Laboratory Allimals (1976) 10, 119-142.

119

NATURAL INFECTIONS OF GUINEA-PIGS

by
CHARLOTTE RIGBY*
Department of Veterinary Microbiology,
Western College of Veterinary Medicine,
University of Saskatchewan, Saskatoon, Saskatchewan S7NOWO,
Canada

The guinea-pig (Cavia porcellus) is an important laboratory animal

whose susceptibility to a wide range of pathogens has made it extremely use-
ful in research and diagnosis. In the United States about 600000 guinea-
pigs are used each year; only mice and rats are more common (ILAR, 1972).
Guinea-pigs are thought to have originated in the mountains of Peru, and
little is known about the diseases which they suffer in their natural habitat.
In the artificial environment of the animal colony a number of infectious
agents may cause disease problems. These are most likely to occur when
the animals are under stress because of overcrowding, a breakdown in sani-
tation, or abrupt changes in temperature, humidity, or diet. Such infections
may be either sporadic or epizootic, and often produce a high mortality. In
this discussion, they are grouped under the heading 'spontaneous infectious
diseases'. Most of the agents involved have been isolated and well-character-
ized, and methods have been devised for their control. With careful manage-
ment, including routine surveillance to detect and eliminate infected animals,
they need not cause many serious problems. Good management practices
have been described (for example, Lane-Petter & Pearson, ]971; UFAW,
1972) and will not be repeated here. BasicalIy they consist of common sense;
if an infectious disease does cause losses, most often it is due to a lapse of
attention to the management routine.
Inapparent infections pose a different type of problem. In these cases a
balanced relationship exists between parasite and host: the infected host
remains healthy and the pathogen is almost always undetected. However,
when stress such as experimental infection disturbs this balance, the pathogen
may multiply or exert its effects. Infected animals may respond unpredict-
ably to experimental treatments, yielding inconclusive or misleading results.
In recent years the increased recognition of the importance of healthy animals
for diagnosis and research has stimulated progress in identifying and charac-
terizing such infettions (Baker, Cassell & Lindsey, 1971; Irving, 1972).
·Present address: Animal Diseases Research Institute (E), P.O. Box 11300, Postal Station
H, Ottawa, Ontario K2H 8P9, Canada.
120 C. RIGBY

Further knowledge of their pathogenesis seems likely, in some cases at least,

to reveal similarities to human diseases which could make them extremely
useful animal models for research into human infections.
This paper will first of all review the better-known spontaneous infectious
diseases causing losses in guinea-pig colonies. Secondly, it will discuss some
examples of inapparent infections which may be important because of their
(often unrecognized) deleterious effects on experimental results, or because of
their possible relationships to human diseases.

SPONTANEOUS INFECTIOUS DISEASES

It is difficult to estimate the incidence of diseases in laboratory animals,

or their economic impact on diagnostic and research funds. In Britain,
Seamer & Chesterman (1967) conducted a survey to identify the most common
disease problems in laboratory animals. They sent a questionnaire to 46
institutions; 33 responded, representing a total of almost 3000 OOOanimals,
including 161 620 guinea-pigs. The problems considered by the respondents
to be important in guinea-pigs included: respiratory problems (11 laboratories);
'pasteurellosis'-presumably Yersinia pseudotuberculosis infection (8); septic
infections (5); otitis media (5); enteritis (5); coccidiosis (3); urinary tract
disease (3); skin and eye disease (3); genital infections (2); mastitis (2); sal-
monellosis (2); lice (2); helminth infestation (1); and staphylococcal infection
(1). 'Undiagnosed disease' was indicated as a major problem by 14 insti-
tutions. Only 20 laboratories routinely necropsied dead or culled animals.
The authors felt that the responses, besides identifying the more common
infectious disease problems, also indicated serious deficiencies in routine
diagnosis and record-keeping facilities in animal colonies.
In our own institution, we reviewed the routine necropsy records of guinea-
pigs which died or were culled from a conventional, closed breeding colony
(consisting usually of between 300 and 400 animals) in the Animal Resources
Centre of the University during the period July 1971 to September 1974. The
principal findings or presumed cause of death were classified as shown in
Table 1. Of 88 guinea-pigs necropsied, 70 (80 %) presumably suffered in-
fection of some kind. Pneumonia was by far the most common problem
(39 animals) and Diplococcus pneumoniae the most common pathogen (17
isolates).
During this period an outbreak of Salmonella typhimurium infection occurred
(Olfert, Ward & Stevenson, 1976). The animals affected or killed during this
epizootic have not been included in Table 1.
Table 2 lists some of the more common infectious disease problems of
conventional guinea-pig colonies, and includes their associated aetiological
agents, common sources of infection, and methods for their diagnosis and
control.
 NATURAL INFECTIONS OF GUINEA-PIGS 121

Table 1. Principal findings at necropsy of 88 guinea-pigs which died or were culled

from the Animal Resources Centre, University of Saskatchewan, guinea-pig colony
from July 1971 to September 1974.
Infectious Non-infectious
findings and no. animals findingsandno.anima~
Pneumonia'" 39 Malocclusion 4
Lymphadenitis and abscessest 7 Pregnancy difficulties 3
Enteritis 5 Pregnancy toxaemia 2
Dermatomycosis 4 Alopaecia 2
Peritonitis 4 Nutritional deficiencies 2
Metritis 3 Congenital abnormalities 2
Otitis media 2 Tumour 1
Nephritis 2 LeuJcaemia 1
Septicaemia 2 Undiagnosed 1
Myositis 1

70 (80%) 18 (20%)

'"Diplococcus pneumonia was cultured from ]6, Pasteurella hemolytica from 1.

t Streptococcus zooepidemiclls was cultured from ], Diplococcus pneumoniae from 1.

Bacteria
Bacterial infections are by far the most common and serious of the diseases
affecting laboratory animal colonies. Without exception, diagnosis depends
on the isolation and identification of the causal agent. For the necessary
bacteriological techniques interested readers are referred to such excellent
manuals as those of Osbaldiston (1973) and Carter (1973).
The nomenclature of many important bacterial pathogens is still unsettled;
we follow that given in Bergey's manual of determinative bacteriology (Buchanan
& Gibbons, 1974).

Bordetella
Bordetella bronchiseptica is probably the most common cause of acute
pneumonia in the guinea-pig (Neil, 1972). Subclinical infection is common,
and the organism may easily be cultured from the nares and trachea of normal
animals. Of 120 healthy guinea-pigs examined by Yoda, Nakagawa, Muto
& Imazumi (1972), 119 carried B. bronchiseptica in the nasal cavity for a
month or longer. Acute pneumonia may result when this 'normal' host-
parasite relationship is disturbed. In one such epizootic, in which 19 % of
the colony's adult breeders died (Woode & McLeod, ] 967), the affected animals
rarely showed any clinical signs before death. At necropsy, an extensive
bronchopneumonia with a reddish-brown pleuritic exudate was the principal
122 C. RIGBY

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finding. In all cases, cultures of affected lung tissue yielded a pure growth
of B. bronchiseptica. The epizootic was finally controlled by identifying and
sacrificing all nasal carriers and their contacts, then thoroughly disinfecting
their quarters.
Autologous vaccines have been used to control similar epizootics with
encouraging results (Ganaway, Allen & McPherson, 1965; Nikkels & Mullink,
1971; Nakagawa, Yoda, Muto & Imazumi, 1974). Rabbits and rats also
frequently carry B. bronchiseptica in the upper respiratory tract, and cross-
infection may occur-a reason for housing different species of animals in
separate rooms.

Salmonella
Salmonellosis is a serious threat to laboratory animals and to their human
attendants. Both chronic and acute salmonellosis occur as epizootics in
animal colonies; both are a hazard to human health.
In the early stages of an outbreak of acute salmonellosis due to Salmonella
enteritidis, which occurred in a lOOO-animal commercial guinea-pig colony,
only a few animals succumbed after a brief, acute illness or severe diarrhoea
(Fish, Fletch & Butler, 1968). After a month losses suddenly increased to
more than 20 animals a day. Gravid sows showing signs of infection had an
abortion rate of 60 %. S. enteritidis was cultured from the tissues of dead
animals; and treatment with sulphamethazine, neomycin, and nitrofurazone
attempted, unsuccessfully. After 2 months over 500 animals had died and,
during this time, 3 human cases of salmonellosis were traced to this outbreak.
The disease was finally c~:mtrolled by depopulating the colony.
In such acute outbreaks death may occur in a few hours, preceded only by
roughening of the fur, weakness, and mild diarrhoea. Few changes (possibly
only a slightly enlarged spleen) are seen at necropsy. It is important to culture
the organism from affected tissues in order to differentiate salmonellosis from
other acute septicaemias. This is best accomplished by the inoculation of
specimens into an enrichment medium (such as selenite or tetrathionate broth)
in addition to routine bacteriological media.
Chronic salmonellosis causes a general wasting of the affected animals, with
death occurring after several weeks. At necropsy, the spleen, liver and lymph
nodes are enlarged, an ulcerative enteritis is common, and a peritonitis may
be present. As in acute salmonellosis, the chronic form is diagnosed by
isolating the organism from affected viscera.
Several species of Salmonella have been incriminated in epizootic guinea-pig
salmonellosis, including the common animal types S. enteritidis, S. dublin, S.
florida, S. poona, and S. bredeney (Haberman & Williams, 1958a; Neil, 1972).
Wild rodents, newly-introduced stock, and contaminated food are frequent
sources of infection. Animals which recover may remain chronic faecal
 NATURAL INFECTIONS OF GUINEA-PIGS 125

excretors of the organism so that control necessitates prompt removal of all

affected animals and their contacts, followed by disinfection of their quarters.
These measures, combined with regular surveillance of the colony by cultural
examination of pooled faecal samples, are an effective means of control (Olfert
et al., ]976).
In an unusual type of salmonellosis, presenting as a mesenteric lympha-
denitis with no visible systemic effects (Paterson & Cook, 1955), S. limete
(Salmonella group B) was isolated from the infected lymph nodes. It was
not pathogenic for mice, and affected guinea-pigs were still very susceptible
to infection with S. typhimurium, another member of Salmonella group B.
The prevalence of such infection by S. limete in guinea-pig colonies is un-
known.

Yersinia
Yersinia pseudotuberculosis, a common pathogen of rodents, produces 3
distinct syndromes in guinea-pigs (Paterson, 1972). Most common is 'classical'
pseudotuberculosis, with chronic caseous lesions in mesenteric and colonic
lymph nodes. Lymphadenitis, emaciation and diarrhoea are seen; death
occurs after several weeks. Newborns may be infected, either congenitally,
or soon after birth. At necropsy the principal findings are enlarged, caseous
lymph nodes, and focal necrosis in the liver and spleen.
A second form is an acute septicaemic pneumonia. Coughing and rapid
breathing, with death in 24 hours, are the principal signs. Severely congested
lungs are seen at necropsy.
Thirdly, Y. pseudotuberculosis may cause a chronic cervical lymphadenitis,
with the animals remaining in good condition. The danger of this infection
is that the lesions may burst externally, disseminating the organism and spread-
ing the infection.
The organism is readily isolated from the tissues and faeces of infected
animals. Wild birds and rodents are frequent sources of infection. In
Britain, pigeon faeces contaminating greens introduced the organism into a
guinea-pig colony (Paterson & Cook, ]963).
An effective control method consists of regular palpation for enlarged
mesenteric and colonic lymph nodes (Paterson, ]972). In this way, animals
in the early stages of infection may be detected and destroyed before they begin
to excrete the organism in faeces.

Strep tobacill us-Fusobac terium

Streptobacillus moniliformis, a common commensal of the throat of rats,
is said to be a rare cause of cervical lymphadenitis in the guinea-pig (ILAR,
1974). The organism is an aerobic, pleomorphic Gram-negative bacillus.
126 C. RIGBY

Since it is difficult to isolate, an aetiological diagnosis has often been made

when such organisms are seen in Gram-stained smears of pus.
The situation is, however, rather confusing. Guinea-pig strains of S.
moniliformis differ from strains isolated from rats in that they require anaero-
biosis for isolation and are rather inert biochemically (Aldred, Hill & Young,
1974). We have also isolated such strains in our laboratory, and we consider
that they conform more closely to the published descriptions of Fusobacterium
necrophorum (formerly Spherophorus necrophorus) (Buchanan & Gibbons, ] 974).
It would be of interest to characterize such strains further, particularly since
guinea-pig lymphadenitis, if it is truly caused by F. necrophorum, could prove
to be a useful model for the study of infections due to this organism in larger
animals and man.

Klebsiella
Klebsiella pneumoniae, a relatively non-pathogenic organism which is widely
distributed in nature, may cause a highly contagious, fatal pneumonia in
guinea-pigs which are debilitated by sudden changes in diet or environment,
or by experimental manipulation (Paterson, ] 972). Emaciation, laboured
breathing, and purulent nasal discharge are followed by rapid death. A
purulent bronchopneumonia is seen at autopsy, and the organism is easily
recovered from affected tissues.

Pseudomonas
Pseudomonas aeruginosa is a ubiquitous, usually non-pathogenic organism
which may be introduced into laboratory animal colonies by contaminated
drinking water. This organism may cause acute septicemia, either as an
epizootic affecting mainly young guinea-pigs, or as sporadic cases in older
animals. Death occurs rapidly, often in as little as 5 or 6 hours. At necropsy,
the lymph nodes may be slightly enlarged, but few other signs are seen. P.
aeruginosa infections are rare, occurring only when conditions are extremely
insanitary.
Bostrom et al (1969) described an unusual fatal botryomycosis in an adult
guinea-pig which had previously recovered from acute P. aeruginosa septicemia.
The lung exudate seen at necropsy contained typical botryomycotic granules,
from which P. aeruginosa was recovered in pure culture.

Streptococcus
Streptococcus zooepidemicus (Lancefield group C) is associated with several
diseases of guinea-pigs. Subclinical infections of the upper respiratory tract
are common (van der Waaij & van Bekkum, 1967), and the organism probably
enters the circulation most frequently through abrasions of the mucosa.
 NATURAL INFECTlONS OF GUINEA-PIGS 127

Most commonly, S. zooepidemicus causes a chronic cervical lymphadenitis

('lumps'). Fraunfelter, Schmidt, Beattie & Garner (1971) observed this con-
dition over a period of 8 months in a group of 42 strain-2 guinea-pigs, where
10 animals developed 'lumps'; all continued to eat well, and grew at the normal
rate. These affected guinea-pigs were necropsied at regular intervals, and S.
zooepidemicus isolated from the purulent material contained in the affected
lymph nodes. Fibrinopurulent pneumonia, pleuritis, pericarditis and inter-
stitial nephritis were seen at necropsy in some of the animals.
Such lymph node abscesses may rupture and drain spontaneously, recurring
throughout the life of the animal (Elwood, 1971). Occasionally, there may be
an associated arthritis and cellulitis, and pregnant females may abort (Cook,
1969).
Bronchopneumonia due to S. zooepidemicus has been observed in guinea-
pigs subjected to irradiation (van der Waaij & van Bekkum, 1967). The
organism may also cause sporadic cases of acute, rapidly fatal pneumonia
and septicemia. Haemoglobinuria is often present, and is pathognomic of
infection with this pathogen (Kunz & Hutton, 1971). Consolidation of the
lungs, congestion of the kidneys, and myocardial necrosis are the principal
findings at necropsy. S. zooepidemicus is easily cultured from the affected
organs, and its isolation is important in order to distinguish this infection
from acute salmonellosis. Control is best achieved by rapid culling of affected
animals and their contacts, and disinfection of their quarters.
Evidence presented by Kunstyr & Matthiesen (1973) suggests that the
route of infection and the type of Streptococcus involved may differ in the two
forms of the disease. 37 of the 39 strains they isolated from sporadic cases
of acute septicaemia produced p-haemolytic, M-phase colonies; they were very
pathogenic for mice. On the other hand, of 8 strains from cases of cervical
lymphadenitis, 5 were a- or y-hemolytic, produced S-phase colonies, and did
not kill mice. Thus, it is possible that this 'disease complex' is really 2 distinct
infections, with different inciting agents and epidemiology.
Mastitis in 2 guinea-pigs was described by Gupta, Langham & Conner
(I 970), who isolated an a-hemolytic Streptococcus sp. from the mammary
secretions. These workers later investigated the flora of the normal guinea-
pig mammary gland (Gupta, Carter, Langham & Conner, 1972), and isolated
cr- or p-haemolytic streptococci from 22 of 298 mammary tissue samples. A
later report characterized 35 streptococcal isolates from mammary glands and
milk of normal females (Gupta & Stark, 1973). A strain each of Lancefield
groups B, C, D, and G and 2 isolates belonging to group F were identified;
29 isolates were not groupable. Their possible importance as agents of mastitis
in the guinea-pig is unknown.
Even though torticollis (otitis media) is a common problem in guinea-pigs,
little information on its aetiology appeared in the literature until the recent
]28 C. RIGBY

report by Kohn (1974). He examined 3] animals with torticollis, from 3

different commercial breeders. S. zooepidemicus was isolated in pure culture
from ]4, Diplococcus pneumoniae from 10, Klebsiella pneumoniae from 4, and
Escherichia coli from 1. A mixed S. zooepidemicus-Proteus sp. infection was
identified in 1 animal, and 1 yielded a mixed culture of Bordetella bronchi-
septica, Streptococcus sp. (group K), and Staphylococcus aureus.
Kohn infected 6 adult guinea-pigs with S. zooepidemicus. 3 of these died
(torticollis was noted in 1) and 3 were killed 20 days later. The organism was
isolated from the middle ear of all 3 that died, and from 1 of those that was
killed. This led Kohn to suggest that the streptococci may localize preferen-
tially in the middle ear and, since the guinea-pig ear is anatomically similar
to that of the human, and it appears that similar organisms may infect both,
he also suggested that guinea-pig otitis media might be a very useful animal
model for the study of the human infection.

Diplococcus

Diplococcus pneumoniae (especially types 3, 4, and 19) is an important

pathogen of guinea-pigs (Romer, 1962). Colonization of the upper respir-
atory tract appears to be common, and when the animals are stressed by
environmental changes or experimental manipulations, disease may result.
Breeding females may be long-term carriers, since the organism tends to
sequester in small abscesses in the genital tract, sometimes causing uterine
infections, stillbirths, and congenital infections of the young.
D. pneumoniae infection is probably most commonly seen as otitis media,
or as the cause of sporadic cases of pneumonia and septicemia. Epizootics
also occur, such as that reported by Keyhani & Naghshineh (1974) in a colony
of 2400 young guinea-pigs. Ventilation and sanitation had been poor in the
colony and were felt to be important predisposing factors. Clinical signs were
few; affected animals were listless and in poor condition, and died within a
few days. During the first month, 450 animals were lost. At necropsy,
extensive consolidation of the lungs, focal necrosis of the liver, fibrinous
pleuritis, and severe congestion of all organs were observed, and D. pneumoniae
type 19 was cultured from the organs and blood. Treatment with tetracycline
controlled the outbreak.
D. pneumoniae grows readily on blood agar in an atmosphere of 10% carbon
dioxide, but we have found that some strains require incubation in an anaerobic
atmosphere for primary isolation.
Because of the tendency of the organism to sequester in genital abscesses of
breeding females, its eradication from guinea-pig colonies by the use of anti-
biotics is likely to be very difficult.
 NATURAL INFECTIONS OF GUINEA-PIGS 129

Staphylococcus
Staphylococcus aureus has been associated with a variety of sporadic in-
fections in guinea-pigs. Chronic staphylococcal pododermatitis in a group
of 35 adult guinea-pigs was described by Taylor, Wagner, Owens & Stuhlman
(1971). The predisposing factor was thought to be the fact that the animals
were housed in cages with dirty wire flooring.
Gupta, Conner & Meyer (1972) investigated 3 cases of osteoarthritis, also
in older animals, and isolated S. aureus from the lesions of 1.
During experiments in which foot-and-mouth disease virus was being in-
jected into the footpads of guinea-pigs, Tessler (1973) noticed purulent lesions
appearing a few days after the injections. S. aureus was cultured from the
pus; probably it was a skin commensal introduced into the subcutaneous
tissue by the inoculation procedure.
From Germany, Weber (1973) described a sudden, severe outbreak of S.
aureus infection in a commercial guinea-pig colony, which resulted in abortions
and the deaths of pregnant females. In this case, the source of infection was
thought to be the 5-year-old daughter of the owner, who was ill with staphy-
lococcal pneumonia.
Fungi
Trichophyton CUld Microsporum
Explosive outbreaks of ringworm due to Trichophyton mentagrophytes are
not unusual in guinea-pig colonies. Microsporum canis infections are also
common. These dermatomycoses are extremely important because they are
zoonoses. Human cases have been ascribed to contact with infected animals;
in fact, according to Wood (1973), trichophytosis is the most common zoonosis
of laboratory animals in the United States. Other authors, however, consider
that the strains which infect guinea-pigs are not very readily communicable
to man (Clifford, 1973), and that human infection resulting from contact with
infected guinea-pigs is a rare event. In at least 1 case, however, T. menta-
grophytes infection was introduced into a guinea-pig colony by an infected
technician (Loosli, 1967).
Infected animals suffer erythema, loss of hair, and scaling in localized patches
which may spread and coalesce. A Wood's lamp may be helpful in identify-
ing animals infected with M. canis.
For diagnosis, affected skin and/or hairs are collected, cleared in 10%
potassium hydroxide solution, and examined for the presence of fungal hyphae
and spores. A portion of the sample is also inoculated onto Sabouraud's and
mycobiotic agars, and fungal growth identified by its characteristic gross and
microscopic morphology.
Guinea-pigs seem to have a greater inherent susceptibility to ringworm in-
fections than do other laboratory rodents. (Smith, Rush-Munro & McCarthy,
130 C. RIGBY

1969). In New Zealand, none of the 44 mice and 71 rats they examined were
infected with dermatophytes; but out of 88 guinea-pigs they found 5 that were
infected with M. canis and 65 with T. mentagrophytes. Of these, only 18 had
grossly visible lesions, and the rest were asymptomatic carriers.
Since such asymptomatic carriers are so common, and probably are an
important source of infection, the elimination of dermatophytes from an
infected colony is very difficult (Menges, Georg & Haberman, 1957). The
administration of griseofulvin in drinking water (Clifford, 1973), combined
with ruthless culling of infected animals and contacts, would seem to be most
effective.
It would be interesting to compare guinea-pig and human strains of T.
mentagrophytes in order to help determine if infection with this organism is
truly a zoonosis.
Absidia
Mucormycosis of guinea-pigs, caused by the 'perfect' fungi Absidia ramosa
and A. corymbifera, is a benign infection which causes swelling and abscessation
of the mesenteric lymph nodes. Usually no loss of condition results, and the
affected lymph nodes return to normal in 3-4 weeks. The importance of the
infection lies in the necessity of distinguishing it from the more serious lympha-
denitis due to Yersinia pseudotuberculosis or Salmonella sp. This may be
done by demonstrating the fungal elements in 10 % potassium hydroxide
solution mounts of pus. The condition is said to be common in some countries,
presumably as a result of feeding hay contaminated with fungal spores (Neil,
1972). Control, therefore, would seem to be essentially a matter of ensuring
that good food is provided.
Protozoa
Eimeria
Coccidiosis of the guinea-pig, caused by Eimeria caviae, presents occasional
problems, although the parasite is not very pathogenic (Flynn, 1973). Its
incidence in commercial colonies in North America has been estimated at
6% (M. J. Walcroft, personal communication). In a severe outbreak in New
York (Kleeberg & Steenken, 1963), the mortality was 5 % per month. Emacia-
tion and enteritis were the principal signs; at necropsy the caecum was found
to be the site of infection.
Predisposing debilitating factors such as changes of environment or diet
are probably required for the production of clinical disease. Since the oocysts
excreted in the faeces require about 6 days to become infective, regular clean-
ing of pens every 4 or 5 days should control the infection.

Cryptosporidium
Cryptosporidium wrairii, a normal inhabitant of the guinea-pig small 1l1-
 NATURAL INFECTIONS OF GUINEA-PIGS 131

testine, may be pathogenic in some circumstances. Jervis, Merrill & Sprinz

(1966) found it multiplying in the small intestine and associated it with an
outbreak of chronic enteritis. In a later report (Vetterling, Jervis, Merrill
& Sprinz, 1971) these workers characterized the parasite further, and estimated
its incidence in their colony at 30-40 %. However, since reports from other
laboratories are lacking, and since infective oocysts were not demonstrated,
the importance of this parasite as a guinea-pig pathogen remains unknown.

Nematodes
Paraspidodera
Paraspidodera uncinata is the only nematode commonly affecting guinea-
pigs. This caecal worm is of low pathogenicity, and is likely to cause problems
only in animals kept in outside pens (Haberman & Williams, 1958b). Its
presence is diagnosed by finding adult worms at necropsy, or by the detection
of ascarid-type eggs in faeces.

Ectoparasites
Mites
The guinea-pig mite Chirodiscoides caviae is widespread in the United States,
particularly in some commercial colonies (Wagner, AI-Rabiai & Rings, 1972).
The guinea-pigs in which these investigators found heavy mite infestations
showed no signs of discomfort, but they suffered a high morbidity from D.
pneumoniae infection. It was suggested that the parasitism could have been
important either in transmission or as a predisposing debilitating factor.
Infestation of guinea-pigs by the mange mites Mycoptes musculinis of mice,
Sarcoptes scabei of rabbits and other animals, and Notoedres muris of rats
have been observed (Ronald & Wagner, ] 976). In each case the outbreak
originated from contact with infected animals of the normal host species.
While such occurrences are very rare, they serve to emphasize the need to
house different species of animals separately.
The mite Demodex caviae has been found in the conjunctival muscle tissue
of guinea-pigs suffering from a conjunctivitis (Ronald & Wagner, ]976). No
other signs of infection were observed, and the prevalence and importance
of this mite in guinea-pig colonies is unknown.

Lice
The biting lice Gyropus ovalis and Gliricola porcelli parasitize the guinea-pig,
often in association with each other. Usually no discomfort is apparent in
the parasitized animal, even in heavy infestations. Trimenopon jenningsi has
been described from guinea-pigs, but has now become so rare that Ronald
& Wagner (1976) were unable to find it in several thousand animals.
132 C. RIGBY

Control of ectoparasites is difficult in guinea-pigs. Paterson (1972) recom-

mended the use of aerial insecticidal vapours, with 2-3 weeks of continuous
treatment.

Viruses
Lymphocytic choriomeningitis
Lymphocytic choriomeningitis (LCM) virus causes widespread subclinical
infection in mice. The virus may cause serious disease in man, and all
laboratory animals are also susceptible. In the guinea-pig, infection may be
asymptomatic, or the virus may cause a generalized pneumonia or a paralyz-
ing meningoencephalitis (Obeck, 1974). Classical persistent latent LCM
infection, as it is seen in mice, is not believed to occur in guinea-pigs (Neil,
1972).
LCM infection is best diagnosed by virus isolation in LCM-free animals,
and by complement fixation (Blackmore, Guillon & Schwanzer, 1972).
Since mice commonly carry the virus, and ticks and mites have been shown
to transmit it experimentally (Maurer, 1958), arthropod control, optimum
sanitation, and the separation of mice from guinea-pigs are important in
control. Since LCM may cause severe disease in humans, such control
measures are very important.

Parainfluenza I (Sendai)
Parainfluenza type I (PI-I) virus (Sendai virus) has been isolated from all
of the common laboratory rodents, and is of importance because it may also
cause upper respiratory tract infections in man. In mice subclinical infection
is common and may develop into a lethal pneumonia when the animals are
stressed (Blackmore et al., 1972). A similar situation is thought to exist in
guinea-pigs, and outbreaks of pneumonia due to PI-I, with 100 % mortality,
have been reported (Kunz & Hutton, 197]).
Diagnosis requires the isolation of the virus in embryonated eggs or cultured
cells, and the demonstration of antibody by haemadsorption-inhibition
(Blackmore et ai., 1972). Since infection is transmitted by the respiratory
route, separation of mice and guinea-pigs is important in the control of this
infection.

INAPPARENT INFECTIONS

For the purposes of this discussion, we have defined an 'inapparent infection'

as a host-parasite relationship which permits the survival, growth and normal
propagation of both host and parasite. Such infections may be 'subclinical'
(the parasite may be demonstrated by routine procedures) or 'latent' (the agent
is not demonstrable by routine methods). This relationship may be altered
 NATURAL INFECTIONS OF GUINEA-PIGS 133

when changes in the delicate host-parasite balance either enable the host to
rid itself of the parasite, or permit the parasite to cause frank clinical disease.
This ill-assorted group of infections, although rarely associated with clinical
disease, is probably far more important economically than the overt pathogens.
Their unsuspected deleterious effects on experimentally-stressed animals may
lead to unexplained deaths, or to misleading or erroneous results.
It is important to learn more about the incidence, pathogenesis and epi-
demiology of these inapparent infections; first, in order to devise adequate
control methods (if such are necessary or, indeed, possible); second because
of their possible importance as zoonoses; and, third, because in some cases
at least, their similarities to known human infections may make them of value
as animal research models.

Bacteria
Escherichia
Enteropathogenic Escherichia coli (predominantly serotype 0119 :B14) was
isolated from 75 % of healthy adult laboratory animals of several species by
Schiff et al. (1972). These authors suggested that undiagnosed neonatal in-
fections with such strains must surely be common, and could be responsible
for many unexplained deaths in newborns. This interesting hypothesis
deserves to be investigated further, particularly since such infections could
serve as useful models for the investigation of neonatal coli bacillosis of large
animals and humans.

Listeria
Even though human listeriosis is rather rare, its high mortality makes it an
important zoonosis. Listeria monocytogenes commonly causes inapparent in-
fections in many animals, and infected animals may excrete the organism in
faeces. Guinea-pigs and rabbits are, according to Wood (1973), the most
common hosts. Gay & Blood (1967) considered listeriosis the most serious
of all the non-primate zoonoses, precisely because of this propensity to cause
inapparent infection. Opinion, however, is divided, and other workers are
not convinced that listeriosis is a zoonosis at all (Medoff, Kunz & Weinberg,
1971). More research is needed into the epidemiology of this infection in
guinea-pigs and other animals, and its relationship to human infection. Then
if control measures are indeed necessary, appropriate precautions can be
recommended.

Leptospira
Leptospirosis is an important zoonosis, affecting domestic and wild animals
and man. Outbreaks of human leptospirosis have be'en associated with
134 C. RIGBY

mouse and rat colonies. The organism may be introduced into laboratory
animal colonies by infected wild rodents which excrete the organism in urine.
Once established the infection is difficult to eradicate, since recovered animals
may remain carriers indefinitely (Wood, 1973).
Leptospirosis is diagnosed in the laboratory partly by the inoculation of
guinea-pigs. This method, of course, presupposes the absence of spontaneous
infection in these animals. There is, however, a report (Mason, 1937) of a
guinea-pig colony becoming infected by wild rats. Only 1 animal had clinical
jaundice: it died, and Leptospira icterohaemorrhagica was isolated from the
liver and lungs. Other apparently healthy adults of the colony were sub-
sequently found to be refractory to experimental infection, and some of these
had agglutination titres as high as 1 :300 to L. icterohaemorrhagica. In view
of this report, if there is any history of a colony having been exposed to wild
rodents, it should be established serologically that the animals are indeed free
of leptospiral infection before they are used for research or diagnosis.

Protozoa
Klossiella
The renal coccidian Klossiella cobayae is common in guinea-pigs and gener-
ally considered to be nonpathogenic. Of 108 guinea-pigs examined by Hof-
mann & Hanichen (1970) in Germany, 30 % were found to be infected. These
authors, however, observed a chronic nephritis and described degenerative
lesions seen on histopathology. They considered that K. cobayae was respon-
sible for these lesions, and suggested that disease due to this coccidian may
be more common than is generally supposed.

Pneumocystis
Pneumocystis carinii is an important cause of interstitial pneumonia in human
infants and debilitated adults (Woodruff, 1971). The protozoon was first
described in microscopic sections of the lungs of normal guinea-pigs, and is a
common commensal of rodents and other animals (Faust & Russell, 1964).
The mode of infection of humans is unknown, but presumably sporozoites
derived from carrier animals are inhaled (Audy & Dunn, 1972). Obviously
there is a great need to clarify the epidemiology of this parasite.

Toxoplasma
Spontaneous Toxoplasma gondii infection of guinea-pigs, with infection rates
as high as 33 % (as detected by the dye test) has been reported from the United
States and Europe (Neil, 1972). Inapparent infections are most common;
with stress of some kind, a chronic wasting may be evident, and the affected
animals very gradually become extremely lethargic and die. Prolonged,
 NATURAL INFECTIONS OF GUINEA-PIGS 135

asymptomatic parasitaemia has been observed in experimentally infected

guinea-pigs (Frenkel, 1973). Diagnosis depends on the dye test or immuno-
fluorescence, or on histological demonstration of encysted parasites in the
tissues.
The effect of T. gondii infection upon the response of guinea-pigs to experi-
mental procedures is, according to Shadduck & Pakes (1971), likely to be con-
siderable.
Since cats are the only animals which have so far been shown to excrete
infective oocysts, guinea-pig toxoplasmosis is not likely to present a threat to
human health.
Intrauterine transmission of T. gondii in experimentally-infected guinea-pigs
has been demonstrated (Wright, ] 972). Because the mode of transmission
of the natural infection is still unknown, specific preventive measures have not
yet been formulated.

Encephalitozoon
Encephalitozoon (Nosema) infection may be difficult to differentiate from
toxoplasmosis. No clinical signs are evident, the organism producing a
mildly contagious, chronic infection (Yost, 1958). The mode of transmission
is unknown. Little is known about the incidence or effects of Encephalitozoon
infection in guinea-pigs (Griffiths, 1971), but some colonies of mice and rats
have infection rates of 20-50% (Shadduck & Pakes, 1971). Sporadic cases
have been described in guinea-pigs, the typical lesions in kidney and brain
being detected as incidental findings by histological examination (Moffatt &
Scheifer, 1973). Their importance lies chiefly in the difficulty of interpreting
histological findings when infected animals are used for experimental purposes.
In rabbits, latent infection with E. cuniculi changes the course of some
experimental infections and transplantable tumours (Shadduck & Pakes, 1971).
These authors (Pakes, Shadduck & Olsen, 1972) have developed a diagnostic
skin test for encephalitozoonosis in rabbits, which they claim is sensitive and
specific, and can be used to select un infected rabbits for breeding stock.
Application of this test to guinea-pigs could yield valuable information on the
incidence of this microsporidianin guinea-pig colonies, and afford a means
of obtaining Encephalitozoon-free stock.

Viruses
Cytomegalovirus
Guinea-pig cytomegalovirus (CMV) was first described almost 50 years ago
by Cole & Kuttner (1926) who found typical CMV-type inclusion bodies in
the salivary gland ductal epithelium of 84 % of normal adult guinea-pigs.
Guinea-pigs less than a month old were free of infection.
CMV infection of guinea-pigs is usually inapparent, but severe generalized
136 C. RIGBY

disease may occur in association with some stress factor such as a concurrent
respiratory infection (Blackmore et aI, 1972). Diagnosis is made by virus
isolation in cultured cells, or by the demonstration of typical CMV inclusion
bodies in the epithelial cells of the kidney, liver and bronchi. Transmission
is probably by direct contact, since CMV is excreted in the urine and saliva.
Guinea-pig CMV is similar to other cytomegaloviruses in its replication
cycle in infected cells (Hartley, Rowe & Huebner, 1957) and as seen by electron
microscopy (Middelkamp, Patrizi & Reed, 1967). The disease is important
in the animal colony principally as it may affect the responses of animals
inoculated experimentally with other agents (Cook, 1969). Guinea-pig CMV
infection may serve as a useful model for the study of human CMV infection
(Ditchfield, 1968).

Oncornavirus
The induction of an oncornavirus by 5-bromodeoxyuridine treatment of
cultured guinea-pig fibroblasts and tumour cells was first reported by Opler
(1967). It has since been studied rather intensively by several workers, and
there is an ongoing controversy about its identity as a type B or type C oncoma-
virus (Dahlberg, Perk & Dalton, 1974). It is similar morphologically and
biochemically to the Gross murine leukaemia virus (Type C), but is serologically
distinct (Murray & Nayak, 1974). Hsiung & Fong 1974) isolated a C-type
virus from all guinea-pigs they tested, both normal and leukaemic. The
possible association of this oncornavirus with leukaemia in the guinea-pig,
and its relationship to other human and animal tumour viruses, remain to be
clarified.

Parvovirus
Green (1974) isolated a parvo-like virus from the tissues of guinea-pigs
which had been treated with penicillin. This antibiotic has long been known
to be lethal to guinea-pigs, and Green has put forth the interesting theory that
this effect may be associated with the activation of a latent parvovirus.

Herpes-like virus
Guinea-pig herpes-like virus (HLV) has been studied rather intensively since
its first isolation from guinea-pig kidney cell cultures (Hsiung & Kaplow,
1969). HLV infection in guinea-pigs is inapparent, and the virus seems to
be truly latent, since its isolation required organ culture or co-cultivation
methods (Bhatt, 1971; Lam & Hsiung, 1973). Its incidence varied in different
strains of animals: 90% of 40 strain-2 (leukaemia-susceptible) but none of 40
Hartley strain (leukaemia-resistant) guinea-pigs carried the virus (Hsiung,
Kaplow & Booss, 1971). Experimentally, HLV could be transmitted trans-
 NATURAL INFECTIONS OF GUINEA-PIGS 137

placentally (Lam & Hsiung, 1971a) as well as orally and intranasally (Lam &
Hsiung,1971b).
The similarities of HLV to human herpesviruses led Booss & Hsiung (1971)
to suggest that it may make a valuable animal model for the study of per-
sistent herpesvirus infections in humans.

Other agents
Chlamydia
A guinea-pig inclusion-body conjunctIvItIs agent was detected by Murray
(1964) in conjunctival scrapings from 32 of 144 guinea-pigs tested. Evidence
of infection was very slight, and disappeared spontaneously in about 3 weeks.
The agent grew in embryonated eggs, and was identified as a member of the
Chlamydiae by the presence of the common group antigen. Watson, Mull,
MacDonald, Thompson & Bear (1973) have taken advantage of its similarity
to human trachoma, and used it in their studies on this infection.
Robinson (1969) isolated a chlamydial agent from 4 of 15 normal adult
guinea-pigs. The organism was identified as the ovine enzootic abortion
agent by complement fixation. 15 years previously, guinea-pigs from the
colony had had access to grass paddocks grazed by sheep experimentally in-
fected with this agent, and this was thought to be the original source of the
infection. If so, the inapparent infection had presumably been maintained
in the guinea-pig colony by mother-offspring transmission for 15 years.

Mycoplasma
Mycoplasma of guinea-pigs were first reported by Hill, Blackmore & Francis
(1969), who isolated a new species, later identified as Mycoplasma caviae
(Hill, ]97]0) from 10 of 232 guinea-pigs examined. The isolates, mostly
from the genital tract, were not associated with disease, except possibly with
a metritis in 1 animal. This association suggested a possible relationship to
abortions and other reproductive problems, as have been postulated for myco-
plasmas in other animals (Razin, 1973).
Since then, several other guinea-pig mycoplasmas have been described. In
Germany, M. pulmonis, a pathogen of the uterus of mice and rats (Casillo &
Blackmore, 1972), was isolated from the uterus of all of 80 pregnant and non-
pregnant females (J uhr and Obi, 1970): the significance of this finding is difficult
to assess (Hill, 197Ib). 2 other un-named species were later isolated from the
vaginas of healthy females (Hill, 1971c) and, more recently, Acholeplasma
laidlawii was isolated from the nasopharynx and vagina of guinea-pigs originat-
ing from 4 different colonies (Hill, 1974). Stalheim & Matthews (1975) con-
firmed Hill's observations, and isolated 3 unidentified species of Mycoplasma
from guinea-pigs in the United States. Thus it appears that the guinea-pig,
138 C. RIGBY

like every other animal so far investigated, is host to a variety of species of

Mycoplasma whose pathogenic potential is still largely unknown.

Haemobartonella
The genus Haemobartonella (formerly thought to be a protozoon) is now
included in the order Rickettsiales. Members of this genus infect many.
laboratory and domestic animals. Haemobartonella tyzzeri (caviae) of guinea-
pigs produces an inapparent infection which becomes evident as an acute
haemolytic anaemia after experimental stress such as splenectomy (Griesemer,
1958). The small beaded bacillus-like organisms are found attached to the
plasmalemma of erythrocytes of infected guinea-pigs, but because overt in-
fections are transient and Haemobartonella may be confused with erythrocytic
inclusions, diagnosis by the examination of blood smears may be unreliable. The
infection is best diagnosed by inoculation of known uninfected splenectomized
guinea-pigs. Haemobartonella infection is transmitted by lice and may occur in
utero; Baker et al. (1971) review research implications due to this type of infection
in rats and mice, and it is possible that similar situations exist in guinea-pigs.
If so, then it is clearly important to define the problem and devise adequate
measures to ensure that biomedical research in these animals is not needlessly
complicated by the unsuspected presence of Haemobartonella.

COMMENTS

Good management is crucial in the prevention and control of infectious

disease in the guinea-pig, as in all laboratory animals. Proper housing,
adequate ventilation, sanitation, and temperature and humidity control are
all-important, as is the provision of well-balanced diets.
There is still, however, a need for improved record-keeping, diagnostic
methods, and a better understanding of the epidemiology and pathogenesis
of agents such as Streptococcus zooepidemicus, Diplococcus pneumoniae and
others. The appropriate knowledge, combined with frequent surveillance, will
permit early detection and diagnosis of infection. Then specific measures
can immediately be taken to ensure that animal losses due to infectious disease
are minimal.
Inapparent infections pose a different type of problem, more difficult to
resolve because more difficult to detect. Such infections are likely to become
more important because of their deleterious effects on experimental animals,
similarities to problems in human medicine, and possible zoonotic significance.
Procedures for the establishment of 'germ-free' or 'S.P.F.' animals may not
ensure their freedom from potential pathogens such as Toxoplasma gondii,
Listeria monocytogenes, and herpesviruses, which may be transmitted trans-
placentally. Clearly, more work is needed to improve our understanding of
these infections and their effects on experimental animals.
 NATURAL INFECTIONS OF GUINEA-PIGS 139

ACKNOWLEDGEMENTS

The author gratefully acknowledges the helpful CrItIcIsm received from Dr

C. H. Bigland, Dr F. M. Loew, and Dr E. D. Olfert during the preparation of
this manuscript.

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