Inter,,aliond Journalfor Pamimio~y, Vol. 21, No. 4. pp. 431437.

1991
CJ I997 Austr&m Society for Parasitology. Published by Elsevier Scmce Ltd
Pergamon Punted in Great Britain
PII: SOO20-7519(96)00190-7 002&7519/97 $l7.OO+O.CKl

The Effect of Fasting on ASCU& suum and

Oesophagostomum spp. in Growing Pigs
S. PETKEVI&JS,*-i P. NANSENI and L. STEPHENSON5

*Lithuanian Veterinary Institute, LT-4230, KaGiadorys, Lithuania

$Danish Centre for Experimental Parasitology, Department of Veterinary Microbiology, Royal
Veterinary and Agricultural University, 13 Btilowsvej, DK-1870 Frederzksberg, Copenhagen,
Denmark
§Cornell University, Division of Nutritional Sciences, 214 Savage Hall, New York, U.S.A.

(Receked 20 May 1996; accepted 8 Notlember 1996)

Abstract-PetkevRius S., Nansen P. & Stephenson L. 19!Z’. The effect of fasting on Ascaris smmi and
Oesophagostomum spp. in growing pigs. International Journal for ParasitoIogy 27 431-437. Experiments
were conducted to study the possible intiuence of fasting on Ascaris WWII and Oesophagostomum spp. in
growing pigs. Forty young crossbred pigs naturally infected with A. ~IIWIJ and Oesophagostomum spp. were
used. ln one expriment 10 pigs were fasted and offered water ud libitum for 6 days, in another experiment for
10 days. Subsequently, these pigs, together with 10 non-fasted control pigs per experiment were slaughtered,
and worm numbers, worm location, sex, developmental stage and female worm fecundity were determined.
Pigs fasted for 10 but not for 6days had decreased numbers of A. suum and Oesophagostomum spp. at
slaughter vs controls, and worms were found in more distal locations in the gastrointestinal tract. Fasting for
both 6 and 10 days significantly Iowered the fecundity of both worm species. 0 1997 Australian Society for
Parasitology. Published by Elsevier Science Ltd.

Key word.y: Pigs; Ascaris suum: Oesophagostomum dentatum; Oesophagostomum quadrispimdatum; fasting.

INTRODUCTION in populations of wild animals during spells of

drought, in severe winters or in injured animals. In
Over the past years many investigations have studied
domestic animals, starvation may occur because of
the effect of host nutrition on the host’s capability to
sudden shortages of feedstuffs, extreme transport con-
resist parasite infection or to withstand or compen- ditions (Mayes et al., 1988) or when animals suffer
sate for losses brought about by the parasites (cf.
anorexia due to severe disease. In human populations,
Stephenson, 1987). However, diet or nutrition may food deprivation and starvation are unfortunately
exert more direct effects on growth, development and
common, especially in developing countries, e.g. when
pathogenicity of gastrointestinal parasites, simply
crops fail, or in association with wartime, civil dis-
because these are dependent on their hosts for their
turbances and mass migrations.
nutrient supply and habitat requirements (Crompton,
Helminth infections have a widespread occurrence
1991; Solomons & Scott, 1994; Petkevicius er al.,
and significant impact on human and animal hosts
1995).
worldwide. In humans, these infections cause health
Fasting or starvation is an extreme nutritional situ-
problems primarily in tropical and subtropical areas,
ation which leads to general and profound physio- where food shortages and famine are most likely to
logical changes in the host, including radical changes
occur. Apart from studies on naturally infected horses
in the gut habitat. Starvation often occurs naturally
which expelled strongyle nematodes in connection
with starvation (Dvojnos G. M. & Timoshenko 0.
tTo whom correspondence should be addressed. Tel: N., unpublished, Abstracts of Eighth International
+45 35 28 2115: Fax: +45 35 28 2174. Congress of Parasitology, Izmir, Turkey, 1994), little
432 S. Petkevicius et ul

is known about how fasting or starvation may influ- of faeces (epg) was estimated with a modified McMaster
technique (Roepstorff & Nansen, in press). On days 6 and
ence gastrointestinal helminths and severity of disease
10, all animals initially euthanised by CO* were exsangui-
they may produce (Beisel, 1982; Shetty & Shetty, nated. The entire small and large intestines were immediately
1993). removed and separated from the mesenteries. The small
The present investigation on pigs was designed to intestine was divided into 4 sections of approximately equal
examine, under controlled conditions, the influence of length (designated sections 1, 2, 3, 4 from the anterior end).
The contents of each section were collected by pressing luke
fasting on persistence, location and fecundity of two warm water through the intact section twice followed by
commonly occurring nematodes, namely ,4~uri~ suum washing over a sieve with a mesh size of 212 pm. The large
and Oesophugostomwn spp. There are close simi- intestine was divided into 5 sections, designated as follows,
larities between these two parasite species and their starting from the anterior end: (1) caecum; (2) O-20% of the
total length of the colon; (3) 2140%: (4) 41-60% and (5)
related species in man, and in addition, the two hosts
6lllOO%. The sections were opened with scissors and the
have many points of resemblance including gas- contents washed off the mucosal surface. All intestinal con-
trointestinal anatomy and physiology. We expect that tents were examined from fasted pigs, and from control pigs
information obtained in a pig model may be of rel- representative subsamples of 10% were examined. Specimens
evance to man, in which, for obvious reasons, this of A. saurn were weighed, their length measured with a ruler
and the sex and developmental stage were noted, Oeso-
type of experimentation cannot be performed.
phugosfomum spp. worms were collected with an agar-gel
method described by Slotved ef al. (1996). The worms
released from the agar gels were transferred to screw-capped
plastic tubes, fixed and stored in iodine solution (6.25%
MATERIALS AND METHODS iodine + 31.25% potassium iodide + 62.5% distilled water).
Subsequently the samples were decolorized with 3% thio-
E.xperimental design, In two separate experiments (I and sulphate solution and the worms counted with a microscope.
II), pigs were fasted for 6 and lOdays, respectively, and Species and developmental stage were determined by random
compared with control pigs given a standard diet. The pigs collection and microscopical examination of 100 worms from
used originated from a traditional Danish pig farm and were each sample. All worms were examined when there was less
Danish Landrace/Yorkshire/Duroc crosses of both sexes. than 100 worms per sample, The morphological criteria of
After transportation to an experimental animal facility pigs Goodey (1926) and Haupt (1966) for differentiation of 0.
to be fasted were placed individually in small separate pens, dentatum and 0. quadrispinulatum were used. The sex of
whereas pigs fed a standard diet were kept in larger pens, 5 adults of both species was noted.
animals in each. The pens had solid concrete floors and were Srufisfical analysis. The experimental data were analysed
separated by partition walls, approximately 1.5 m high. No using analysis of variance (ANOVA) to determine the effect
bedding was offered since previous observations on fasted of 6 and lOdays of fasting on worm faecal egg counts,
pigs have shown that these ingest considerable amounts of numbers, location, and fecundity of female worms. As an
bedding. Also, any bedding would have made recovery of indication of the average position of the worms along the
expelled worms from the floor very difficult. Before fasting small and large intestine of each pig. the “location” was
started there was an adaptation period of 4 days. The diet of calculated by multiplying the number of worms in each sec-
the control pigs, which consisted of a ground barley plus a tion in each pig by the section number of intestines, divided
protein supplement, was similar to that given in the herd of by the total number of worms in all sections.
origin (PetkeviEius ef al., 1995).
Experimental and control pigs were offered water ad Mi- ,i, I x number of worms in section I
rum. In the pens harbouring fasting pigs, faeces were quan- Location =
Total number of worms in all sections
titatively collected at frequent (4-5 times per day) intervals
during the entire experiment to minimize coprophagy and to Female worm fecundity was estimated by multiplying epg
determine the faecal volume excreted daily. From control by total amount of faeces excreted per day, divided by the
pigs a11 faeces excreted were collected for a 24 hour period adult female worm burden for the relevant worm species.
prior to slaughter. The clinical condition of the pigs was Log-transformed fecundity was examined by analysis of vari-
observed daily by a veterinarian and all pigs were weighed ance to test for the effect of starvation.
at the start of the experiment. and again at slaughter on days epg x weight of faeces per day
Fecundity =
6 and 10. To determine parasite egg concentrations faecal Total number of adult females in all sections
samples obtained directly from the rectum were collected
daily for experiment I and every second day for experiment Ethical considerations. The experiments were approved by
II. the Danish Animal Ethical Committee (animal experiment
Experiment I. Twenty pigs, 2426 weeks old, with a mean permission 1994-101-I 15). Meetings were held with the agri-
body weight of 78 kg (range 71-90 kg) collected on the basis cultural and laboratory technicians to explain the purpose
of faecal egg counts of both helminth species were randomly of the experiments and what was required from the persons
divided into 2 comparable groups of 10 pigs each. The exper- handling the pigs.
imental group was fasted for 6 days. On day 6 all pigs were
killed for postmortem worm determinations. RESULTS
Experiment II. Twenty pigs l&l 1 weeks old. with a mean
body weight of 32 kg (range 28-37kg) were collected and Experiment t (6 day fast)
randomly divided into 2 groups and examined as above, the
experimental group being fasted for 10 days. Faecul egg counts. Ascaris sum and Oesophago-
Parasitological techniques. The number of eggs per gram stomum spp. egg counts are given in Figs 1 and 2, and
 Impact of fasting on A. XMTI and Oesophagostomum spp. in pigs

I, ! # I, 8 11 1 t
0 1 2 3 4 5 6 r 6 9 10
‘1 2 3 4 5 6 7
Days of fasting
Fig. 1. Geometric mean Ascaris suum egg counts in pigs Section of intestine
fasted for 6 days (U), fed for 6 days (m), fasted for 10 days Fig. 3. Distribution of Ascaris swrn in pigs fasted or fed
(m) and fed for lOdays (0). for 6 days. Sections of intestine: 1425%; 2-2&50%; 3-
51-75%; &7&100% of small intestine; SLarge intestine;
&Expelled; 7-Total.

m Faatad
: 20
u
k, 16
n
E 12

g5 8

II 11 L 11 11 1 I g 4
0 I 2 3 4 5 6 7 6 9 10

0 n w n
Days of faxding
1 2 3 4 5 6 7
Fig. 2. Geometric mean Oesophagostomum spp. egg counts
in pigs fasted for 6days (O), fed for 6days (O), fasted for Section of intestine
lOdays (m) and fed for lOdays (0). Fig. 4. Distribution of Ascaris wurn in pigs fasted or fed
for lOdays. Sections of intestine: l&25%; 2-2650%;
3-51-75%; &7&100% of small intestine; 5-Large intes-
tine; &Expelled; 7-Total.
did not differ significantly by species at the outset.
Numbers of A. suum eggs (epg) in the fasted group
increased gradually and differences between the fasted in control pigs, and was reduced 60.7% by fasting
and fed groups were statistically significant on the (P=O.Ol).
5th and 6th day of starvation (P c 0.05). Numbers of Oesophagostomum spp. burdens and distribution
Oesophagostomum spp. eggs (epg) in the fasted pigs are shown in Fig. 5. The total number of Oeso-
were not significantly different from those in fed pigs phagostomum spp. was not significantly affected by
from days 2 to 6 of fasting (P>O.O5). fasting (P> 0.05).
0. dentatum constituted on the average 92% and
95% of the total Oesophagostomum worm burden in
Worm burdens experimental and control pigs, respectively. The
Ascaris suum burdens and distribution are given in remaining worms were identified as 0. quadris-
Fig. 3. There were no statistically significant differ- pinulatum. The mean (S.D.) location of 0. dentatum
ences in total worm burden between the groups was sections 2.5 (1.0) and 2.2 (1.2), of 0. quadris-
(P> 0.05), but worm location was significantly affec- pinulatum sections 1.9 (0.4) and 1.5 (1.0) of the large
ted by fasting (P=O.O4). In the fasted pigs, 27% of intestine in experimental and control pigs, respec-
the total worm burden was found in the proximal tively. 0. quadrispinulatum showed a more proximal
parts of the large intestine. The back-transformed location in the large intestine, but did not differ sig-
mean (S.D.) of A. suum fecundity was 155,794 (77,791) nificantly from the location of 0. dentatum. In the
eggs per female worm per day in experimental pigs experimental group, 35% of 0. dentatum and 22% of
and 396,259 (184,420) eggs per female worm per day 0. quadrispinulatum were found in the posterior part
434 S Petkevieius et al.

2100 m OAenfafum
I
- O.quadrispinuiatum
ci 1800
z 1 Fasted m Odentatum
1500
.z 0.quadrispinulatum
E 1mo
9 900

123456
Section of large intestine
Fig. 5. Distribution of Oesophagostomum and 0. quadrispinutatwn in pigs fasted or
denta?um
fed for 6days. Sections of large intestine: 1-Caecum; 2420%; 3-2140%; -l-60%; 5-
61-100%: &Total.

(section 5) of the large intestine, in which no worms worm per day in the experimental pigs and 461,369
were found in the controls. All Oesophagostomum (194,184) eggs per female worm per day in the control
examined were adults and for both species in both pigs, showing a 85.2% reduction due to fasting
groups there was an equal distribution of males and (P=O.O05).
females. The back-transformed mean (SD.) of Oeso- Oesophagostomum dentatum and 0. quadrispinu-
phagostomum spp. fecundity was 86.5 (1348) eggs per datum burdens and distributions are shown in Fig.
female worm per day in fasted pigs and 2927 (2577) 6. The total number of worms was not significantly
eggs per female worm per day in fed pigs, showing a affected by lOdays of fasting (P>O.O5). 0. dentutum
reduction of 70.4% (p=O.O3). constituted on the average 94% and 92% of the total
Oesophugostomum worm burdens in experimental and
control pigs, respectively. The remaining worms were
Experiment II (10 duyfast)
identified as 0. quudrispinuhztum. The mean (S.D.)
Faecal egg counts. Mean egg counts of A. suum and location of 0. dentatum was sections 2.8 (0.6) and 2.5
Oesophagostomum spp. are depicted in Figs 1 and 2, (0.6), of 0. quadrispinukztum sections 1.6 (0.9) and 1.2
and did not differ significantly by species at the outset. (1 .O) of the large intestine in experimental and control
There was a gradual rise in Oesophagostomum spp. pigs, respectively. 0. quadrispinulatum showed a more
and A. suum counts in experimental pigs up to day 6 proximal location in the large intestine and differed
and 8, respectively. For A. suum, the difference significantly from the location of 0. dentatum
between experimental and control pigs was statis- (P = 0.04). All Oesophagostomum worms recovered
tically significant (P=O.O3) on days 8 and 10; it was were adults and for both species in both groups there
significant for Oesophagostomum spp. on day 6 was an equal distribution of females and males. The
(p=O.O4). Towards the end of the period, however, back-transformed mean (S.D.) of Oesophagostomum
there were marked reductions in egg counts for both spp. fecundity was 184 (256) eggs per female worm
species which were more pronounced for Oeso- per day in experimental pigs and 3390 (2608) eggs per
phagostomum spp. where counts approached zero. female worm per day in control pigs, showing a 94.6%
Worm burdens. The total worm burdens and dis- reduction (P = 0.0005).
tribution of A. suum are shown in Fig. 4. As expected
large individual variations in A. suurn burdens were
observed between pigs within experimental and con- Clinical and other observations
trol groups, but the mean total A. suum burden of the The behaviour of fasting pigs in both experiments
experimental pigs was significantly lower as compared differed from fed pigs but not significantly. During
with the control pigs (P=O.OOl). The mean (S.D.) the first days the fasted pigs were restless and nervous,
location of A. suum in the experimental and control but later they became less active and were lying/
pigs was sections 2.4 (1.3) and 1.4 (0.4) of the small sleeping most of the time. Some cases of coprophagy
intestine, respectively. and an average of 0.9 (2.8) were observed in the experimental animals. The mean
worms were found in the large intestine and 1.2 (2.5) live weight of fasted pigs had decreased 1.8 kg per day
were expelled. The back-transformed mean (S.D.) of after 6 days of fasting and 1.O kg per day after 10 days,
A. suum fecundity was 68,502 (72,791) eggs per female while mean live weight of fed control pigs had
 Impact of fasting on A. ~w;;z and Oesophagostomwn spp. in pigs 435

- O.dentatum
1600 m O.quadrispitwlatum
5
F = O.dentatum
2 1200 m O.quacirkpinulatum
E
; 600
2
g 400

0
123456 123456
Section of large inteetine
Fig. 6, Distribution of Oesophagostomunz dentatwn and 0. quudrispinulatum in pigs fasted
or fed for lOdays. Sections of large intestine: I-Caecum; 2420%; 3-2140%; Wl-
60%; 5-61&100%; &Total.

increased 1.2 kg per day by day 6 (Experiment I) and time numbers of A. suum, but not Oesophagostomum
0.7 kg per day by day 10 (Experiment II), respectively. spp., were significantly reduced in the fasted pigs as
As expected the experimental pigs after fasting for well.
6 and lOdays were significantly lighter that control Fasting of pigs for both 6 and 10 days led to a more
animals (P=O.OOl). The faeces of the fasted pigs wide distribution of both species along the gastro-
gradually developed a hard consistency and after 6 intestinal tract. The most striking feature was the
days of fasting faeces appeared in “pelleted” form. recovery of worms from gut sections where they are
The mean (S.D.) weight of excreted faeces in the fasted normally not found, i.e. for A. suum in the large intes-
pigs decreased from 892 (819) g at the start of the tine; and for Oesophagostomum spp. in the very distal
experiment to 63 (29) g at the 6th day of fasting and part of the large intestine. These were the regions
from 669 (169) g to 39 (18) g at the 10th day of fasting, where small amounts of gut contents were still found.
respectively. At slaughter intestinal contents in the In both experiments 0. dentatum constituted 92-95%
guts of fasted pigs were sparse and seen only in the of the Oesophagostomum species.
large intestines, primarily in the most distal parts. 0. quadrispinzdatum had a more proxima1 location
in the large intestine than 0. dentatum, which is in
agreement with findings of Roepstorff & Nansen (in
DISCUSSION
press) and Christensen et al. (in press).
The results of these experiments show a significant Few studies have been designed to analyse the
effect of fasting on parasitism. Fasting of pigs for 6 influence of fasting on helminth parasites. In the faeces
and 10 days influenced worm numbers, fecundity and of domestic horses starved for ten days, the number
location in the gut. A. suurn and Oesophagostomum of strongylid eggs decreased, and expulsion of adults
spp. epg in the experimental pigs started to increase and larvae of the genera Delafondia and Alfondia was
from the first day of fasting, and high levels were observed (Dvojnos & Timoshenko, unpublished,
observed until day 6 for Oesophagostomum spp. and Abstracts of Eighth International Congress of Para-
day 8 for A. suu?n, after which marked decreases sitoIogy. Izmir, Turkey, 1994). Under natural winter
occurred (Experiment II). The initial increase may conditions in Mongolia wild horses had great diffi-
obviously be explained by the significant reduction in culty finding food and showed a significant expulsion
the volume of faeces excreted, while the drop in epg of adult strongyhds (Dvojnos & Timoshenko. unpub-
towards the end of the study might be a consequence lished. Abstracts of 2nd European Congress of Mam-
of worm expulsion or lowered fecundity of remaining malogy, Southampton, U.K., 1995).
females, or both. Both possibilities were confirmed by The results of our experiments show that young,
determinations of worm numbers and worm fecundity healthy pigs can sustain withdrawal of feed for 6 and
at slaughter. On day 6, numbers of A. suuw and Oeso- 10 days without effects on health and behaviour, pro-
phagostonzum spp. were not affected by fasting com- viding that they are adequately housed and allowed
pared with fed controls, but fecundity was access to water ad libitum. There was a higher level of
significantly reduced, especially for Oesophago- general behavioural activity of the fasting pigs than in
stomu~. On day 10 the fecundity of both A. suum and control animaIs during the first few days of fasting
Oesophagostomum spp. was even lower, and at this in both experiments, before lethargy set in. Similar
436 S. Petkevieius et al.

observations on the behaviour and health status of malnutrition on gastrointestinal helminths is justified
pigs fasted for IO days were made by Anderson (1988). for what we can learn about the complex ways in
The mechanisms by which fasting decreases the sur- which parasites and hosts co-exist, even in unusually
vival and fecundity of A. suum and Oesophagostomum adverse situations.
spp. in the gastrointestinal tract of pigs are unknown.
One likely explanation is that the reduction in intes-
tinal contents in the small and large intestine during
Acknowledgements-‘IIe study was supported by the Danish
fasting deprives the worms of their normal nutrient National Research Foundation.
supply or otherwise creates unfavourable physio-
logical or biochemical conditions. Wills (1985) states
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