Preventive Veterinary Medicine, 2 (1984} 411--421 411

Elsevier Science Publishers B.V., Amsterdam -- Printed in The Netherlands

BOVINE LEPTOSPIROSIS IN THE~ TROPICS: P_REVALENCE, PATHOGENESIS AND CONTROL

W. A. ELLIS
Veterinary Research Laboratory., Stoney Road, Stormont, Belfast, BT4 3SD
(Northern Ireland)

ABSTRACT
W. A. Ellis, 1984. Bovine leptospirosis in the tropics: prevalence, pathogene-
sis and control. Prev. Vet. Med., 2: 411-421.
Evidence suggests that bovine leptospirosis may be a cause of enormous
economic loss from abortion, stillbirth, death, decreased milk production
and infertility in tropical and sub-tropical countries. This paper reviews
the epidemiological features, prevalence, pathogenesis, clinical features
and control measures for leptospiral infections in cattle in these regions and
the shortcomings of the methods used to assess these parameters. The possible
financial losses which may accrue from only one aspect of bovine leDtospirosis-
reproductive wastage-are illustrated.

INTRODUCTION
Leptospirosis is a common zoonotic disease which can infect most manuals and
occurs throughout the world. It causes economic loss to the cattle industry
from abortion, stillbirth, death, decreased milk production and infertility.
It can be caused by any of the parasitic organisms classified within the genus
Leptospira. The last official list (WHO 1967) recognised 16 serogroups and
more than 150 serovars have now been identified.
~4any aspects of leptospirosis in cattle are still inadequately defined.
This is particularly so in the cattle population of tropical and sub-tropical
countries where the literature indicates widespread serological evidence of
infection but where there are relatively few microbiological or clinical
disease reports. Most of what we know of leptospirosis in cattle in the latter
countries has been as a result of work carried out in North Eastern Australia
and in Central and South America, and by extrapolation of data from temperate
ecosystems to the tropical situation. Consequently this paper reflects the
bias in published information.

EPIDEMIOLOGICAL CONSIDERATIONS
Theoretically any parasitic leptospire may infect any animal species, but
in practice only a small number of serovars will be endemic in any particular
region or country. Furthermore, each serovar tends to be maintained in

0167-5877/84/$03.00 © 1984 Elsevier Science Publishers B.V.

412

s p e c i f i c m a i n t e n a n c e h o s t s (BlacI~norc- and Scbolltm~, 19S0). T b e F e l o r e in an\"

region cattle w i l l be i n f e c t e d b y : -
(a) strains a d a p t e d t o and m a i n t a i n e d hx, c a t t l e (eg s e r o w ~ l~Fd.jo)~tnd
where d i r e c t cow t o cow t r a n s m i s s i o n is prol~abl,x: ol- g r e a t e s t importance.
I n f e c t i o n by s t r a i n s o f h a r d j o a p p e a r s t o he i n d e p e n d e n t o f ~'egion and
the results o f V,l d e r and l~'ard i n 1978 i n d i c a t e t h a t it is a l s o
independant of rainfall;
(b) strains m a i n t a i n e d hy o t h e r d o m e s t i c and f r e e - l i v i n g anianals, t',q~at
these incidental i n f e c t i o n s a r e w i l l be d e t e r m i n e d bv what o t h e r animal
s p e c i e s a r e p r e s e n t i n t h e a r e a and what s e r o v a r s t h e y m a i n t a i n . Most
tropical c o u n t r i e s have a wide r a n g e o f f r e e - l i v i n g a n i m a l s and g e n e r a l
farm management p r a c t i c e s and c o n d i t i o n s p r o v i d e :~n~le o p p o r t t m i t y f o r
direct or indirect contact with cattle (and s u b s e q u e n t t r a n s m i s s i o n ) .
The p r e v a l e n c e o f i n c i d e n t a l i n f e c t i o n s w i l l a l s o depend on w h e t h e r t h e
environment favours stnwival of leptospires outside the host. Optimt~n
s u r v i v a l o c c u r s i n wa~m wet c o n d i t i o n s w i t h a ~1t c l o s e t o n e u t r a l . Since
such c o n d i t i o n s o c c u r more conm~onlv in t r o p i c a l regions than elsewhere
incidental infections are relatively more i m p o r t a n t i n t r o p i c a l countries
than in other regions. The warm wet c o n d i t i o n s f a v o u r a h l e t o t h e t r a n s -
mission of incidental i n f e c t i o n s w i l l even be found i n dry d e s e r t r e g i o n s
i n c e r t a i n m i c r o - e n v i r o n m e n t s such as w a t e r - b o l e s and t h e i r immediate
s u r r o u n d i n g s where animal c o n c e n t r a t i o n s may he h i g h . l',1~ere r a i n f a l l is
seasonal, transmission of incidental i n f e c t i o n s may a i s o be s e a s o n a l
( D o h e r t y , 1967).

LIMITATIONS IN THE AVAIIAkBLE DATA

The frequency of recognition of leptospiral infections to a large extent
reflects the interest of investigators (Sullivan, 1974) and the investigation
facilities and ftmding available within a country. Any con~nent on the
prevalence and clinical significance of particular serovars in cattle must
also be balanced by an appreciation of the limitations in the teclmiques used
by investigators to assess these parameters. These assessments are usually
made by a combination of clinical and laboratory investigations and the
limitations in either effects the other.
Clinical investigations in tropical countries have been handicapped by the
broad spectrum of and non-specific nature of the clinical signs found in acute
and chronic bovine leptospirosis. Acute clinical signs may be confused with
those of babesiosis (Tabel and Losos, 1979) and anaplasmosis both of which are
prevalent in many tropical countries.
Laboratory investigations have utilised microbiological, serological and
 413

experimental methods. All these have inherent limitations.

Microbiological methods
These have included attempts to identify those leptospires which (I)
cause clinical disease in cattle, (2) are maintained by cattle and (3) can
incidentally infect cattle. This has mainly been done by the attempted
isolation of leptospires from ~]) tissues of clinical cases (2) the kidneys
of unaffected cattle (abattoir surveys) and (3) the kidneys of those species
likely to come into contact with cattle, respectively.
Such isolation attempts have been limited by: the difficulties
experienced in isolating many leptospires especially from clinical material;
the age of the animal examined; a lack of suitable isolation media; a lack
of rmiformity in media used in different studies and the lack of suitably
experienced laboratory staff in most veterinary laboratories particularly in
tropical countries with restricted budgets. These shortcomings have meant
that there has been an absence of meaningful microbiological studies in tropical
com~tries and they make direct comparison of reports from different
laboratories in more temperate regions rmreliable. The strains isolated must
be considered as those which the methods used could isolate and not
necessarily the most prevalent leptospire present.
Serological methods
These have consisted of surveys in which a number of sera from healthy and/
or suspect leptospirosis cases have been examined for leptospiral antibodies by
a variety of serological tests. This type of approach has been the one most
widely used and has been the basis for most of the reports of bovine
leptospirosis from tropical and sub-tropical countries. The value of such
surveys has been restricted by a number of factors:-
(i) The lack of meaningful microbiological data in tropical co~mtries has
meant that the antigens used in serological surveys may not adequately
represent the serovars present in the particular region.
(2) Serological prevalences may not reflect economic importance.
(3) The methods of population sampling have frequently been limited, biased
and unsuitable for statistical analysis.
(4) Arbitrary significant titres have been used in the interpretation of
serological survey data. While useful in eliminating some of the
confusion which may arise due to cross reactivity of certain L_eptospira
serogroups the use of such arbitrary values has no merit in the
investigation of the cattle adapted serovar (hardjo) ~llis et al., 1981;
1982b).
Experimental studies
The pathogenicity of a particular serovar may vary from strain to strain
414

found within a ~mtry and between strains fom~d in different parts of tl~e
world. There has been a reluctance to carry out e~]erimental studies ~si~g
strains isolated front cattle in tropical and sub-tropical countries.
Recently, ~Iadruga (]980) and Aycardi et ai.(1982) have s h o ~ that South
American hardj9 strains have a sim]lar experimental pathogenic ity to str;ii~s
isolated in temperate regions (Ellis and }~icbna, 1977; Tl~ie~nam~, ll)S2).

PREVALENCE
There is widespread serological evidence of infection of cattle throughout
the tropical and sub-tropical regions, l#here suitable antigens have been used
these indicate that infections by strains belonging to the Hebdomadis serogrou!~
almost always predominate. In Africa, Ball (1966), Van Riel et a]. (]971),
EI-Vali (1979) and Ezeh and Agba (1982) found such a ~redominance of Ilebdomadis
serogroup reactions in Kenya, Senegal, Sudan and Nigeria respectively. Simil;~r
findings were reported in South and Central ~%~erJca by Acha et al. (1963),
Clarke et al. (1966), Jelanbi et al. (1976), Lara ~1978), ~.loreira et ~il. ~]979),
_ _ _

Edwards et al. (19~I) and Rivera et al. (]981) (in Guatemala, Nicaragua,
Venezuela, Mexico, Brazil, Bolivia and Columbia respectively). IIigh serological
preva]ences have also been reported in the Oueensland region of Austr~lia (Elder
and Ward, 1978), ~{alaya (Wisseman et al., ]g55)and in the l'bilippines (C~rlos
et al., 1970).
I~hile there has been a lack of suitable microbiological surveys to supplement
these serological surveys, extrapolation of data from other areas of the world
(Hellstrom, 1978; Ellis et al., 1981; I~ite et al., ]982) would indicate that
the majority of these Hebdomadis serogroup reactions are probably due to the
cattle adapted strain serovar hardjo. In South and Central /~nerica, where a
large number of Hebdomadis group serovars have been isolated from ~ variety of
species (Szyfres, 1976), incidental exposure of cattle to such strains may
contribute significantly to this preponderance of serological reactions to
Hebdomadis serogroup m~tigens.
The prevalence of serum antibodies to serovars belonging to other serogroups
varies from region to region and country to country and reflects the different
distribution of incidental infections in these regions. In North ilastern
Australia (Elder and Ward, 1978), Brazil (}4oreira et al., ~979), ~{exico
(Lara, 1978), Nicaragua (Clarke et al., 1966) and Senegal (Van Riel et al., 1971%
Pomona g~-oup antibodies were the second most commonly detected antibodies, while
in Bolivia (Edwards et al., 1981) and the Suda[~ (E]-Vali, 1979) this role was
filled by Tarassovi serogroup antibodies, and in Nigeria (Ezeh and Agba, 1982)
by reactions to the B a l l ~ serogroup.
Occasionally in some countries envirorm~ental contamination by free-l~ving
 415

animals with strains belonging to one serogroup may be at such a level that
this is reflected in serological surveys of cattle by antibodies to such
incidental infections being the ones most commonly detected. Examples of this
are to be found in Barbados (Damude et al., 1979) where Autumnalis serogroup
infections predominate and in Vietnam (Spinu et al., 1978) where Tarassovi
infections are most prevalent.

PA~HOGENESIS
Infection of susceptible cattle can occur though mucous membranes and
abraded skin and is followed after a 4 to IO day incubation period by a
bacteraemic phase M~ich may last from a few hours up to 7 days. This phase
is characterised by pyrexia, excretion of leptospires in milk and with some
serovars by functional damage to the internal organs. Acute clinical
infection occurs at this stage.
With the appearance of circulating antibody bacteraemia ceases but
leptospires localise and persist in a number of organs, especially in the
proximal renal tubules and in the female genital tract. Relapses of pyrexia
may occur. I~hether these are associated with persistent loci of infection
giving rise to recurring bacteraemia is unknown.
Leptospires are excreted in urine for a variable period depending on the
infecting serovar and the age of the animal. In incidental infections urinary
excretion usually only lasts for several weeks but with the cattle maintained
strain (serovar hard, o) excretion can last as long as 542 days (Thiermann, 1982)
and possibly for the animal's lifetime. Urinary excretion is very important in
the transmission of leptospirosis.
Localisation of leptospires in the pregnant and non-pregnant uterus has
been shown to persist for up to 142 days and 97 days post-infection
respectively (Thiermann, 1982). Localisation in the pregnant uterus may in
turn be followed by fetal infection With subsequent chronic reproductive
wastage and excretion of the leptospires in the post-calving uterine discharge
for up to 8 days and with persistance in fallo~ian tubule for up to 22 days
post-calving, (Ellis, unpublished data).
Whether localisation occurs in the male genital tract is not certain but
Sleight et al. (1964) demonstrated ~omona in the semen of a bull 18 and 38
- - _ _

days post-infection. Vaz and Oliveira (1978) detected leptospiral antibodies

in the semen of 849 of bulls examined by them indicating possible local
antibody production in the genital tract.
Strains belonging to the Hebdomadis serogroup may also localise and persist
in the mammary gland as Thiermarm (1982) recovered hardjo from milk of an
experimentally infected cow on days 30 and 91 post-infection.
416

CLINICAI~ FFATURES
Clinical bovine leptospirosis can be d i v i d e d i n t o two d i s t i n c t phases:
firstly, an a c u t e p h a s e whose o n s e t c o i n c i d e s ~'ith t h e b a c t e r a e m i c !~base o~
infection, and s e c o n d l y , a c h r o n i c p h a s e xd~ich o c c u r s much l ~ t e r and ~,hose
affects a r e most n o t i c e a b l e on t h e r e p r o d u c t i v e ~ r a c t .
Acute infection
The c l i n i c a l signs exhibited i n t h e a c u t e pha~o can v a r y m a r k o d l v in
severity. The most s e v e r e f o r e is n s u a l l y o b s e r v e d in young c a l v e s
(English, 1979) a l t h o u g h o u t b r e a k s in a d u l t s have a l s o been rel~ortcd
(Burdin et al.,1958; T a b e l and Losos, 1979). Tho c l i n ~ c a I ~gns a r e ghoso off
severe septicaemia with pyrexia, anorexia, an actltC h;tomolyt ic a n a e m i a ,
haemoglobinuria ~d jaundice. The m o r t a l t r y r a t e i s h i g h and i f r e c o v e r y does
occur there is a prolonged convalescence period. In l a c t a t h ~ g cows m i l k
p r o d u c t i o n a l m o s t s t o p s and what m i l k t h e r e is i s p i n k or r e d c o l o u r e d and
contains blood clots. A necrotic dc~m~atitis lnay occtlr i n some a n i m a l s
infected with Grippot~hosa strains (lklrd~n e t a l . , 19~8) and o c c a s i o n a l
c a s e s may be c h a r a c t e r i s e d by s i ~ s of meningitis (English, 1979).
Such p e r a c u t e o u t b r e a k s a r e a s s o c i a t e d ~vith i n c i d e n t a l infection of
cattle by h a e m o l y s i n p r o d u c i n g s t r a i n s o f ~he Pomona, ( ~ r i p p o t y p h o s a ,
Icterohaemorrhagiae and A u t ~ a l i s serogroups.
T h e r e may be a d e g r e e o f b r e e d s u s c e D t i b i l i t y t o such a t t i r e i n C e c t i o n s .
Burdin et al. (1958) o b s e ~ e d a higher mortality i n European and European X
Zeb u c a t t l e than in native Boran c a t t l e d u r i n g a (;ril~oty~i~osa o u t h r c a k in
Kenya.
There is a gradation o~ c l i n i c a l s i g n s from t h o s e o b s e ~ e d in very
acute cases to the mildest fo~ of acute infection M~ich i s s e e n i n a d u l t d a i r y
cattle. This is characterised by p y r e x i a , which i s f r e q u e n t l y ~detected
and by a marked d r o p i n m i l k flow. This mild clinical s~drome is usually
associated with serovar hardjo infection, a l t h o u g h i t c a n be c a u s e d by s t r a i n s
belonging to other serogroups. All f o u r q u a r t e r s o f t h e udde~ a r e a f f e c t e d
and t h e u d d e r f e e l s s o f t and f l a b b y and n o t h a r d as i n m a s t i t i s cases
(Ellis et al, 1976; H i g g i n s e t a l , 1980). Milk from a f f e c t e d coors a p p e a r s
thick, yellow ~d colost~-like. It contains clots and h a s a h i g h ~ h i t e
cell co~t. Agalactia lasts f o r 2-10 days a f t e r which p r o d u c t i o n u s u a l l y
returns to almost no~al although i£ infection occurs late in lactation milk
p r o d u c t i o n may n o t r e t u r n .
The n ~ b e r o£ a n i m a l s a f f e c t e d on a £ a ~ w i l l v a r y £rom m o r e t h a n 50$ o f
cows o v e r a two month p e r i o d o£ a n e p i d e m i c i n a s u s c e p t i b l e herd to sporadic
cases ~ animals in first or second lactation in endemically infected herds
(Hathaway aM L i t t l e , 1983).
 417

While agalactia is not readily recognised in beef cattle it may become

apparent by the presence of weak calves that do not receive nutrition
(Ilanson, 1980).
In many animals the acute phase of infection is apparently sub-clinical.
Hoare and Claxton (1972) observed that the drop in total herd milk production
could not be explained by the number of clinically affected cows. A
comparison of milk production predictions with actual production figures in
infected herds (lliggins et al., ]980) and a comparison of pre and post-
vaccination milk production fignres (Carroll, 1966) would suggest that there
may be considerable losses in milk production associated with such apparently
s~-clinical infections.
Chronic infection
In an animal recovering from the most acute form of the disease there is
a prolonged convalescence with associated liver and kidney damage contributing
to reduced growth rate. Occasionally relapses may occur, especially in
Grippotyphosa infections (Bernkopf et al., 1947; Burdin et al., 1958).
The most important chronic sequels to leptospirosis are seen in the
pregnant cow, where consequent fetal infection with resultant abortion,
stillbirth and premature live birth of weak calves may occur. Such chronic
events occur one to twelve weeks and occasionally longer after the acute
phase of infection, however, such animals have frequently shown no clinical
evidence of acute infection.
With incidental infections these events usually occur in the last 3 months
of gestation although Damude et al. (1979) observed abortion in the second
trimester in cases of Autumnalis infection. With serovar hardjo infection,
abortion has been diagnosed at all stages from the fourth month through to term
and circumstantial evidence indicates that it may also cause embryonic death
(Ellis, 1983).
It has been very difficult to define the importance of leptospirosis
relative to other factors in the aetiology of bovine abortion, stillbirth and
premature live birth in national herds because of the difficulties in diagnosis.
Recent studies indicate that this role may be considerable. Mazzonelli and
Jelambi (1980) attributed a leptospiral aetiology to 40.8% of 1,526 abortions
investigated by them in Venezuela. More recently in Northern Ireland, Ellis
et al. (198~) were able to demonstrate leptosDiral infection (almost entirely
serovar hardjo) in 49.7% of 348 aborted fetuses which they examined by a
combination of isolation of leptospires, i~unofluorescence and fetal serology.
These latter figures may be directly relevant to many tropical and semi-tropical
countries since hardjoinfection is endemic in many of those countries and
experimental studies with South American har~o strains have shown that they are
418

pathogenic for the bovine fetus(~.ladruga, 1980; @ c a r d i et a l, 1982).

The incidence of abortion on individual farms can be very bigb (up to 50
per cent) following an epidemic of an i n c i d e n t a l ]eptosT~iral i n f e c t i o n such as
pomona (Knott and Dadswel], 1970). l~ith s e r o v ar hardjo i n f e c t i o n , a b o r t i o n
r a t e s tend to be much lower, however, hardjo i n f e c t i o n is probably the most
important l e p t o s p i r a l i n f e c t i o n as i t is a s s o c i a t e d with long-te~z~ herd
i n f e c t i o n s on farms with new i n f e c t i o n s o c c u r r i n g year a f t e r year (Hanson, 1972)
Hathaway and L i t t l e (1983) have shown how important t h i s r e c u r r i n g loss can
be, where management p r a c t i c e s r e g u l a r l y ensure the intro&~ction of s u s c e p t i b l e
animals i n t o an i n f e c t e d herd: out of a group of 13 s e c o n d - c a l f h e i f e r s 3
aborted (hardjo was i s o l a t e d from the two a v a i l ~ b l e f e t u s e s ) and another d e l i v e r .
ed a premature l i v e c a l f . This episode occurred approximately 14 months a f t e r
an outbreak of acute hardjo a g a I a c t i a in the main herd.
Infertility in hardjo_ i n f e c t e d herds has been a common f i e l d o h s e r v a t i o n
by v e t e r i n a r i a n s (Hanson and Brodie, 1967) and improvements in breeding
e f f i c i e n c y have been noted in herds f o l l o w i n g v a c c i n a t i o n (ttanson e t a l . , 1!)72).
S i m i l a r o b s e r v a t i o n s have been made with i n f e c t i o n by s t r a i n s belonging to
o t h e r serogroups ( C a r r o l l , 1966).

ECONOMIC LOSSES
Economic losses due to leptospirosis can be caused by any of the acute,
chronic or sub-clinical features outlined above. It is impossible to assess
what the total cumulative losses are either in an individual herd or in
national herds, as there are too many variables. However, if only one aspect
is considered - pre-natal death - it is possible to envisage the enormity of
the problem. Holroyd (1980) reported that in a six year, split herd,
vaccination eN)eriment with a serovar hardjo bacterin conducted in tropical
Northern Queensland that vaccination caused a significant (P< O.l) reduction
of pre-natal loss (from 3.6 to ].4%/year). Should this 2.2% reduction
represent the losses due to hardjo in cattle in tropical countries (and it
could be an underestimate) then in South and Central Nnerica with an
estimated total cattle population of 250 million the annual calf loss due to
leptospirosis would be enormous.

CONTROL MEASURES
Control measures have been reviewed by Hanson (1977, 1980, 1982).
V a cci n at i o n with and without supplementary a n t i b i o t i c therapy ( d i h y d r o s t r e p t o -
mycin a t 25 mg/kg) o f f e r s the only e f f e c t i v e method of c o n t r o l l i n g and
p r e v e n t i n g c l i n i c a l l e p t o s p i r o s i s in c a t t l e herds. In closed herds v a c c i n a t i o n
of a l l members o f the herd should be c a r r i e d out annually while in open herds
a l l members should be r e v a c c i n a t e d every s i x months. During ~m episode of
 419

clinical leptospirosis, Icptosniral vaccines can be administered

simultaneously with dihydrostreptomycin (South and Stoenner, 1975).
The vaccine used should be a multi-valent bacteria containing the
serovars endemic in the region. This requirement can be a major drawback to
the use of vaccines in tropical countries since the endemic serovars have not
been adequately identified in many such countries.

FUTURE I)E~q~LOP~~NTS
Future research progr:nns in the tropics must take cognisance of the
limitations which have applied to previous studies. At a national level there
is a need for : - (I) detailed microbiological investigation of carrier and
suspect clinical animals to identi~y those serovars M~ich affect the food
producing animals: and (2) long-term clinical and laboratory' studies to assess
the actual extent of economic loss in infected herds and to evaluate the
e f f e c t s of v a c c i n a t i o n ~'ith s u i t a b l e b a c t e r i n s . This should be supported
i n t e r n a t i o n a l l y by the development of more e f f e c t i v e i s o l a t i o n media and by
the p r o v i s i o n of adequate r e f e r e n c e tyq~ing f a c i l i t i e s ~or i d e n t i f y i n g serovars
i s o l a t e d from animals.

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