Table Of ContentTHE PHYSIOLOGY OF CRUSTACEA
VOLUME I
METABOLISM AND GROWTH
VOLUME II
SENSE ORGANS, INTEGRATION, AND BEHAVIOR
THE P H Y S I O L O G Y
OF CRUSTACE A
EDITED BY
TALBOT H. WATERMAN
DEPARTMENT OF ZOOLOGY, YALE UNIVERSITY
ι
VOLUME
METABOLISM AND GROWTH
ACADEMCI PRESS ,
NEW YORK A ND LONDON
I960
Copyright ©, 1960, by Academic Press Inc.
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PRINTED IN THE UNITED STATES OF AMERICA
VOLUME I
METABOLISM AND GROWTH
1. GENERAL CRUSTACEAN BIOLOGY. Talbot H. Waterman, Department of
Zoology, Yale University, New Haven, Connecticut, and Fenner A.
Chace, Jr., Division of Marine Invertebrates, U. S. National Museum,
Washington, D. C.
2. RESPIRATION. H. P. Wolvekamp, Zoologisch Laboratorium der Rijksum-
versiteit te Leiden, Leiden, The Netherlands, and Talbot H. Water
man, Department of Zoology, Yale University, New Haven, Connecti
cut.
3. BIOCHEMISTRY OF PIGMENTS. T. W. Goodwin, Department of Biochem
istry, The University of Liverpool, Liverpool, England.
4. BLOOD CHEMISTRY. Marcel Florkin, Laboratoires de Biochimie, l'Uni-
versite de Liege, Liege, Belgium.
5. CIRCULATION AND HEART FUNCTION. Donald M. Maynard, Department
of Zoology, University of Michigan, Ann Arbor, Michigan.
6. FEEDING AND NUTRITION. S. M. Marshall and A. P. Orr, Marine Station,
Millport, Isle of Cumbrae, Scotland.
7. VITAMINS. L. R. Fisher, National Institute for Research in Dairying,
University of Reading, Reading, England.
8. DIGESTION AND METABOLISM. H. J. Vonk, Laboratorium voor Vergelij-
kende Physiologie der Rijksuniversiteit, Utrecht, The Netherlands.
9. OSMOTIC AND IONIC REGULATION. James D. Robertson, Department of
Zoology, The University, Glasgow, Scotland.
10. EXCRETION. Gwyneth Parry, Freshwater Fisheries Laboratory, Ministry
of Agriculture, Fisheries and Food, London, England.
11. TERRESTRIAL ADAPTATIONS. Ε. B. Edney, Zoology Department, University
College of Rhodesia and Nyasaland, Salisbury, Southern Rhodesia.
12. ECOLOGY AND METABOLISM. Marcel Florkin, Laboratoires de Biochimie,
TUniversite de Liege, Liege, Belgium.
13. SEX DETERMINATION. Helene Charniaux-Cotton, Laboratoire de Gene-
tique Evolutive et de Biometrie, Gif-sur-Yvette (Seine-et-Oise), France.
14. INTEGUMENT AND EXOSKELETON. Ralph Dennell, Department of Zoology,
University of Manchester, Manchester, England.
15. MOLTING AND ITS CONTROL. L. M. Passano, Department of Zoology,
Yale University, New Haven, Connecticut.
16. RELATIVE GROWTH. Georges Teissier, Laboratoire de Zoologie, Universite
de Paris, Paris, France.
17. AUTOTOMY AND REGENERATION. Dorothy E. Bliss, Department of Fishes
and Aquatic Biology, American Museum of Natural History, New
York, New York.
ν
VOLUME II
SENSE ORGANS, INTEGRATION, AND BEHAVIOR
1. LIGHT SENSITIVITY AND VISION. Talbot H. Waterman, Department of
Zoology, Yale University, New Haven, Connecticut.
2. MECHANORECEPTION. Melvin J. Cohen, Department of Biology, Univer
sity of Oregon, Eugene, Oregon, and Sven Dijkgraaf, Laboratorium voor
Vergelijkende Physiologie der Rijksuniversiteit, Utrecht, The Nether
lands.
3. CHEMORECEPTION AND THERMORECEPTION. Saul B. Barber, Department
of Biology, Lehigh University, Bethlehem, Pennsylvania.
4. PIGMENTARY EFFECTORS. L. H. Kleinholz, Department of Biology, Reed
College, Portland, Oregon.
5. LIGHT PRODUCTION. E. Newton Harvey, formerly Department of Biol
ogy, Princeton University, Princeton, New Jersey.
6. THE NEUROMUSCULAR SYSTEM. C. A. G. Wiersma, Division of Biology,
California Institute of Technology, Pasadena, California.
7. REFLEXES AND THE CENTRAL NERVOUS SYSTEM. C. A. G. Wiersma,
Division of Biology, California Institute of Technology, Pasadena,
California.
8. NEUROHUMORS AND NEUROSECRETION. John H. Welsh, The Biological
Laboratories, Harvard University, Cambridge, Massachusetts.
9. LOCOMOTION. /. H. Lochhead, Department of Zoology, The University
of Vermont, Burlington, Vermont.
10. KINETIC AND TACTIC RESPONSES. L. Pardi, Istituto e Museo di Zoologia,
Universita di Torino, Torino, Italy, and F. Papi, Istituto di Zoologia
dell Universita, Pisa, Italy.
11. PHYSIOLOGICAL RHYTHMS. F. A. Brown, Jr., Department of Biological
Sciences, Northwestern University, Evanston, Illinois.
12. MIGRATIONS. Richard Bainbridge, Department of Zoology, University of
Cambridge, Cambridge, England.
13. COMPLEX BEHAVIOR. Hermann Schone, Max-Planck-Institut fur Ver-
haltensphysiologie, Seewiesen uber Starnberg, Germany.
14. COMPARATIVE PHYSIOLOGY. Talbot H. Waterman, Department of Zool
ogy, Yale University, New Haven, Connecticut.
vi
PREFACE
This book endeavors to add the Crustacea to the small but growing list
of major animal groups which are beginning to be well known physiologi
cally. Although literally its subject is the special physiology of the arthro
pod class in question, our work may be considered essentially as a con
tribution to comparative physiology, an important, if often somewhat
vaguely defined, field of biology. Such vagueness may be circumvented
pragmatically if the field is defined in terms of what those people do who
claim it as their own. Thus comparative animal physiology can be de
scribed in a heuristic way by citing the research of a few comparative
physiologists (outside our present company) whose work is well known:
Jordan, Winterstein, G. H. Parker, Krogh, Crozier, von Buddenbrock, James
Gray, von Frisch, Munro Fox, Prosser, von Hoist, Pantin, Fontaine, Bul
lock, and Zeuthen. Their research is broadly characterized by the kind of
physiological approach that we have wished to follow in this volume.
Prosser has suggested that comparative physiology is that part of physi
ology which uses the organism studied as an important experimental
parameter. In so doing it seeks out the functional analogies and homologies
to be found among living things and attempts to correlate them in terms
of some basic pattern such as their adaptive significance or phylogenetic
origin.*
In this sense comparative animal physiology is a borderline field lying
between physiology, as it is commonly practiced, and zoology. As such it
provides the stimuli of freshness and panoramic scope that often character
ize the areas between conventional disciplines; yet it sometimes seems to
lack its fair share of attention and understanding just because it does cut
across the traditional organization of biology and because it has the in
herent difficulty of any subject which encompasses more than one complex
area of science. These drawbacks are at once the cause and, in part, the
* A more explicit analysis of comparative physiology and an evaluation of the
present work's contributions thereto are presented in Chapter 14, Vol. II.
vii
viii PREFACE
effect of the scattered and fragmentary state of present knowledge in this
field. The organization and synthesis of such information is a formidable
task but one whose accomplishment is essential for the understanding of
general principles and the direction of future fruitful experiment.
One may protest that treating the special physiology of various system
atic groups of animals in succession is not the procedure most likely to
result in a rapid, effective development of comparative physiology. In a
sense this may be valid, but there are two important points to consider in
refuting such an objection. To begin with, so large and varied an assem
blage of animals as the Crustacea provides within the class itself a broad
scope for comparative physiology much as do the insects and the verte
brates. In addition the breadth of the entire field for all animals is so vast
that a fully adequate synthesis may not be possible until the way is pre
pared by a considerable series of studies of more restricted extent.
With these ideas in mind we have written a book intended for many
biologists. Most directly we have addressed advanced students and research
workers who are already interested in problems of comparative physiology.
Beyond this we have aimed at stimulating those whose outlook might profit
from developing a new interest in this field: physiologists who may have
forgotten the fruitfulness of the comparative method for William Harvey
or Johannes Muller, and zoologists for whom a fresh insight into the
functional significance of familiar systems may produce a valuable stimu
lus. Perhaps we may dare hope that in providing a coherent review of the
physiology of Crustacea we may also succeed in epitomizing some of the
basic principles of the biological sciences.
Every reasonable effort has been made to keep the coverage of our book
up to date even though its preparation has extended over three years. Ex
tensive alterations in galleys were made by many authors in 1958 and
important new material was incorporated in page proofs during the first
half of 1959. While the very latest developments are sometimes considered
the only ones of interest in science, this can become a fetish. Biological
revolutions seldom take place overnight since large bodies of facts and
complex correlations are involved in the development of almost all new
ideas in the field. Even casual acquaintance with the work of many nine
teenth century and early twentieth century biologists bears almost em
barrassing witness to their ability to grasp the basic problems and propose
their likely solution.
Inevitably in a book like this hundreds of species and higher taxa of
Crustacea as well as other organisms are cited. To minimize the difficulties
and confusion resulting from different schemes of classification and am
biguous nomenclature for particular species, we have selected a particular
system of classification (presented in detail on pp. 24-26), and used it as
PREFACE ix
consistently as possible. In some cases this scheme is rather different from
those our authors ordinarily employ, but in most instances they have co
operated fully in its use. Also for each species cited what appears to be
its valid scientific name is employed throughout the text. Some authors
would have preferred to use whatever names appeared for a species in the
papers quoted but in the Editor's opinion this has serious drawbacks. If
such a practice is followed the same species would be referred to in differ
ent chapters or even in various parts of the same chapter by more than
one name and quite different forms would be labeled with the same name.
Since adequate identification of entities dealt with is a primary requisite
of any system of knowledge, such nomenclatural confusion must be mini
mized.
While problems of this kind often seem unduly vexing to physiologists
they cannot be solved by ignoring them. The section on systematics in
Chapter 1 has been written with this in mind. In addition a Systematic
Index has been prepared (pp. 613-633). This is cross referenced so that
synonymous names which have been cited from the physiological literature
can be identified with their valid names used here in the text. In many
instances the common names of animals are also given.
Fortunately we have had the benefit of collaboration in these matters
with Fenner A. Chace, Jr., of the U. S. National Museum, who not only
has co-authored Chapter 1 (for which he also prepared the illustrations)
but in addition has acted as a referee on all taxonomic citations. Where
his own knowledge was not first hand, he obtained the advice of various
colleagues expert in the different groups. Among those who assisted in this
way John L. Brooks should be mentioned since he monitored the names
of cladocerans mentioned. In this group particularly, but often in others
such as the crayfishes, the citations in the physiological literature may be
quite inadequate or even patently incorrect in identifying the organism
studied. This emphasizes the key position of taxonomy in biology, a re
lation of particular importance to the comparative physiologist or bio
chemist. Obviously the comparison of things which are not recognizable
or things which are the same is nonsense.
The Editor is indebted to a great many people who have collaborated
on this work. In first place stand the co-authors themselves who have
labored hard and effectively to produce a coherent volume with a reason
ably uniform approach to the subject. The cooperative efforts necessary to
achieve this have repeatedly tested their patience and strained the Editor's
abilities. Then a large number of colleagues and friends, including the
authors, have been most generous in providing advice and helpful criticism
on many phases of the book's preparation from initial planning to final
production. Of these a considerable group has been of major assistance:
χ PREFACE
Dietrich Bodenstein, John L. Brooks, Fenner A. Chace, Jr., H. Munro Fox,
Isabella Gordon, G. Evelyn Hutchinson, Stephen W. Kuffler, Sidnie M.
Manton, Robert J. Menzies, C. F. A. Pantin, Grace E. Pickford, C. Ladd
Prosser, Charles L. Remington, Rolf Slewing, R. E. Snodgrass, and C. A. G.
Wiersma should be particularly thanked. Needless to say the shortcomings
of the final result cannot fairly be attributed to these helpful collaborators
but must ultimately depend on the Editor's stubbornness or ignorance.
In the laborious task of preparing manuscripts and proofs for publication
the Editor is especially grateful to Miss Elizabeth A. Livingston who has
acted as his effective editorial assistant from the beginning. Thanks are
also due on many counts to Miss Jean G. Caldwell, the Editor's Research
Assistant, and to the staff of Academic Press who have been concerned
with this book.
Finally a word of gratitude should be expressed to the Office of Naval
Research which has substantially supported the Editor's general research
program for a number of years. Yale University has also contributed
greatly in time and facilities of many kinds. Without all of these and
others unnamed this book would have been impossible.
TALBOT H. WATERMAN
January, i960
New Haven, Connecticut
CHAPTER 1
TALBOT Η . WATERMAN
FENNER A. CHACE, JR .
GENERAL CRUSTACEAN BIOLOGY
This chapter considers in broad outline the kind of animals crusta
ceans are. The material presented and its emphasis are intended mainly
to introduce the physiologist to the rudiments of morphology and sys-
tematics as well as to the general features of crustacean ontogeny and
phylogeny. Knowledge of these fields is crucial for an effective com
parative physiology. Biophysics and biochemistry may provide an
adequate basis for general physiology, but the comparative field requires
a broader foundation, particularly of a zoological sort. Hence our in
sight may be significantly increased by adequate appreciation of the
structural, developmental, ecological, and evolutionary correlates of
physiology. Obviously such large topics can be but briefly examined
within the scope here appropriate.*
I. DIAGNOSTIC FEATURES OF THE CLASS
Crustacea may be simply defined as the class of arthropods which
includes the lobsters, crabs, shrimps, prawns, barnacles, water fleas,
and their relatives. A more technical definition would concisely de-
* In addition to the more specific citations, valuable general reference to the Class
Crustacea can be found in: Caiman,4 Manton,35 Snodgrass,62 Tiegs and Manton,68
Vandel,70 Waterlot,72 Weber,73 and especially the crustacean sections of W. Kuken-
thal's "Handbuch der Zoologie" (3 (1), pp. 277-1074, de Gruyter, Berlin, 1926-1927),
written by many authors, and of "Bronn's Tierreich." Band 5, Abteilung 1 of the
latter contains the pertinent material, first written by A. Gerstaeker and A. E.
Ortmann (2639 pp., 1866-1901), but since 1938 being completely redone by many
contributors (e.g. references29,30).
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