

tion
Ambystoma mexican urn Overfeeding during limb regen- Inhibition of histogenesis
(axolotl). eration

High Level at Specific Periods

Pig and other mammals Early postnatal period Promotes muscle production 4,5
Later juvenile period Promotes fat production 4,5
At various periods Differentially increases relative 6, 7
size of organs differentiating in
that period

Different Levels at Specific Periods

Polys tom a (Monoge- Young fluke on young host Rapid maturation as self-fertilizing 8
nea) (abundant food) form
Young fluke on older host, Slower development with change 8
near metamorphosis of site; maturation into cross-fer-
tilizing adult
Ge/is; Pezomachus Larval development in large Macropterous imago with thorax, 9, 10
(Parasitic Hymenop- host(abundantfood) ocelli, etc. different from those of
tera) micropterous form
Development in small host Micropterous form 9, 10
Vespa(wasp) female Low intake early in season Small sterile imagines ("workers") II
larva High intake later in season Large fertile imagines (Queens) 11
Apis(honey bee) female High level (Royal jelly) Large queen imagines 12
larvae throughout
High level for first three days Small sterile workers 12
only
Pheido/e(ant) female Amount of food stored in egg High level: queens; intermediate 13, 14
larvae level: soldiers
Leptothorax(Hyme- High level of larval intake after 940Jo developed into females 15
noptera) hibernation
Low level Only30% developed into females 15
Hirudinaria granulosa Overfeeding on mammalian Failure of reproductive maturation 16
blood of leech
Hirudo medicinalis Undernourishment of young Failure of reproductive maturation 16
Campanularia (Hydro- Overfeeding and underfeeding Both induce casting and regenera- 17
zoa) tive replacement of hydranths
Chrysaora (Scyphozoa) High dietary level, July to Oc- Normal strobilization to produce 18
tober medusae
Subsequent fasting Autotomy of tentacles, each of
which then develops into a minute
new scyphistoma (polyp form)

Note: Caste-determination in Apis and probably in other Hymenoptera depends also on the qualitative
composition of the diet, but at present there is little knowledge of the actions of specific constituents.
The results in items with reference numbers 1 to 9 and 12 are probably direct effects of nutrient level.
In items with reference numbers 10 and 11, the paradoxically similar result of either extreme of
dietary level presumably is an indirect response triggered by any sufficiently unfavorable condition.
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