The Biology of the Guinea Pig

317 Pages·1976·9.285 MB·English

by  Joseph E. Wagner (Eds.)

CONTRIBUTORS
James Ε. Breazile
Esther M. Brown
Raymond D. Ediger
Michael F. W. Festing
H. L. Foster
James R. Ganaway
Lawrence V. Harper
Richard M. Hoar
Charles E. Hunt
James G. McCormick
Patrick J. Manning
Juan M. Navia
Alfred L. Nuttall
L. R. Robinette
N.C. Ronald
Dudley B. Sisk
Ronald F. Sprouse
G. L. Van Hoosier, Jr.
John M. Vetterling
Joseph E. Wagner
Richard B.Wescott
The Biology of
the Guinea Pig
EDITED BY
Joseph E. Wagner Patrick J. Manning
Department of Veterinary Pathology Department of Laboratory Medicine and Pathology
Veterinary Medical Diagnostic Laboratory and Research Animal Resources
College of Veterinary Medicine Unit of Comparative Medicine
University of Missouri-Columbia University of Minnesota Health Sciences
Columbia, Missouri Minneapolis, Minnesota
ACADEMIC PRESS New York San Francisco London
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COPYRIGHT © 1976, BY ACADEMIC PRESS, INC.
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United Kingdom Edition published by
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Library of Congress Cataloging in Publication Data
Main entry under title:
The Biology of the guinea pig.
"All chapters . . . were delivered in part at one of
three symposia co-sponsored by ACL AM."
Includes bibliographies and index.
1. Guinea-pigs. 2. Guinea-pigs-Diseases.
3. Guinea-pigs as laboratory animals. I. Wagner,
Joseph Ε. II. Manning, Patrick. J. III. American
College of Laboratory Animal Medicine.
[DNLM: 1. Guinea pigs. 2. Animals,
Laboratory. QY50B615]
QL737.R634B56 599Λ3234 75-13096
ISBN 0-12-730050-3
PRINTED IN THE UNITED STATES OF AMERICA
List of Contributors
Numbers in parentheses indicate the pages on which the authors' contributions begin.
James E. Breazile (53), Department of Anatomy-Physiology, Sciences, Minneapolis, Minnesota
College of Veterinary Medicine, University of Missouri- Juan M. Navia (235), Institute of Dental Research and Depart-
Columbia, Columbia, Missouri ment of Comparative Medicine, School of Dentistry and
Esther M. Brown (53), Department of Anatomy-Physiology, Medicine, University of Alabama, Birmingham, Alabama
College of Veterinary Medicine, University of Missouri- Alfred L. Nuttall (281), Kresge Hearing Research Institute,
Columbia, Columbia, Missouri Department of Otolaryngology, University of Michigan
Raymond D. Ediger (5), Litton Bionetics, Inc., Frederick School of Medicine, Ann Arbor, Michigan
Cancer Research Center, Frederick, Maryland L. R. Robinette* (137), Department of Veterinary Pathology
Michael F. W. Festing (99), Medical Research Council Labora- and Office of Laboratory Animal Resources, Washington
tory Animals Centre, Carshalton, Surrey, England State University, Pullman, Washington
H. L. Foster (21), The Charles River Breeding Laboratories, N. C. Ronald (201), Department of Veterinary Parasitology,
Inc., Wilmington, Massachusetts Texas A & M University, College of Veterinary Medicine,
James R. Ganaway (121), Comparative Pathology Section, College Station, Texas
Veterinary Resources Branch, National Institutes of Health, Dudley B. Siskt (63), Department of Anatomy-Physiology,
Bethesda, Maryland College of Veterinary Medicine, University of Missouri-
Lawrence V. Harper (31), Department of Applied Behavioral Columbia, Columbia, Missouri
Sciences, University of California, Davis, California Ronald F. Sprouse (153), Department of Veterinary Micro-
Richard M. Hoar (13, 269), Department of Toxicology, Hoff- biology, College of Veterinary Medicine and Departments
mann-La Roche, Inc., Nutley, New Jersey of Microbiology and Medicine-Dermatology, School of Medi-
Charles E. Hunt (235), Department of Comparative Medicine cine, University of Missouri-Columbia, Columbia, Missouri
and Institute of Dental Research, School of Dentistry and G. L. Van Hoosier, Jr., (137), Department of Veterinary
Medicine, University of Alabama, Birmingham, Alabama Pathology and Office of Laboratory Animal Resources,
James G. McCormick (281), Department of Surgery, Section Washington State University, Pullman, Washington
of Otolaryngology, Bowman Gray School of Medicine, Wake
Forest University, Winston-Salem, North Carolina
Patrick J. Manning (211), Department of Laboratory Medicine
•Present address: Alpine Animal Hospital, Route 3, Moscow, Idaho
and Pathology and Research Animal Resources, Unit of tPresent address: Central Kentucky Animal Disease Diagnostic
Comparative Medicine, University of Minnesota Health Laboratory, Lexington, Kentucky
χ LIST OF CONTRIBUTORS
John M. Vetterling (163), Animal Parasitology Institute, lege of Veterinary Medicine, University of Missouri-
Beltsville Agricultural Research Center, Agricultural Columbia, Columbia, Missouri
Research Service, United States Department of Agriculture, Richard B. Wescott (197), Department of Veterinary Pathol-
Beltsville, Maryland ogy, College of Veterinary Medicine, Washington State Uni-
Joseph E. Wagner (1, 21, 201, 227), Department of Veterinary versity, Pullman, Washington
Pathology, Veterinary Medical Diagnostic Laboratory, Col-
Preface
The American College of Laboratory Animal Medicine vestigators using or considering the use of guinea pigs in
(ACLAM) was founded in 1957 to encourage education, train- research, veterinarians, students of veterinary medicine, and
ing, and research in laboratory animal medicine. Its primary others professionally concerned with the care and management
goals include professional certification of veterinarians in labo- of guinea pigs (supervisory personnel, animal technicians, and
ratory animal medicine (diplomates) and continuing educa- technologists concerned with day to day applied care of re-
tion. This monograph represents a part of a program developed search guinea pigs) as well as commercial producers of guinea
to further the educational goals of ACLAM. pigs and cavy fanciers. The contributors were assembled based
"The Biology of the Guinea Pig" is patterned after the first on their recognition as experts in their specific disciplines.
ACLAM book, "The Biology of the Laboratory Rabbit," With their detailed presentations we have hopefully assembled
edited by S. H. Weisbroth, R. E. Flatt, and A. L. Kraus, an authoritative reference work that will be of interest to
published by Academic Press in 1974. Development of this almost anyone utilizing guinea pigs.
book was formulated on the belief that there was a need for a The strengths of this book lie in its comprehensive coverage
comprehensive review of literature pertaining to the use of the of material related to applied care and management of guinea
guinea pig as a substrate in research. All of the contributions pigs and their diseases. The chapters on guinea pig behavior,
were delivered in part at one of three symposia cosponsored genetics, specific pathogen-free technique, biomethodology,
by ACLAM. The chapters on husbandry, history, physiology, and colony husbandry represent exhaustive and authoritative
behavior, genetics, reproduction, biomethodology, and specific treatises. The reader will also find that noninduced diseases of
pathogen-free and germfree techniques were presented as an guinea pigs have been emphasized in an organized review form.
ACLAM cosponsored seminar at the 23rd Annual Session of They constitute potentially one of the most limiting factors to
the American Association for Laboratory Animal Science in the utility of the guinea pig in biomedical research, which
St. Louis, Missouri, in October of 1972. Chapters on research makes their early detection and elimination or recognition per
uses of guinea pigs were presented at the annual meeting of the se of paramount importance. Other noninduced diseases of
Federation of American Societies of Experimental Biology in guinea pigs are of substantial value as models for the study of
Atlantic City, New Jersey, in April of 1973. Chapters on similar diseases in man.
diseases of guinea pigs were presented at the July, 1973 The compilation of chapters on research uses of the guinea
meeting of the American Veterinary Medical Association in pig proved to be a difficult task. Thousands of research papers
New Orleans, Louisiana. are published annually in which the guinea pig has been the
The goal of ACLAM and the editors has been to produce a experimental subject. We initiated a series of computer-assisted
monograph on the guinea pig useful to the widest possible literature searches of current and recent journals keying only
audience, i.e., the general scientific community including in- on the words "guinea pig." Several of the topics and authors
xii PREFACE
were selected as a result of these searches, based on the tive superiors for allowing us freedom to assemble this mono-
number of citations in which guinea pigs were used in certain graph: Dr. W. H. Eyestone, Chairman, Department of Veteri-
research areas. We perceived use of the guinea pig in nutrition nary Pathology; Dr. L. G. Morehouse, Director, Veterinary
research, otologic research, toxicology, and teratology as the Medical Diagnostic Laboratory; Dr. C. C. Middle ton, Director,
areas of research in which the guinea pig is most widely used, Sinclair Comparative Medicine Research Farm; and Dean K. D.
and selected these topics for review in this book. Obviously Weide, all of the College of Veterinary Medicine, University of
research in dozens of other disciplines could have been Missouri-Columbia, Columbia, Missouri. We acknowledge the
selected as well. editorial assistance of Joel Morganstern and Ellen Maring and
While this represents the first attempt at assembling a mas- the typographical assistance of Sylvia Bradfleld. Last but not
sive amount of authoritative information on the guinea pig per least we are indebted in a special way to all the contributing
se, we look forward to new editions. Certainly, as research use authors who so generously gave of their time and talent in
of the guinea pig changes, topics not included will deserve writing their respective chapters. Furthermore they, like the
emphasis in future editions. We therefore solicit and welcome editors, have signed royalities from this work to the coffers of
suggestions and comments on errors, significant omissions, and ACLAM for further professional and educational missions in
content. Laboratory Animal Medicine.
We particularily would like to express appreciation to the To economize, we have kept illustrations to a minimum in
1972 ACLAM Board of Directors for committing the resources the belief that readers with continued interest in a topic will
and support of ACLAM to this endeavor and for selecting us as consult the original works liberally referenced in this mono-
editors. We acknowledge our indebtedness to our administra- graph.
Joseph E. Wagner
Patrick J. Manning
Chapter 1
Introduction and Taxonomy
Joseph E. Wagner
I. Introduction 1
II. History and Etymology 1
III. Taxonomy 3
References 4
I. INTRODUCTION culinary delight by the natives of the Bolivian and Peruvian
high country, where the animal is killed, scalded, and scraped
to remove the fur and then either roasted or fried whole and
The importance of the guinea pig to medical research is served with a hot sauce known as picante.
symbolized by its synonymity with "experimental subject" in
the English language. One of the earliest records in English
concerning the biology of the guinea pig is that of Goldsmith
II. HISTORY AND ETYMOLOGY
(1791). Lavoisier in 1780 (Lane-Petter, 1963) is commonly
credited with being one of the first to use the guinea pig in
research, having used it to measure heat production. The most extensive inquiry into the history of the guinea pig
Weir (1974) reviewed the origin of the domestic guinea pig outside of South America is that of Cumberland (1886), and
and its interrelationships with other species of cavies. The wild the second edition of that work by Gardner (1913). Cumber-
guinea pig (Cavia apered) is widely distributed in Argentina, land (1886) concluded that the original importers of the cavy
Uruguay, and Brazil (Weir, 1970). Another wild ancestor, C. into Europe were probably responsible for the nomen "guinea
cutleri, is still found in Peru (Paterson, 1972). Cumberland pig." According to Cumberland (1886):
(1886) stated "I do not believe that the Cavy ever existed in
Brazil as a domestic animal, in the sense of being used for Previous to the arrival of Europeans on the West Coast of
South America, the only domestic animals which could be used
food, as stated by some naturalists, or that it was known there
as food were the llama, alpaca, Coy or Guinea Pig, and a bird
at all before the arrival of the Portuguese."
called Tuya, about the size of a large duck. Under these circum-
Wild guinea pigs inhabit open grassland, where they nest and stances, Cavies must have been of great importance as a food
make paths in taller vegetation. They feed in more open areas supply, and are in fact so mentioned by the early writers, who,
however, generally speak of them as rabbits (Conejo), the name
at dawn and dusk (Weir, 1970). They live in a small societies
by which they are still called by those Peruvians and Bolivians
of from several to several dozen individuals. Waterhouse
who speak Spanish.
(1848) related the guinea pig's one useful property was that of Velasco, in his "Historia de Quito," says that the Indians had
banishing rats from its vicinity. The guinea pig is considered a great numbers of Cavies in their houses. We may fairly conclude
JOSEPH E.WAGNER
that they would bestow the same elaborate care in "breeding and causes rouses them to activity. The date of the first edition of
selecting these small rodents as we know they bestowed upon Pennant's "Hist. Quad." is 1781. It is evident that Pennant has
the llamas and alpacas." We are told that these latter animals followed Buff on, who, in this instance at least, is not a safe
were kept in separate flocks of different colours; and if, by any guide. He says: "Colour white, varied with orange and black in
chance, a young one was born of a wrong colour, it was immedi- irregular blotches; no tail; inhabits Brazil; a restless, grumbling
ately moved into a different flock. little animal, perpetually running from corner to corner; breeds
If this system were applied to the Cavies, many distinct varie- when two months old, has from four to twelve young at a time;
ties would be evolved. This careful breeding of the Cavy, and a would be innumerable, but numbers of the young are eaten by
very remote period for its first domestication, are circumstances cats, others killed by the males; are very tender, multitudes of
both needed to account for the extremely artificial colour and them dying of cold." A great part of Pennant's description of the
marking of the animals; and we may also reason inversely, from habits of the Guinea Pig is incorrect. There are many curious
the permanence and obstinate irregularity of the markings, as misstatements in Buffon, and these have been repeated, upon his
well as from the colours, so distinct from any wild form, that the authority, by Pennant and subsequent naturalists, evidently
domestication took place at a very remote epoch. without any practical acquaintance with the subject.
We are at first puzzled to know how this little animal, which is In Peru, the Cavy seems to have been reared running loose in
neither a pig nor a native of Guinea, came by its absurd mis- the kitchens of the Indian houses; and this appears to be still
nomer. This is, however, soon explained, if we consider the done in the more rural parts of the country, where the Guinea
circumstances under which Europeans must have made its ac- Pig still holds its own in the struggle with European cattle and
quaintance. The Spaniards, on their first going into the Peruvian poultry.
markets, would see exposed for sale, in large numbers, a little
animal looking remarkably like a sucking pig, and would give it
It is apparent that Cavia cutleri had been domesticated for
that name, to which they would add the distinctive term, "Indi-
some time by Andean Indians when Pizarro conquered Peru
an," because the early navigators spoke of South America as a
part of the Indies: thus, we have Porco da India, Porcella da around 1530 (Sire, 1968). Up until that time little is known
India, Cochon d'Inde, and Topsell's term, "the Indian Little Pig about the domestication of the guinea pig in Peru; however,
Coney." The name of "Guinea Pig" is of a later date, and was, the Incas, induced by an absence of large meat providing
probably a further confusion of our own, caused by the circum-
animals, used the guinea pig for food and sacrifices to their
stance that we had more traffic with the coast of Guinea than
gods. After the Spanish conquest the domestic guinea pig
with that of South America, and also were accustomed to
consider Guinea as a part of the Indies. The pig-like appearance became established for food and fancy in many places of the
of which I have spoken results from the mode of preparing Spanish Colonial Empire. It was apparently in the sixteenth
Cavies for cooking-namely, by scalding and scraping them in the century that Dutch sailors introduced the small, easily trans-
same manner as we should treat a pig.
ported guinea pig into Europe. From there the guinea pig
The Guinea Pig was first described scientifically by Aldrovan-
spread rapidly to other parts of Europe and the British Isles
dus, and his contemporary, Gesner. Aldrovandus was born A.D.
but failed to gain widespread acceptance as a food source in
1522, and died in 1607. Gesner was born in 1516, and died in
1605. The great victory of Pizarro over the Peruvians, when he the latter countries. As a result of this travel, it acquired many
seized the person of the Inca, and decided the fate of the common names. They were bred for show and fancy and as
country, was gained on 16th November, 1532, shortly after the pets in Europe for up to 300 years before they were intro-
Spaniards arrived in Peru. It is clear, therefore, that the figures of
duced into research. Their introduction into medical research
the Guinea Pig, and the descriptions of them, which still accu-
was a matter of convenience. They probably first reached the
rately represent and describe the animal, must have been done
soon after its arrival in Europe. It is usual to fix this event, United States in the early part of the seventeenth century from
vaguely, as occurring about fifty years after the discovery of Europe as pets and fancy animals. Their movement from
Peru, or about A.D. 1580. continent to continent was sporadic and incidental to other
In Topsell's "Historie of Foure-footed Beastes, from Gesner
travel. Therefore there are no accurate and reliable records of
and Others," 1607, which contains what I believe to be the first
their movement.
English reference to the Guinea Pig, it is called "the Indian Little
Pig Coney." The description, which I conclude is Gesner's, is as The common name of the guinea pig, cavy, probably derives
follows: "Five claws upon a foot behind, and six before: teeth from its generic name, Cavia. However, the native Indian name
like a mouse, and no tail, and the colours variable. I have seen
coüy, is easily formed into cavy or cavia which may have
them all white and all yellow, and also different from both
influenced its initial generic name. It is interesting to note that
these." Topsell's "Gesner" makes a curious mistake as to the
number of claws-mine certainly have only three on a foot people with a "fancy" interest in Cavia porcellus have adopted
behind, and four before. It is clear that the pure white breed the more "scientific" terminology cavy, while the research
existed at this early period, and was, probably, a result of community continues to utilize guinea pig, a term of obscure
Peruvian cultivation under the Incas.
origin.
Pennant, feeling, no doubt, the inappropriateness of the term
The origin of the name guinea pig is indeed obscure. Most
"Guinea Pig," called this animal the "Restless Cavy." This name
European nationalities perceived the animal as a small pig that
is, however, inaccurate, for Cavies, when there is no exciting
cause, such as fear or hunger, are very quiet animals, and will lie arrived from across the sea and adopted fitting names: Meer-
side by side for hours, until one or other of these disturbing schweinchen (little sea pig); cochon de mer (sea pig); lapin de
1. INTRODUCTION AND TAXONOMY 3
Barbarie (Barbary rabbit). Paterson (1972) noted that the Phiomorpha of the Oligocène epoch, Caviidae first appeared in
name guinea may have come from "Guiana" or more simply South America in the Late Miocene (26 to 7 million years ago)
that in another era may have meant "foreign." Could it be that or a later period. More recently Lavocat (1974) reviewed many
the guinea pig was so named because it arrived in Europe via of the osteological features and other characters of the histri-
ships from Guiana, a country in northeastern South America? comorphous rodents and discussed the use of the word "hys-
There is no reference to the domestic cavy ever having been tricomorph." At the same symposium Wood (1974) also dis-
found there. Could it have arrived in the English speaking cussed the evolution of the Old and New World histrico-
world by slave ships via the coast of Guinea in West Africa? morphs, the sciurognathous hystricomorphs and hystricog-
Could it have been named for the coin, guinea, for which it naths. He also discussed the distribution among rodents of
could be purchased in England in the sixteenth and seven- features supposed to characterize the "hystricomorpha."
teenth centuries? Much of the cultivation of the guinea pig as a fancy may be
Sire (1968) suggested that porcellus may have resulted from attributed to the excellent publication of Cumberland (1886)
comparing the animal's nails to small wooden shoes (petits who predicted cultivation of the cavy as a fancy because a
sabots). However, porcellus is the Latin name for "little pig." wire-haired breed, inappropriately named the Abyssinian, had
Also, the guinea pig, when prepared for the table, resembles a appeared in England. Also a long silky-haired cavy had ap-
whole roasted suckling pig. peared in Paris, which was later named Peruvian by T. L.
Obviously the origin and name of the guinea pig as well as Sclater, Esq., of the Zoological Society of London. Daubigny
the history of its introduction into Europe has been the (quoted by Cumberland, 1886) stated that "the Cobaye has
subject of much curiosity. The names by which the common been remarkably modified by cultivation," and "cultivation
domestic guinea pig, Cavia porcellus Linn., is known through- has greatly increased its size." House (1906) expounds at
out the world include: length on the very active fancy cavy industry in England
before 1906.
English Indian Little Pig Coney (TopselPs "Gresner,"
Many laboratory guinea pigs derive from a strain established
1607), restless cavy (Pennant), Domestic cavy
(Cumberland), cavy, guinea pig by Dunkin and Hartley in 1926 (1930). During a life time of
French Cobaye, cochon d'Inde, cochon d'Inde d'An- genetic studies Sewell Wright (1960) founded a number of
gora (Peruvian) inbred strains of guinea pigs. The genetics of the guinea pig are
Spanish Conejillo de Indias
covered in Chapter 8 of this treatise.
Italian Procella de India, porchita da India
Many North American sources of guinea pigs for research,
Portuguese Porquinho da India
German Meeerschwein testing, and teaching are listed in a handbook "Animals for
Dutch Indianach varken Research" compiled and edited by the Institute for Labora-
Spanish Peruvian Conejo, cuis, curso tory Animal Resources (Joseph Henry Building, 2100 Pennsyl-
Peruvian Indian Cotii, couy
vania Avenue, N.W., Washington D.C.), National Research
or Quichua
Council, National Academy of Sciences. Today most of the
Russian Morskaya svinka
guinea pigs raised in the United States are for laboratory use;
Just as the matter of nomenclature is unsettled so is the however, guinea pigs raised for fancy and companion uses are
matter of from which wild species of Cavia was Cavia porcellus also popular. While the path of the guinea pig from barren
descended. Based primarily on morphological features, Water- floor of the earthen hut of the Andean native to the research
house (1848) held that Cavia cobaya or porcellus, C. aperea, laboratory cannot be traced with accuracy, we fully appreciate
and C cuteri might all be placed in the same species. Darwin that this path has involved its use as a staple food, a fancy
(quoted by Detlefsen, 1914) held that C aperea was not an show species, and a pet or companion animal.
ancestor of the domestic guinea pig because a distinct genus of
lice infested each form. Detlefsen (1914) suggested that C.
aperea was more closely related to the domestic guinea pig
HI. TAXONOMY
than C rufescens because the latter gave sterile male offspring
in a cross with the tame guinea pig. Male offspring from C
aperea were fertile. Thomas (1917) discussed 12 species and Taxonomic Outline (after Simpson, 1945)
subspecies of the genus Cavia. Ellermen (1940) reviewed the
family Caviidae. He referenced and listed 17 "named forms" Kingdom-Animal
or species in the genus Cavia. Phylum-Chordata: Animals with notochord and gills
Subphylum—Craniata (Vertebrata): Chordates with organized head
Landry (1957) and Patterson and Pascual (1972) have re-
region
viewed the paleontology of the South American hystrico- Class-Mammalia: Warm-blooded craniates with hair coat. Young
morph rodents. While caviomorphs are descendents of African nourished from mammary glands