Lee 1975

Intracellular (Muscle-Fiber) Habitat of Ancylostoma caninum in Some Mammalian Hosts
Author(s): Keun Tae Lee, M. D. Little and P. C. Beaver

Source: The Journal of Parasitology, Vol. 61, No. 4 (Aug., 1975), pp. 589-598
Published by: Allen Press on behalf of The American Society of Parasitologists
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THE JOURNAL OF PARASITOLOGY
Vol. 61, No. 4, August 1975, p. 589-598
INTRACELLULAR (MUSCLE-FIBER) HABITAT OF

ANCYLOSTOMA CANINUM IN SOME MAMMALIAN HOSTS*
Keun Toe Lee, M. D. Little, and P. C. Beaver

Department of Parasitology, Yonsei University College of Medicine, Seoul, Korea, and Tulane University
Medical Center, New Orleans, Louisiana 70112
ABSTRACT: The persistence and precise location of Ancylostoma caninum larvae in tissues of vertebrate
paratenic hosts and the nature of host responses were studied in mouse, cat, and monkey. Mice were
infected percutaneously and examined at various intervals up to 260 days after exposure. Long-persisting
larvae were found only in the muscles. Histologic sections revealed that within 4 hr after exposure
some larvae had migrated through the skin and had entered individual fibers of the underlying muscles.
After the 1st day nearly all larvae found in muscles were within fibers. Granuloma formation and
encapsulation were not observed, suggesting that inside the fibers the larvae produced no direct in-
flammatory reaction. Only diffuse infiltration of inflammatory cells was observed in muscles and
this appeared to be in response to destruction of muscle fibers. Larvae were similarly located in muscles
of a cat and a rhesus monkey examined 16 and 17 days, respectively, after cutaneous exposure. The
histologic changes observed in muscle fibers invaded by A. caninum larvae are similar to those ob-
served in early Trichinella spiralis infections. The larvae of A. caninum lying coiled within the muscle
fibers also bear superficial resemblance to the larvae of T. spiralis. A. caninum larvae were also re-
covered by tissue digestion from muscles of naturally infected dogs, which suggests that larvae reside
in this location in the bitch prior to transfer to the neonate via the milk.
Several workers have demonstrated that the larvae been described. This study was initi-
third-stage larvae of Ancylostoma caninum can ated mainly to clarify these two points and to
persist in the tissues of rodents for relatively further investigate the behavior of A. caninum
long periods of time. Kamioka (1938) and larvae in other mammalian paratenic hosts.
Matsusaki (1951) recovered A. caninum
larvae from the tissues of rats 224 and 377 MATERIALS AND METHODS
days after infection, respectively, and Nichols Infective larvae of Ancylostoma caninum were
(1956) recovered larvae from mice 392 days obtained from 10-day-old, or older, Harada-Mori
after infection. In studies on the migration or charcoal cultures of feces from naturally infected
dogs. The recovered larvae were applied to the
of A. caninum larvae in rodents, it was found shaved or clipped abdominal or thoracic skin of
that by either the oral or the cutaneous anesthetized animals by the gauze pad method of
route of infection almost all larvae persisting Beaver (1955). Albino mice, about 25 g, were
in the body for long periods were located in each exposed to 500 to 1,000 larvae on the ab-
domen and examined at various intervals from 1
the musculature (Inatome, 1932; Matsusaki,
hr to 260 days postinfection. Mice were killed by
1951; Soh, 1958; Kono and Sawada, 1961). ether asphyxiation. From mice examined 1, 2, 4,
In oral infections the larvae apparently mi- 8, or 12 hr after exposure the abdominal skin and
grate through the lungs before reaching the muscles were removed and placed in Bouin's fixa-
tive for subsequent histologic examination. Each
musculature, but in cutaneous infections they
mouse examined at 24 hr or later was skinned,
go directly to the musculature without mi- eviscerated, the head and tail removed, and the
grating to the lungs. The exact location of A. remaining body was divided into right and left
caninum larvae in the muscles of rodents and halves. One half was placed in Bouin's fixative
and the other was separated into 5 parts: neck,
other vertebrates has not been determined, foreleg, thorax, abdomen, and hind leg. Each of
nor has the type of host tissue response to these parts as well as the liver, lungs, and kidneys
from each mouse were finely minced and digested
separately in 0.5% pepsin solution for 4 to 6 hr
Received for publication 16 January 1975.
at 37 C with periodic agitation. The digestion
* Supported in part by grant AI-04919 from the
fluid from each sample was then centrifuged and
NIAID, NIH, Bethesda, Maryland, and by grant
aliquots of the sediment were placed on glass slides
71-207-1 from the China Medical Board, New
York. and examined for larvae at low magnification. The
feet, tail, head, and the remaining visceral organs
Address reprint requests to: M. D. Little, Para- were discarded.
sitology, Tulane University Medical Center, 1430 A mature domestic cat was exposed to about
Tulane Avenue, New Orleans, Louisiana 70112. 35,000 infective larvae on the skin of the upper
589
590 THE JOURNAL OF PARASITOLOGY, VOL. 61, NO. 4, AUGUST 1975
TABLE I. Distribution of Ancylostoma caninum larvae in carcasses of mice at various periods after
percutaneous infection.*
Distribution (% of larvae recovered) t

Day of No. No. larvae
infection mice recoveredt Neck Foreleg Thorax Abdomen Hind leg
1 1 229 1 1 6 89 3
2 1 117 2 3 23 69 3
5 2 477 2 3 31 63 1
10 2 263 1 7 32 54 6
15 2 423 6 10 39 40 5
20 1 76 10 8 46 29 7
30 2 214 11 15 37 29 8
70 1 102 10 17 29 31 13
90 1 151 19 7 29 30 15
120 2 306 15 6 32 40 7
200 1 38 18 16 37 21 8
* Mice of about 25 g infected with 500, 700, or 1,000 larvae each.

t Total number of larvae recovered by tissue digestion from the five parts of one-half of the carcass (or carcasses)
of the mouse (or mice) examined on each day.
: Percentages expressed to nearest whole number.
right thorax and examined after 16 days. A juvenile tion of larvae from the abdominal muscles to
rhesus monkey was exposed to about 36,000 larvae muscles in other parts of the body, particu-
on the skin of the lower left abdomen and ex-
larly the anterior parts. From postinfection
amined 17 days later. Each animal was killed
with an overdose of sodium pentobarbital given day 20 to postinfection day 200, the end of
intraperitoneally. From these 2 animals 10-g sam- the experiment, there was little change in the
ples of muscle were taken from each of 5 locations distribution of the larvae in the muscles.
(neck, foreleg, thorax, abdomen, and hind leg), In this experiment, primary interest was on
and were digested in 1% pepsin solution for the the distribution of the larvae that remained in
recovery of larvae as described above. Smaller
samples from each location were placed in 10% the skeletal muscles at various periods of in-
formalin for subsequent histologic examination. fection and not on the percentage of the
Fixed tissues were processed by standard his- inoculum recovered. However, it was evident
tologic procedures, and sections were stained with that in each animal a considerable portion of
hematoxylin and eosin or Giemsa's stain.
the inoculum was not recovered. Larvae ap-
In another experiment, tissues were obtained at
autopsy from 9 dogs, 5 males and 4 females, that parently were lost when, after migrating
had been used for experimental purposes other through the lungs to the intestine, they were
than parasitologic by researchers in other depart- passed out of the body in the feces. The re-
ments of the medical school. The histories of the
dogs were unknown except that they had been sults of the tissue digestion indicate that most
purchased from a supplier in a nearby southern of the migration of larvae through the lungs
state. Samples of 30 to 100 g each were obtained occurred during the early period of infection.
from the thoracic muscles, abdominal muscles,
Nine larvae were recovered from the lungs of
diaphragm, kidneys, liver, and spleen, and each
was digested separately in pepsin solution as a mouse examined after 2 days of infection
described above. and one larva each was recovered from the
lungs of mice examined 5 and 10 days after
RESULTS
exposure, respectively.
The distribution of A. caninum larvae in the
Larvae were also occasionally found in the
skeletal muscles of the mice changed gradu-
liver and kidneys, but always in small num-
ally during the first 15 days of infection, then
bers. Two larvae each were recovered from
remained fairly stable throughout the remain-
the liver of mice examined 1 and 2 days after
der of the experiment (Table I). After 1 day
of infection, 89% of the larvae recovered from exposure, respectively, and one larva each was
the carcass were from the abdominal muscles, recovered from the liver of mice examined
the muscles immediately below the cutaneous 5, 15, and 120 days after exposure, respec-
site of infection. During the subsequent 14 tively. Two larvae were recovered from the
days of infection there was a gradual migra- kidneys of a mouse examined at 2 days, and
LEE, LITTLE, AND BEAVER-ANCYLOSTOMA CANINUM LARVAE IN MUSCLE FIBERS 591
I
*,
.,
I!
,i,
ll
FIGURES 1-4. Histologic sections of abdominal muscles of mice showing Ancylostoma caninum
larvae within muscle fibers at 4 hr to 7 days after percutaneous exposure. HE stain, X 650. 1. 4 hr.
2. 12 hr. 3. 3 days. 4. 7 days.
it>v_ I
i
sb
Ii
.
II
, l.
.7
4t
,*'
I
L . , F-..
I. ~ ??
ki
aP, ' t' * < ' i ,> '..
5
?4
, :. ~. 4,
ing a larva.
larva. Giemsa
Giemsa stain,
stain,XX325.
325.
'V . I I ? dW I i :,1 1
.% .
JLi
-- .
FIGURES 9-12. Histologic sections of muscles of mice showing changes in fibers due to invasion
of A. caninum larvae. HE stain. 9. Thoracic muscle at 7 days of infection showing increase in size and
number of sarcolemmal nuclei. X 325. 10. Abdominal muscle at 70 days of infection showing basophilic
granular alteration of a fiber. X 325. 11. Abdominal muscle at 180 days of infection showing
long chains of sarcolemmal nuclei. X 160. 12. Abdominal muscle at 180 days of infection showing
inflammatory cells (bottom center) along fiber containing a larva. Note degenerating fiber in upper
left of photo. X 325.
4) iS
It
I,
I? ?
Ii;: yEt.
IL J441t*~~
tI : i" "
. I I"r
:i
:
; j
I i I !I l .
, I Jl:
FIGURES 13-16. Histologic sections of muscles of a cat and a rhesus monkey at 16 and 17 days of
infection, respectively. HE stain. 13. Neck muscle of cat showing coiled A. caninum larva within a
fiber. X 650. 14. Abdominal muscle of monkey showing coiled larva within a fiber. X 650. 15.
Neck muscle of cat showing increase in size and number of sarcolemmal nuclei. (Compare with Fig. 9.)
X 325. 16. Neck muscle of cat showing basophilic granular alteration of a fiber and invasion of
inflammatory cells. X 325.
TABLE II. Distribution of Ancylostoma caninum of the sarcolemmal nuclei also occurred and
larvae in the muscles of a cat and a rhesus monkey this was first observed in mice at 5 days of
16 and 17 days, respectively, after percutaneous
infection.
infection. These nuclei usually were enlarged
and could be found in the center of the fibers
No. larvae recovered/10 g muscle* as well as at the periphery (Fig. 9). Invaded
Animal Neck Foreleg Thorax Abdomen Hind leg fibers subsequently underwent basophilic
Catt 49 37 116 28 56 degeneration (Fig. 10). Interstitial invasion
Monkey* 2 2 11 444 21
of affected muscles by inflammatory cells,
primarily neutrophils, and lymphocytes, was
* Larvae recovered by digestion of 10 g muscle from each
location. noted at 24 hr and was present to some extent
t Approximately 35,000 larvae applied to skin of right in all mice examined subsequently. However,
axillary region.
after 5 days eosinophils began appearing in
t Approximately 36,000 larvae applied to skin of lower
left abdomen. the muscles and were prominent thereafter.
Invasion of destroyed muscle fibers by in-
flammatory cells was first observed at 3 days
four larvae were recovered from the kidneys of infection. Numerous mast cells were pres-
of a mouse examined at 30 days of infection. ent in the muscles of mice at 30 days
Since only one cat and one rhesus monkey postinfection and thereafter (Fig. 8). A strik-
were examined after cutaneous exposure, ing feature in older infections was the long
limited information was obtained on the
chains of sarcolemmal nuclei that were present
migration and distribution of larvae in these in many fibers (Fig. 11).
animals. However, although in each animal After the first few days of infection all
larvae moved away from the original site of stages of muscle fiber degeneration were evi-
infection, the migration in the monkey was dent, which indicated that larvae were contin-
markedly less than that in the cat after a ually leaving destroyed fibers and entering
comparable period of time (Table II). new ones. This was true throughout the
Histologic study of the tissues of mice re- period of observation, i.e., 260 days.
vealed that at 1 and 2 hr of infection larvae
In no case was a thickened, cystlike struc-
were restricted to the skin and hypodermis at ture observed around a larva, nor was any
the site of exposure on the abdomen. At 4, 8, larva observed that was immediately sur-
and 12 hr of infection larvae were still located
rounded by inflammatory cells. However, in
in the epidermis, dermis, and hypodermis, but some cases a fiber occupied by a larva was
some had now invaded the underlying muscles nearly surrounded by inflammatory cells
(Figs. 1, 2). Larvae found in muscles at 24 (Figs. 8, 12).
hr and at all subsequent times of examination The histologic features of the muscles of the
were almost always located within individual infected cat and monkey were similar to those
muscle fibers (Figs. 3-8). They were typi- observed in the mice. Larvae were found only
cally in a coiled position within the fiber. within muscle fibers (Figs. 13, 14), and the
Various changes in the invaded muscle various stages of degeneration of invaded fi-
fibers were observed. These ranged from bers were observed (Figs. 15, 16). Accumula-
practically no observable alteration, except for tions of inflammatory cells, mainly neutrophils,
the displacement of myofibrils by the larva, eosinophils, and lymphocytes, were present
to complete destruction of the fiber. Although around affected muscle fibers and in some
the sequence of changes could not be deter- perivascular areas. A few mast cells were ob-
mined with absolute certainty it appeared that
served in the muscles of the monkey but not
the initial change was a loss of striation in the those of the cat.
immediate area around the larva (Figs. 2, 5).
This could be observed in mice examined From the dogs examined for larvae by tissue
within 48 hr after infection. Changes ob- digestion, A. caninum larvae were recovered
served subsequent to this included cloudy from both the thoracic and abdominal muscles
swelling and hyaline degeneration which pro- of four, two males and two females, and from
gressively involved a greater portion of the the thoracic muscles only from a male dog.
fiber (Figs. 3, 4). An increase in the number Usually, one or two larvae per 50 g of muscle
were recovered from the positive dogs. older infections it was not uncommon to find
Toxocara canis larvae also were recovered from the larvae within the fibers. In the present
seven dogs, four males and three females. A study, A. caninum larvae were found within
few T. canis larvae were recovered from the individual muscle fibers as early as 4 hr after
thoracic and abdominal muscles and the liver, cutaneous infection and at later times of ex-
but most were recovered from the kidneys. amination almost all larvae in the muscles were
In one male dog, 48 T. canis larvae were re- found to be within the fibers.
covered from 68 g of kidney. Fewer larvae The intracellular location of A. caninum
were recovered from the kidneys of the other larvae, i.e., in the muscle fiber in the paratenic
dogs. host, is a unique one for a nematode larva.
Few other nematode larvae are known to per-
DISCUSSION
sist in the muscles of their host. Toxocara
It is evident from the distribution of A. canis larvae can, but instead of entering the
caninum larvae in the muscles of mice at vari- individual fibers they reside in fibrous capsules
ous periods after percutaneous infection that that they have induced the host to form around
the larvae tended to migrate away from the them. Norris (1971) reported that the third-
initial site of infection until they were dis- stage larvae of A. braziliense and A. tubae-
tributed throughout the body. This migration forme could persist in the anterior musculature
appeared to occur during the first 20 days of of mice for 18 and 10 months, respectively,
infection, since after that time the distribution but their exact location in the tissues was not
remained rather stable. The factors influencing determined. Later, Norris (1973) reported
the larvae to migrate away from the original that the larvae of A. braziliense which persist for
site of infection are not known. A "crowding long periods in mice are found primarily in the
effect" may play some role in their tendency to salivary glands and nasopharyngeal epithelium.
disperse. The gradual change in the distribu- Apparently, the only other nematode known to
tion of the larvae during the first 15 days of invade the muscle fibers of its mammalian host
infection suggests that larvae migrate directly is Trichinella spiralis. The early larva of T.
through the tissues rather than via the circula- spiralis similarly enters the muscle fiber, but
tory system. unlike the larva of A. caninum it subsequently
Matsusaki (1951) found that at various undergoes tremendous growth and induces the
periods after oral infection in rats the larvae host to produce a capsule around it. On the
were in greater numbers in the anterior part other hand, the histologic changes observed in
of the body but that after cutaneous infection, the muscle fibers of animals infected with A.
on the abdomen, the majority were in the caninum larvae are remarkably similar to those
posterior part of the body. In our studies with described for the early stages of a T. spiralis
mice, the majority of the larvae were found in infection (Drachman and Tuncbay, 1965;
the anterior part of the body in infections of Gould, 1970). Cloudy swelling, hyaline
20 days or more. These results, as well as the changes, basophilic granular alteration, and an
limited information obtained from our experi- enlargement and increase in number of sar-
mental infections of a cat and a rhesus monkey, colemmal nuclei and their displacement to-
indicate that the migratory behavior of A. wards the center of the fiber are seen in both
caninum larvae may differ in different mam- infections. However, the sequence of these
malian paratenic hosts. changes in A. caninum infection is not clear.
Although Inatome (1932) was one of the Ribas-Mujal and Rivera-Pomar (1968) con-
first to study the behavior of A. caninum larvae cluded that in T. spiralis infections such
in the tissues of animals the significance of his
changes were not degenerative in nature but
work was largely overlooked. He demonstrated
were part of a process of redifferentiation in
the presence of larvae in the muscles of mice
in which the fiber adapts itself to the presence
up to 142 days after cutaneous infection and
apparently was the first to observe that the of a larva and its metabolites. They also
larvae can invade the individual muscle fibers. thought that these changes in a fiber were
He stated that in early infections larvae were reversible once the poles of the trichina cyst
located between the muscle fibers but that in had been closed. Whether changes in muscle
fibers invaded by A. caninum larvae are ever the larvae are stored had not previously been
reversible is not known, but it is apparent that demonstrated, but in our studies the recovery
portions of fibers, if not entire fibers, are com- of larvae by tissue digestion from the muscles
pletely destroyed and are subsequently invaded of naturally infected dogs indicates that they
by inflammatory cells. may reside within the muscle fibers of the dog
The period of time that a larva of A. caninum as they do in other mammals that serve as
may spend within a single muscle fiber is not paratenic hosts.
known, although it would appear to be longer
LITERATURE CITED
in older infections than it is during the early
period of infection when the larvae are in the BEAVER, P. C. 1955. Observations on Necator
process of migrating away from the site of in- infections resulting from exposure to three lar-
fection. Unlike the T. spiralis larva, which vae. Rev Iber Parasitol Tomo Extraordinario,
p. 713-721.
eventually becomes encapsulated within the 1969. The nature of visceral larva mi-
muscle fiber it has invaded, the A. caninum grans. J Parasitol 55: 3-12.
larva apparently moves on to another fiber after DRACHMAN, D. A., AND T. O. TUNCBAY. 1965.
the previous one has been destroyed. The The remote myopathy of trichinosis. Neurol-
presence of destroyed fibers and inflammation ogy 15: 1127-1135.
around them in mice infected for as long as GOULD, S. E. (ed.). 1970. Trichinosis in Man
and Animals. Charles C Thomas, Springfield,
260 days is considered to be evidence that the Ill., 540 p.
larvae continue to move from one fiber to
INATOME, T. 1932. (On the behavior and des-
another. The infiltration of inflammatory cells tiny of the hookworm larvae in muscles of
into the muscle tissues is interpreted to be cutaneously infected animals.) Keio Igaku
more of a response to the destruction of the 12: 419-451. (In Japanese.)
KAMIOKA, S. 1938. On the development of
muscle fibers than to the presence of the larva.
Ancylostoma caninum in the body of non-
Inflammatory cells were never observed im- specific host. Keio Igaku 18: 55-70. (In
mediately around a larva although they were Japanese, English summary.)
present around degenerating fibers containing KONO, M., AND T. SAWADA. 1961. Studies on
a larva. Possibly, by residing within muscle hookworm immunity (Ancylostoma caninum).
3. On the migration of larvae in the body of
fibers, larvae escape being detected as "foreign
mouse infected with larvae. Kitakanto Igaku
bodies" by the host. 11: 431-438. (In Japanese, English summary.
The possible role of A. caninum larvae in Abstr. in Excerpta Med, Med Microbiol Im-
human disease has been reviewed by Beaver munol Serol, Vol. 16, Pt. 1, No. 1936, 1963.)
(1969). In addition to their causing cutaneous MATSUSAKI, G. 1951. Studies on the life history
of the hookworm. Part VII. On the develop-
larva migrans, it is also known that they can
ment of Ancylostoma caninum in the abnormal
pass through the lungs and produce pulmonary host. Yokohama Med Bull 2: 154-160.
symptoms. The findings reported here indicate NICHOLS, R. L. 1956. The etiology of visceral
that these larvae may also behave in man as larva migrans. II. Comparative larval mor-
they do in other mammalian hosts, with some phology of Ascaris lumbricoides, Necator
entering and persisting in the muscle after americanus, Strongyloides stercoralis and
oral or cutaneous infection.
Ancylostoma caninum. J Parasitol 42: 363-
399.
The complexity of the life cycle of A. NORRIS, D. E. 1971. The migratory behavior of
caninum was recently demonstrated when it the infective-stage larvae of Ancylostoma bra-
was shown that infection of the newborn pup ziliense and Ancylostoma tubaeforme in rodent
occurs primarily by the ingestion of infective paratenic hosts. J Parasitol 57: 998-1009.
larvae in the colostrum and milk of the mother 1973. Migratory behavior of infective
stage larvae of Ancylostoma species in rodent
(Stone and Girardeau, 1968). It is assumed paratenic hosts. (Abstr.) Proc 9th Int Congr
that in such infections A. caninum larvae are Trop Med Malaria 1: 175-176.
stored in somatic sites in the female dog and RIBAS-MUJAL, D., AND J. M. RIVERA-POMAR. 1968.
then at or near the time of parturition they Biological significance of the early structural
alterations in skeletal muscle fibers infected
migrate to the mammary glands where they
by Trichinella spiralis. Virchow's Arch Abt
enter the colostrum or milk (Stone and Smith, A Pathol Anat 345: 154-168.
1973). The location of the somatic sites where SoH, C. T. 1958. The distribution and persis-
tence of hookworm larvae in the tissues in ninum larvae in dogs. J Parasitol 54: 426-
relation to species and routes of inoculation. 429.
J Parasitol 44: 515-519. , AND F. W. SMITH. 1973. Infection of
STONE, W. M., AND M. H. GIRARDEAU. 1968. mammalian hosts by milk-borne nematode
Transmammary passage of Ancylostoma ca- larvae: A review. Exp Parasitol 34: 306-312.
RESEARCH NOTE . . .
Susceptibility of the "Bush Baby" (Galago crassicaudatus

panganiensis) to Brugia malayi and B. pahangi
The "bush baby," Galago crassicaudatus Three of the four inoculated with B. malayi
panganiensis, is one of 10 subspecies of thick- produced microfilaremia within 3 months; the
tailed galagos. Its natural habitat is the open one with B. pahangi exhibited microfilaremia
and mixed forest and woodland savannah of within 2.5 months. After patency, micro-
Angola and south central Africa down the east filaremia levels reached 870 per 0.2 ml in 12
coast from the equator to Natal. This small weeks for B. malayi and 300 per 0.2 ml in 8
(2 to 3 lb), nocturnal lower primate (pro- weeks for B. pahangi and were still rising.
simian) is insectivorous but is more or less Microfilarial periodicity studied in one B.
omnivorous in captivity. This species re- malayi-infected galago is shown in Figure 1.
produces readily in captivity and has a lon- The pattern is distinctly diurnal (coinciding
gevity record of about 15 years. with the indoor light period), while periodicity
A Galago species has been reported as both of this same strain of B. malayi has been found
a natural and an experimental host for Brugia to be nocturnal in Macaca irus (Edeson, 1959,
patei (Nelson et al., 1962, Trans Roy Soc Trop Ann Trop Med Parasitol 53: 381-387) and in
Med Hyg 56: 202-217). In our laboratory, M. mulatta, the rhesus (Wong, unpublished
G. crassicaudatus panganiensis has been tested data). Thus, several advantages in using this
recently for susceptibility to B. malayi and B. primate model for studies which do not require
pahangi. Four animals were subcutaneously large quantities of blood are obvious: e.g., the
inoculated with 50 to 240 B. malayi infective possibility of sampling maximum microfilarial
larvae (L3) and one with 250 B. pahangi L3. populations during normal working hours and
the ease in handling this small animal as com-
pared to other known experimental African
mf/0.2 ML
primate hosts for B. malayi, namely, the patas
monkey, Erythrocebus patas (Orihel and
Pacheco, 1972, Program abstract, Workshop
on Filariasis, Trop Med and Parasitol Study
Section, U. S.-Japan Cooperative Medical Sci-
ence Program, U. S. Panel of Parasitic Diseases,
Tulane University, New Orleans, Louisiana,
13 September 1972), the green monkey,
Cercopithecus aethiops (Wong, unpublished
data), and the baboon, Papio cynocephalus
(Tullock et al., 1974, Int J Parasitol 4: 109-
110).
I1 G H T
7 10 IPM 4 7PM 10
Ming M. Wong and K. C. Lim, California Primate
FIGURE 1. 24-hr microfilaremia pattern of B. Research Center, University of California, Davis,
malayi in a galago as determined by averaging 6 95616. This work was supported by NIH research
20-mm3 blood counts for each time period, X 10. grants RR 00169 and R22 A109741.

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Intracellular (Muscle-Fiber) Habitat of Ancylostoma caninum in Some Mammalian Hosts

Author(s): Keun Tae Lee, M. D. Little and P. C. Beaver

INTRACELLULAR (MUSCLE-FIBER) HABITAT OF

Keun Toe Lee, M. D. Little, and P. C. Beaver

Distribution (% of larvae recovered) t

* Mice of about 25 g infected with 500, 700, or 1,000 larvae each.

Susceptibility of the "Bush Baby" (Galago crassicaudatus

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