Livestock Science 132 (2010) 1–12

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Livestock Science
j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / l i v s c i

Review article

Genetics of adaptation in domestic farm animals: A review

T. Mirkena a,b,c, G. Duguma a,b,d, A. Haile a, M. Tibbo e, A.M. Okeyo f, M. Wurzinger b, J. Sölkner b,⁎
a
International Livestock Research Institute (ILRI), PO Box 5689, Addis Ababa, Ethiopia
b
Division of Livestock Sciences, Department of Sustainable Agricultural Systems, University of Natural Resources and Applied Life Sciences (BOKU),
Vienna, A-1180 Vienna, Austria
c
Department of Animal and Range Sciences, College of Agriculture, Hawassa University, PO Box 05, Hawassa, Ethiopia
d
Bako Agricultural Research Center, PO Box 03, Bako, Ethiopia
e
International Center for Agricultural Research in the Dry Areas (ICARDA), PO Box 5466, Aleppo, Syria
f
International Livestock Research Institute (ILRI), PO Box 30709, 00100 Nairobi, Kenya

a r t i c l e i n f o a b s t r a c t

Article history: This review summarizes available information on genetics of adaptation in major livestock
Received 26 January 2010 species focusing on small ruminants. Adaptation to humans and consequences of
Received in revised form 30 April 2010 domestication on predator aversion, mechanisms of adaptation to available feed and water
Accepted 7 May 2010
resources, severe climates and genetic evidence of disease tolerance or resistance have been
presented. The latter focuses on gastrointestinal parasites and bacterial diseases. The resource
Keywords:
allocation by the animal to production and fitness traits under both optimal and sub-optimal
Genetics
conditions has a genetic background. Such information would help in identifying the most
Domestication
Adaptation appropriate and adapted genotypes capable of coping with the environmental challenges
Disease resistance posed by the production systems or, wherever possible, in adapting the environments to the
Review requirements of the animals.
© 2010 Elsevier B.V. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Domestication . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
3. Adaptation to the environment and production system . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3.1. Adaptation to humans . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3.2. Behavior towards predators . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3.3. Adaptation to available feed resources . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3.3.1. Low metabolic requirements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3.3.2. Ability to reduce metabolism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3.3.3. Digestive efficiency and ability to utilize high-fiber feed . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3.3.4. Fat deposition as feed reserve . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3.4. Adaptation to severe (hot/cold) climates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
3.5. Adaptation to water scarcity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
3.6. Tolerance/resistance to disease. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
3.6.1. Parasitic diseases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
3.6.2. Bacterial diseases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
3.7. Resource allocation theory (production vs. fitness traits) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

⁎ Corresponding author. Tel.: +43 1 47654 3271; fax: +43 1 47654 3254.
E-mail address: soelkner@boku.ac.at (J. Sölkner).

1871-1413/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.livsci.2010.05.003
2 T. Mirkena et al. / Livestock Science 132 (2010) 1–12

4. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

1. Introduction mances of zebu cattle adapted to the harsh tropical climatic

conditions. Similarly, successful dairy and beef programs have
Adaptability of an animal can be defined as the ability to been developed in Brazil (Madalena, 2000).
survive and reproduce within a defined environment In the dairy industry, decline in average fertility and health
(Prayaga and Henshall, 2005) or the degree to which an of cows have been associated with increased genetic merit for
organism, population or species can remain/become adapted milk yield (e.g. Pryce et al., 2004). Goddard (2009) gives two
to a wide range of environments by physiological or genetic main reasons for the decline in fitness traits: (1) these traits
means (Barker, 2009). Smallholders, pastoralists and their were ignored in the construction of selection indices because
animals often live in harsh environments which may be hot they were considered to have lower heritability or not easy to
and dry, hot and humid, or high in altitude and cold. record and (2) use of inappropriate breeding programs while
Moreover, these environments can be characterized by the underlying genetic process (selection and inbreeding
scarce feed and water resources and high disease pressure depression) is well understood. However, the low heritability
with large seasonal and annual variation (Sölkner et al., of some fitness traits does not imply negligible genetic
1998). Adaptation to these factors is largely based on variance; often heritability is low because the phenotypic
genetics, but animals can “learn” to live under such stressful variance is rather larger than the genetic variance is small as
conditions. In order to match genotype with the environ- evidenced by as high genetic coefficient of variation for fitness
ment, breeders can follow two alternative strategies: adapt traits as for some production traits (Goddard, 2009; Hill and
the environment to the need of the animals as is the case in Zhang, 2009). Inclusion of functional traits (functional longev-
industrial animal production systems or keep animals that ity, persistency, fertility, calving ease, still birth and somatic cell
are adapted to the respective environment as is the case in count) in a total merit index has been reported to have a
low input smallholder and pastoral systems. The physical positive effect on the annual monetary genetic gain (Sölkner et
environment greatly differs between locations and produc- al., 1999; Willam et al., 2002; Veerkamp et al., 2002; Philipsson
tion systems based on available resources and economic and Lindhe, 2003; Samore et al., 2003; Weigel, 2006).
conditions. Because of this, smallholders and pastoralists Tick counts, fecal worm egg counts (FEC), rectal tempera-
need different and diverse animal genotypes, species mix tures and coat scores have been used as indicator traits of
and types. As a result of thousands of years of adaptation to adaptability of beef cattle to assess the suitability of particular
region specific conditions, a much larger variety of livestock genotypes to tropical environment (Prayaga and Henshall,
breeds with diverse and unique traits have been created than 2005). In some cases (e.g. Silanikove, 2000) the physiological
what are currently being used in commercial agriculture basis of adaptation has been investigated in great detail.
(NRC, 1993). However, more commonly such assessments are not made
There is ample evidence that livestock breeds and popula- and information on the genetic parameters for adaptive traits in
tion that have evolved over the centuries in diverse, stressful livestock populations managed in tropical environments are
tropical environments have a range of unique adaptive traits lacking. The purpose of this paper is to review the current state
(e.g. disease and heat resistance, water scarcity tolerance, of knowledge on genetics of adaptation in major livestock
ability to cope with poor quality feed, etc) which enable them to species with emphasis on small ruminants. The genetics and
survive and be productive in these environments (Fitzhugh and related information on breeds of sheep and goats that are
Bradford, 1983; Devendra, 1987; Baker and Rege, 1994; Baker resistant or resilient to a variety of disease infections, feed and
and Gray, 2004). Marked genetic distinction between taurine water scarcity and climatic stressors are reviewed.
and zebu cattle has been reported by MacHugh et al. (1997)
through phylogenetic analysis of microsatellite polymorphisms
with estimated divergence between the two subspecies being 2. Domestication
the order of 610,000–850,000 years. It is possible that reason-
able levels of functional genetic diversity exist between the Keeping and breeding animals was practiced by ancient
two, especially with respect to adaptation (Hanotte et al., societies before the recorded history of animal domestication
2003). A prudent conservation and utilization of the diverse in which our present civilization has its roots. To domesticate
genetic resources is of special concern to livestock breeders so means to adapt the behavior of an animal to fit the needs of
that their special characteristics may be adapted to unforeseen people. Thus, domestication is defined as a process by which a
social/commercial needs, changing climate, and also to population of animals becomes adapted to human and to the
researchers who could study them and expand society's captive environment by some combination of genetic changes
understanding of domestication, selection, genetics and evolu- occurring over generations and environmentally induced
tion (NRC, 1993). The Australian beef industry provides one of developmental events reoccurring during each generation
the best examples of production systems where adapted zebu (Price, 1984). Domestication is the first step of selection and
breeds are utilized through crossbreeding with taurine cattle to has to be distinguished from taming, in that domestication
form synthetic breeds. Prayaga and Henshall (2005) report that means that breeding (by choice of the reproducers and
this system is evolving as an efficient tool to improve the isolation from wild counterparts), care (shelter, food, protec-
productive (meat quality) and reproductive (fertility) perfor- tion against predators) and feeding of animals are more or
 T. Mirkena et al. / Livestock Science 132 (2010) 1–12 3

less controlled by humans (Hale, 1969). It is believed that the Ayalew et al. (2003) compared productivity of indigenous
most important decisive factor for selection during the early breeds of goats (Hararghe Highland and Somali) with that of
times was adaptation (live/survive, reproduce and produce) crossbred (Anglo-Nubian × Somali) goats in Ethiopia and
to a given environment (Gillespie, 1997). The small number concluded the crossbred did not improve households' income
of domesticated species indicates the characteristics required in the mixed crop–livestock production system. The authors
for domestication including traits such as diet, reproduction, indicated that there were increased net benefits per unit of
social relationships and behavior toward man of which most land or labor from mixed flocks (i.e. both indigenous goats
important are a strong gregariousness, feeding regimes that and Anglo-Nubian crosses) under improved management
can be easily supplied by humans, and precocious young compared with indigenous goats under traditional manage-
(Mignon-Grasteau et al., 2005). ment. In flocks using improved management package, the
Consequences of domestication could be investigated crossbreds did not produce more net benefits than indige-
using methods such as comparison of wild and domestic nous goats either in mixed or separate flocks. The improved
stocks, longitudinal analysis of wild animals kept in captivity, management package, however, increased net benefits of
and molecular genetics techniques (Mignon-Grasteau et al., farmers keeping indigenous goats; findings that explained
2005). The consequences of domestication that resulted in the low adoption rate of exotic crosses by smallholder
modifications of many traits determining the capacity of farmers and superior adaptability of indigenous goats to the
adaptation of animals including behavior, physiology and prevailing production system.
morphology include: proportion of white color has increased Karugia et al. (2000) analyzed the impact of crossbreeding
in domestic population as a result of attraction to humans and zebu with exotic cattle breeds for dairy improvement in Kenya
relaxation of natural selection on predation (Pielberg et al., using sector- and farm-level approaches. The agricultural sector
2008); size has been increased in small species to boost meat model showed that dairy technology that involved crossbreed-
quantity but reduced in larger species to make them easier to ing and complementary nutrition and management improve-
handle; fat location has been modified (it is stored under the ments has had a positive impact on Kenyan economy and
skin and around kidneys in wild animals, and in muscle and welfare but this approach ignored important social cost
around the tail in domestic animals); head or brain size has components of crossbreeding. The farm-level approach, how-
decreased in most domestic species; behavior has changed ever, indicated that farm performance was little improved by
quantitatively rather than qualitatively (behavior traits did replacing the indigenous zebu with exotic breeds. On the other
not appear or disappear, but the threshold of their expression hand, this analysis indicated that breeding program that
changed (Price, 1999) indicating that if the opportunity is concentrates on improving the local zebu breeds would
offered to them, domestic species can revert to behaviors improve the financial performance of the farm level with
observed in related wild species, as the genetic variability is still important implications for the conservation of farm animal
present in domesticated populations); relaxation of natural biodiversity. Philipsson (2000) presents several case studies on
selection and natural selection in captivity are partially con- limitations of crossbreeding and/or replacing indigenous with
trolled by humans through determining environmental condi- exotic cattle breeds. For instance, the Sahiwal breed currently
tions (Mignon-Grasteau et al., 2005). suffers from small population size and high degree of
inbreeding due to indiscriminate crossbreeding with exotic;
3. Adaptation to the environment and production system Kenana x Holstein crossbreds with higher exotic blood levels
(75 and 87.5% Holstein) did not keep up with F1 animals, rather
The external environmental stimuli (physical, chemical, had problems with resistance to diseases and suffered from
climatic and biological) to which animals respond interact environmental stresses in Sudan; and in West Indies, where
with their genotypes to determine level of performance. All exotic breeds were introduced, reproductive problems oc-
species respond to changing natural environments through curred due to tick-borne diseases and heat stress resulting in a
altering phenotype and physiology; in livestock production shortage of females for herd replacements. These examples are
the situations become more complex since human interven- in agreement with other studies that emphasize more use could
tion influences both genotype and external environment be made of adaptive characteristics, such as parasite resistance
(King, 2006). The North Ronaldsay, a breed of sheep (Preston and Allonby, 1979) and disease tolerance (Trail et al.,
indigenous to an island in the northeast coast of Scotland 1988).
possesses unique adaptive characteristics. This sheep survive
exclusively on a diet of seaweed and obtain all nutritional 3.1. Adaptation to humans
requirements from limited freshwater and abundant kelp
beds along the shore; mastered the physiological challenge of The process of animal domestication involves adaptation
handling elements present in excess (e.g. sodium) and hence particularly to human and the environment provided. Adapta-
are very salt tolerant in the face of very limited supply of tion to humans is reflected by showing low reactions to humans
freshwater; and are adapted to the very low concentration of (short flight distance for instance) and low fear reactions; low
copper present in Limnaria (their most preferred feed). Other flight times indicate animals with poor temperaments and high
breeds found in Scotland, which normally feed on grass or flight times indicate desirable docile temperament (Prayaga
hay, would die from lack of copper if fed Limnaria (NRC, and Henshall, 2005). Defensive reactions against humans are
1993). Thus, preserving unique qualities in such and many still observed in domestic ruminants even though reduced fear
other livestock breeds will ensure a wealth of genetic of humans is generally considered to be a major component of
resources for future use in basic scientific research and the domestication as routine management procedures (e.g. shear-
advancement of the agricultural sciences. ing, castration, tail docking, dehorning, vaccination, herding
4 T. Mirkena et al. / Livestock Science 132 (2010) 1–12

and transportation in cattle and sheep) can still trigger negative ewes, also influenced how animals attempt to reduce preda-
emotions, such as fear, which are generally considered to affect tion risk (Rieucau and Martin, 2008).
animal welfare negatively (Boissy et al., 2005). Excessive fear
may reduce productivity. For instance, fear-related reactions
affect sexual and maternal behaviors and social dominance 3.3. Adaptation to available feed resources
ability in cattle and sheep (Boissy et al., 2005). Lankin (1997),
using 467 rams and 1617 ewes of the Soviet meat-and-wool Adaptation to periods of feed scarcity can be in one or
breed at various ages studied the influence of environmental more of the following ways: developing low metabolic
factors on the manifestation and diversity of withdrawal from requirement, ability to reduce metabolism, digestive efficien-
man in sheep, and also investigated the polymorphism of cy and ability to utilize high-fiber feed, and deposition of
domestic behavior in 11 breeds. He found that the manifesta- nutrients in the form of fat as feed reserve.
tion and population variability of withdrawal reactions in sheep
are under the influence of farming factors which affect their
feeding behavior. The author concluded that the common 3.3.1. Low metabolic requirements
direction of development of adaptive domestic behavior in Having low metabolic requirements is an advantage if feed
different breeds presupposes the existence of a universal quality and/or quantity are low. The improved temperate
physiological mechanism of ontogenetic inhibition of fear of breeds produce more than indigenous tropical breeds if
man in animals. supplied with high quality feed; however, they lose weight
Boissy et al. (2005) summarized estimates of heritability and fail to survive when fed poor quality grass or straw,
of fear in dairy and beef cattle and sheep. The estimate ranged whereas adapted indigenous animals still grow, give some milk
between 0.09 and 0.53 for dairy cattle while a moderate and reproduce. Adapted tropical animals recycle nutrients
heritability of 0.22 was estimated for reactions to handling in more efficiently than do improved temperate breeds (Bayer
beef cattle. It ranged between 0.28 and 0.48 in sheep. Thus, and Feldmann, 2003) and can also reduce their basic
genetic selection in ruminant livestock based on reduced metabolism during periods of weight loss. The energy require-
fearfulness to increase their adaptive abilities could be as ment of a mammal, a function of body mass0.75, implies the
significant for their welfare as the systems in which they are requirement per kg weight of body tissue in small mammals is
managed. Genetic selection programs for reducing fear greater than that in large mammals which means their
responsiveness to handling could be implemented without metabolic requirements cannot be met by diets rich in cellulosic
adverse effect on other desirable productive traits rather matter. Thus, small ruminants have to balance their compar-
could possibly improve some other adaptive behavioral traits atively higher energy requirements by eating more food of a
such as maternal behavior. higher nutritional value. However, small desert breeds such as
the black Bedouin goat have been found to be efficient
3.2. Behavior towards predators exploiters of high-fiber low quality roughage and their energy
requirement is lower than that predicted from their body mass
As domestication involves human protection of animals in comparison to relatives from non-desert areas (Silanikove,
from predators, they express a lower incidence of anti-predator 1986, 2000). Silanikove (2000) indicated that the energy
behaviors, probably due to relaxed selection on these traits. requirements of five desert goats weighing each 20 kg are at
Consequently, it might be expected that there would be greater about the same level as those of goats from European breed,
losses than wild animals when faced with predation (Mignon- weighing 100 kg. Thus, the ability to maintain a larger number
Grasteau et al., 2005). A few studies in birds have confirmed this of animals on the same area provides an obvious advantage in
hypothesis. Hill and Robertson (1988) showed that captive- terms of survival to the desert goats.
reared pheasants were three times more susceptible to
predation than wild birds. White Leghorn chickens also showed
less anti-predator behavior than Jungle Fowl. 3.3.2. Ability to reduce metabolism
Hansen et al. (2001) studied three breeds of sheep The ability of some mammals to maintain steady body
representing light, medium and heavy weight breeds using weights under less energy intakes than their voluntary intake
blind and carnivore stimuli. The light breed showed longest levels may be due to their ability to reduce metabolism. This
recovery time, the longest flight distance and the tightest ability may vary from species to species or among breeds. For
flocking behavior indicating that lighter sheep breeds display instance, Silanikove (2000) compares the capacity of non-
stronger anti-predator reactions than heavier breeds. In a desert Saanen goats and Bedouin goats to maintain steady
study of Anderson et al. (1998) that compared behavioral body weights when their consumption was restricted but fed
adaptation of sheep, it was found that sheep bonded to cattle on high quality roughages. Saanen goats were able to cope up
from young age remained as one interspecific group when to 20–30% lower than their voluntary intake while the
threatened by trained dog and reduced the chance of being Bedouins tolerated an intake level that was 50–55% lower
attacked by positioning themselves among the cattle and than their voluntary consumption. The Bedouins had a 53%
away from the dog. On the other hand, non-bonded sheep lower fasting heat production under such feed restriction. The
reduced their intraspecific space by flocking together, moved author also reports similar capacity to adjust to a low energy
away from cattle and rendered themselves more exposed to intake by reducing energy metabolism in other herbivores,
predator attack. In wild sheep (e.g., bighorn sheep), however, such as zebu cattle and llama, which are annually exposed to
gregariousness or flocking together was found to increase long periods of severe nutritional conditions in their natural
safety though reproduction status, i.e., lactating versus barren habitats.
 T. Mirkena et al. / Livestock Science 132 (2010) 1–12 5

3.3.3. Digestive efficiency and ability to utilize high-fiber feed adaptation to periodic feed fluctuations in fat-tailed sheep
The digestive efficiency of ruminants and their ability to breeds. Ermias et al. (2002) reported a heritability estimate of
utilize high-fiber feed has been extensively reviewed by 0.72 ± 0.19 for the combined weight of tail and rump fat in
Silanikove (2000). Goats have better digestive efficiency than Menz breed indicating opportunities for selective breeding.
other ruminants with high-fiber low quality forages, and one
of the main reasons is the longer mean retention time in the 3.4. Adaptation to severe (hot/cold) climates
rumen (Devendra, 1990; Tisserand et al., 1991). Goat breeds
indigenous to semi-arid and arid areas are able to utilize low When animals are exposed to heat stress the biological
quality high-fiber feed more efficiently than other types of functions affected include depression in feed intake and
indigenous ruminants, or exotic breeds of goats (Silanikove utilization, disturbances in the metabolism of water, protein,
et al., 1993). The digestive efficiency of desert black Bedouin energy and mineral balances, enzymatic reactions, hormonal
goats fed on roughage diets under controlled environment in secretions and blood metabolites (Marai et al., 2007), resulting
comparison with Swiss Saanen goats (Silanikove et al., 1993; in the impairment of production and reproduction perfor-
Silanikove, 1986) and under exposure to the full impact of mances. The effect is aggravated when heat stress is accompa-
their natural environment (Brosh et al., 1988) was found to nied by high humidity. McManus et al. (2009a) comparing
be superior. Such digestive efficiency of Bedouin goats fed Santa Inês (with three different coat colors — brown, black and
wheat straw (also which enables them to utilize efficiently white), Bergamasca and Santa Inês x Bergamasca breeds of
high-fiber low nitrogen desert pastures) has been observed in sheep in Brazil used physiological traits (sweating, respiratory,
other ruminants only after chemical processing of the straws and heart rates, rectal and skin temperatures) and blood
(Silanikove, 1986). This characteristic is an important asset parameters (PCV, total plasma proteins, red blood cell count,
for their capacity to exist and produce in extreme arid areas and hemoglobin concentration). The authors reported signifi-
and in the face of changing climate (Rischkowsky et al., 2008; cant differences between animals due to breed, skin type and
Tibbo et al., 2008a, b). time of the day and concluded the Santa Inês (hair sheep) with
white color were shown to be better adapted to higher
3.3.4. Fat deposition as feed reserve environmental temperatures while Bergamasca, wool sheep,
Ruminants store energy in adipose tissues when the quality were least adapted. Finocchiaro et al. (2005) reported the
and quantity of feed is ‘adequate,’ and mobilize it to meet genetic correlation between the general additive effect of milk
energy demands during periods of scarcity (Ball et al., 1996; production and the additive effect of heat tolerance to be
Ørskov, 1998; Nigussie et al., 2000; Ermias et al., 2002). In a negative (r = −0.8) during all periods considered indicating
tropical environment, where wet seasons alternate with dry that selection for increased milk production will reduce heat
seasons that are long and characterized with low quantity and tolerance.
quality of pasture, the ability to store fat during ‘favorable’ Turner (1980) extensively reviewed the genetic and
seasons, and its subsequent use for maintenance, pregnancy biological aspects of zebu adaptability and attributed their
and lactation during ‘unfavorable’ season is an essential unique suiting to hot climates to coat, hide, skin, hematolog-
strategy for survival. The ability of sheep to survive in hilly ical characteristics, form, growth, and physiological aspects
environments had been associated with greater fat deposition which are unique genetic attributes of zebu compared to Bos
in the internal fat depots (Kempster, 1980). taurus cattle. Zebu cattle are smooth coated, have primary
Nigussie et al. (2000) compared the patterns of fat depo- hair follicles, have better developed sweat and sebaceous
sition in Horro and Menz sheep breeds of Ethiopia and found glands than B. taurus cattle and can lose more moisture by
that subcutaneous fat and gut fat were the major fat depots in evaporation and hence have the ability to maintain thermal
Menz and Horro, respectively. Genotype variation in amount equilibrium that is a necessary factor for normal function and
of carcass and non-carcass fat was also found: the former performance (Turner, 1980). McManus et al. (2009b),
represented the largest proportion of total fat in Menz while working with five naturalized and two exotic cattle breeds
the latter represented the largest proportion in Horro. in Brazil, found that the Junqueira and Nellore breeds were
However, proportion and distribution of tail fat was similar the most adapted to the climatic condition while Mocho
in both breeds (Nigussie et al., 2000; Ermias et al., 2002). Nacional and Holstein breeds were least adapted.
Comparing the different stages of growth and maturity, Adaptation to cold involves physiological responses affect-
Nigussie et al. (2000) indicated that the growth phase at six ing the thermoregulation of animals by making them more able
months of age that represented the period where a loss in to maintain euthermia during a subsequent cold challenge
body condition and reserves occurred in both breeds co- (Young et al., 1989). Development and retention of a long,
incided with a marked reduction in the proportion of tail fat thick-winter hair coat contributes to thermal insulation. In
compared to all the body fat depot types, indicating its cattle seasonal changes in hair cover are influenced by daily
selective mobilization in order to fill the gap of prevailing photoperiod and ambient temperature (Young et al., 1989). In
energy deficiency. Both breeds probably differ less with temperate environment, the rate of growth of new hair is
regard to fat deposition to anticipate fluctuation in nutrient inversely related to day length while the rate of shedding is
supply, but more in their adaptation to climatic factors. Both associated with thermal status of the animal (Webster, 1974).
may be suited to environments where there is periodic feed Morphological modifications (shorter legs and smaller ears)
fluctuation but Horro is better suited to warmer climates than were observed in growing sheep (Young et al., 1989) and swine
Menz. As tail and rump fat depots are the most responsive (Dauncey and Ingrain, 1986) probably due to reduced blood
ones to nutritional changes, the mass of these depots relative flow in peripheral tissues of animals in the cold. Alterations
to the others, may be the ‘best’ indicator of variations in in the distribution of body fat with enhanced subcutaneous
6 T. Mirkena et al. / Livestock Science 132 (2010) 1–12

deposition have been reported to occur during cold stress to The physiological mechanism that enables desert goats to
increase peripheral insulation (Webster, 1974). Sheep origi- cope with severe water deprivation lies in their ability to
nating and living in cold areas deposit more of their body fat withstand dehydration, and to minimize water losses via
under skin compared to those adapted to warmer areas where urine and feces. The water losses of Barmer and Bedouin goats
degree of heat load is higher (Kempster, 1980; Farid, 1991; by the fourth day of dehydration may exceed 40% of their
Bhat, 1999; Nigussie et al., 2000; Ermias et al., 2002). body weight (Khan et al., 1979a,b; Silanikove, 2000); how-
Nigussie et al. (2000) and Ermias et al. (2002) in their ever, when maintained under an intermittent or a partial
studies that compared Horro and Menz sheep breeds reported watering regimen during the summer, the Barmer goats
that the combined weight of tail and rump fat accounted for a usually gain in body weight at the end of the season.
large proportion of total body fat in the Horro, while the
subcutaneous and intramuscular depot accounted for a large 3.6. Tolerance/resistance to disease
proportion of fat in the Menz confirming preferential deposi-
tion as a result of adaptation to specific environmental con- In livestock, genetic diversity with respect to disease
ditions. Hence, the greater deposits of subcutaneous and resistance is important given that disease-causing organisms
intramuscular fat in the Menz may be adaptation mechanism evolve continuously and develop resistance to drugs. If a new
to the lower temperature of its typically cooler habitat com- strain of a disease or a new disease occurs in a country, animals
pared to the warmer environment at the slightly lower altitude with a narrow genetic base may all be affected whereas in
of the Horro habitat. Sheep adapted to arid conditions often genetically diverse livestock, the chances that some animals
deposit less fat under the skin; almost all of the fat deposited is survive, when others die, increase. Some native livestock are
on the rump and/or the fat tail (Bhat, 1999), also an adaptation less affected by ticks and worms than imported ones. In tsetse
strategy to overcome thermal stress since the relative positions infested areas of Africa, for instance, indigenous cattle have
of these depots (tail and rump fat) do not impede heat loss from developed tolerance to tsetse and trypanosomosis challenge,
the body. It has also been reported that animals adapted to cold whereas imported ones die if not treated with chemicals.
climates have increased circulating erythrocytes and plasma Similarly, local cattle, sheep and goats in West Africa are
concentration of substrates (glucose and free fatty acids) and resistant to heart water, a deadly disease for imported animals
hormones (catecholamines and thyroid) associated with or crossbreds (Bayer and Feldmann, 2003). In this review, we
energy metabolism (Young et al., 1989). are limited to presenting some genetic evidences of tolerance to
or resistance against parasitic and bacterial diseases.

3.5. Adaptation to water scarcity 3.6.1. Parasitic diseases

Resistance to infections with endoparasites involves the
Breeds of ruminants native to arid lands are able to with- initiation and maintenance of responses provoked in the host
stand prolonged periods of water deprivation and graze far to suppress the establishment of parasites and/or eliminate
away from watering sites at times 50 km or more far apart parasite burdens (Baker and Gray, 2004). Resilience/tolerance
(Silanikove, 1994; Bayer and Feldmann, 2003). Livestock is the ability of the host to survive and be productive under
which need little water and do not have to go back to a water parasite challenge (Albers et al., 1987; Woolaston and Baker,
point every day can access larger areas of pastures and thus 1996; Baker and Gray, 2004). The degree of resistance to
get more feed during periods of drought. For example, camels gastrointestinal (GI) nematode parasites has usually been
can undergo as long as 17 days of water deprivation con- assessed in terms of worm counts at necropsy or fecal parasite
suming dry food in the hot summer (Schmidt-Nielsen et al., egg counts (FEC) during an infection period in live animals. In
1956) or can go without drinking water for 30 to 60 days lambs FEC are highly correlated with worm counts (Woolas-
when grazing on green vegetation (Schmidt-Nielsen, 1955). ton and Baker, 1996). Packed red cell volume (PCV) and
There are also donkey, goat, sheep and cattle breeds that can mortality rates have also been used as proxies for resilience
get along without drinking for several days (Bayer and (Baker et al., 2003). Albers et al. (1987) treated both FEC and
Feldmann, 2003). Such animals drink large amounts of water PCV as two different measures of resistance.
quickly but their overall water intake is lower than that of As extensively reviewed by Bishop and Morris (2007),
animals which are watered daily. Reduced water intake genetic differences between host animals in nematode
reduces feed intake and metabolic rate; hence, livestock can parasite resistance have been observed in all major produc-
survive longer during a drought, when feed is very scarce. tion environments and for a variety of parasite species in-
Desert goats have been reported to be the most efficient cluding Haemonchus contortus, Trychostrongylus colubriformis,
among ruminants in terms of ability to withstand dehydra- Teladorsagia circumcincta and various Nematodirus species. In
tion (Silanikove, 1994). The black Bedouin and the Barmer most cases, it is the impact of nematode parasites on the
goats, herded in the extreme deserts of Sinai (Middle East) growing lamb or kid that is of interest. However, nematode
and Rajasthan (India), often drink only once in every 4 days infections are also problematic for reproductive females
(Khan et al., 1979a,b,c; Silanikove, 2000). The Bedouin goats undergoing the stress of late gestation and early lactation
are also able to maintain good level of milk production under and some attention has been given to host genetic variation in
water deprivation. The small black Moroccan goats use a low resistance during the peri-parturient period.
water turnover as a mechanism to economize on water In Africa, several studies compared sheep breeds for
(Hossaini-Hilali et al., 1993); a combined strategy of main- resistance to GI nematodes (Preston and Allonby, 1978, 1979;
taining a frugal water economy and the capacity to endure ILCA, 1991; Baker et al., 1994, 1998, 1999, 2002, 2003). Main
severe dehydration and rapid rehydration. findings indicate that the Red Maasai breed is more resistant and
 T. Mirkena et al. / Livestock Science 132 (2010) 1–12 7

resilient to endoparasites, particularly to H. contortus than lower FEC post-weaning but no breed difference for PCV was
Dorper lambs as reflected by their significantly higher PCV found (Baker et al., 1994, 1998). It is possible that the
(ability to control anemia), lower FEC (lower worm burden), and mechanisms or level of resistance may be different in sheep
lower lamb mortality (Baker et al., 1994, 1998). In another and goats. Goats are predominantly browsers; hence they are
report, Baker et al. (2002) concluded that there is little difference likely under less intense natural selection for resistance
between the two breeds in overall output or efficiency in semi- (Baker et al., 2001).
arid conditions with a low parasite challenge; however, under Many studies have quantified heritabilities of relative
humid conditions where parasite (H. contortus) challenge is high, nematode resistance in sheep usually using FEC as an indicator.
Red Maasai has an output per hectare three times greater than Appropriately transformed FEC is a moderately heritable trait in
the Dorper and is five times more efficient. Baker et al. (1994, lambs and responds to selection (Morris et al., 1997a, 2000;
1998) also compared four crossbred sheep genotypes and found Bishop et al., 1996, 2004; Gruner et al., 2004). FEC tends to be
an additive genetic breed effect for both PCV and FEC indicating less heritable in kids and does (Woolaston et al., 1992; Morris
that crossbreds with higher proportion of Red Maasai blood are et al., 1997b; Mandonnet et al., 2001; Vagenas et al., 2002) but
more resistant; they found no heterosis both for PCV and FEC. the last authors showed that responses to selection for
Other tropical breeds considered resistant based on anec- decreased FEC can be achieved over a short time period. In
dotal evidences that they survive and thrive in the stressful the peri-parturient ewe, FEC is also moderately heritable
environments where they are found under severe disease (Woolaston et al., 1992; Morris et al., 1998; Bishop and Stear,
challenge include the West African Djallonke sheep which may 2001) and is genetically correlated with resistance in the lamb
be resistant to both endoparasites and trypanosomosis (Baker, (Morris et al., 1998). Resistance to different species of
1995; Osaer et al., 1999) and the Garole sheep in India (Ghalsasi nematodes tends to be related, with genetic correlations
et al., 1994). Nimbkar et al. (2003) compared the resistance to between the FEC values arising from different species or genera
H. contortus of F1 Garole crossbred lambs with that of Bannur, of parasites generally being close to 0.5 (e.g. Bishop et al., 2004)
Deccani and 50% Bannur/50% Deccani lambs in India and found or higher in some cases (e.g. Gruner et al., 2004). Douch et al.
that lambs with 50% Garole genes were significantly more (1995), working with Romney ewes in New Zealand, studied
resistant than the other breeds and crosses tested. Boyce et al. the antibody levels against antigens from infective larvae of
(1987) found significant breed differences in FEC and fluke C. curtecei, H. contortus, O. circumcincta, or T. colubriformis and
counts after five breeds of sheep were experimentally infected immunoglobulin G1 (IgG1) specific to C. curtecei or T. colubri-
with Fasciola hepatica. Barbados Blackbelly sheep were the formis and reported heritability values for antibody and IgG1
most susceptible to infection while St. Croix and Florida Native ranging from 0.18 to 0.37 with average of 0.26 (for details, refer
sheep were the most resistant. Wiedosari and Copeman (1990) to the article). Heritabilities of loge (FEC + 100) and dag score
reported relatively high resistance to F. gigantica in Javanese (measure of breach soiling) were 0.28 and 0.13, respectively.
Thin Tail sheep. Roberts et al. (1997a,b) compared the Phenotypic correlations among the 6 antibody and IgG1 traits
resistance to F. gigantica of Indonesian Thin Tail with St. Croix, averaged 0.55, whilst the genetic correlations among them
F2, and F3 crosses between these breeds and concluded that the were even higher, averaging 0.83. Phenotypic and genetic
Indonesian Thin Tail were more resistant than St. Croix. The correlations between antibody or IgG1 and loge (FEC + 100)
authors also stated that resistance may be controlled by a major were all negative and generally small, with genetic correlations
gene with incomplete dominance. In contrast, the Indonesian averaging −0.15. Antibody and IgG1 were positively correlated
Thin Tail sheep were as susceptible to F. hepatica as the Merino genetically with dag score (average value 0.35).
sheep that they were compared with (Roberts et al., 1997a). Studies in sheep to detect quantitative trait loci (QTL) for
Menz and Horro sheep exposed to natural pasture nematode resistance or detect associations with candidate
challenge in the central highlands of Ethiopia showed no genes are now well advanced in New Zealand, Australia, Kenya,
difference in resistance to endoparasites (Baker et al., 1994, US and Europe (UK, France, Italy and Spain) although results are
1998; Tembely et al., 1998; Rege et al., 2002). However, under not readily available in the public domain (Bishop and Morris,
artificial challenge there was some evidence that the Menz 2007) and much success has not been achieved in this area (e.g.
may be more resistant than Horro lambs (Haile et al., 2002). Marshall et al., 2009). Several studies have also looked at
Asegede (1990) compared four Ethiopian sheep breeds (Afar associations between specific genes or markers and FEC.
and Blackhead Somali native to semi-arid lowlands, Horro Coltman et al. (2001) found significant associations with a
and Arsi from humid highlands) for their resistance to endo- microsatellite within the interferon gamma gene in feral sheep,
parasites, mainly H. contortus, at Awassa in southern Ethiopia and various associations with microsatellites in or near the
and found that the Blackhead Somali were the most major histocompatibility complex (MHC) have been observed
susceptible while the Arsi were the most resistant. (Schwaiger et al., 1995; Janssen et al., 2002). The genetic
The evidence for genetic variation for resistance to variation in many aspects of host resistance to nematodes is
endoparasites among goat breeds is limited. As for sheep, it well documented. However, the use of molecular information
is usual that the indigenous goat breeds (e.g. the Alpine goats (mainly QTL) that has long been advocated as a promising tool
in France and the Small East African (SEA) goats in Kenya) are to improve difficult traits, contributed little to genetic improve-
more resistant (Baker and Gray, 2004). The SEA kids were ment in livestock breeding schemes up to now (Bijma, 2009).
more resistant than the Borana1 kids as evidenced by their Reasons are that QTL have often been detected in experimental
crosses, the number of QTL soon becomes impractically
1
This nomenclature has been used in this article (the breed is so named
large, linkage phase between markers and QTL may change
in its home tract of Southern Ethiopia) instead of the offensive term used by over time, and QTL effects may change over time. A major step
the authors of the quoted article. forward will come from the implementation of marker assisted
8 T. Mirkena et al. / Livestock Science 132 (2010) 1–12

breeding value estimation (MA-BVE) using dense maps counts reflected levels of infection, and that lower cell counts
covering the entire genome (Meuwissen et al., 2001). Further- did not indicate any lowered ability to fight infection.
more, microarray studies do have the ability to detect genes Footrot, a bacterial disease caused by Dichelobacter
differentially expressed between ‘resistant’ and ‘susceptible’ (Bacteroides) nodosus (D. nodosus), is a common cause of
animal, with pathways implicated in these differences includ- lameness in both lambs and mature sheep, and it is
ing the development of acquired resistance and the structure of considered to be one of the major welfare problems in
the intestinal smooth muscle (Diez-Tascon et al., 2005). The sheep; it is also a major cause of economic loss. Currently it is
interest in MA-BVE approach is solely in the breeding value of estimated to have economic costs to the UK industry of £31 M
the candidates with the objective to estimate the breeding per annum (Nieuwhof and Bishop, 2005; Bishop and Morris,
value with the highest possible accuracy using all phenotypic 2007). Assessing the genetic control of footrot and subse-
and genomic information. There is no interest in the location or quently breeding for resistance is simple due to the fact that
effect of individual QTL. Using a mixed model, the method gives footrot severity is relatively easily scored under field condi-
an estimate for each marker haplotype and the breeding value tions. As reviewed by Bishop and Morris (2007), Egerton and
of an individual is the sum of the effects of its markers or Roberts (1971) developed a footrot lesion scoring method
haplotypes (Bijma, 2009). using Australian Merino sheep which was later refined by
Raadsma (2000a) into a system that separated clinical signs
into 8 categories. Using this system, Raadsma et al. (1994)
3.6.2. Bacterial diseases demonstrated substantial genetic variation in resistance both
Mastitis is an inflammation of the mammary gland to challenge with virulent isolates of D. nodosus, and also to
resulting from bacterial infections particularly staphylococci. natural challenge. Heritabilities of individual assessments of
Subclinical mastitis is generally diagnosed by an increase in severity of the disease were low to moderate; however,
somatic cell counts (SCC) in the milk of cows and ewes, genetic correlations between indicators were high, approach-
although in goats the predictive value of SCC is less ing unity, and heritability estimates from repeated measure-
established (Bergonier et al., 2003). SCC may also be used to ments approached 0.30. A practical application of this
help select for increased resistance to mastitis though recent approach has been described by Patterson and Patterson
estimates of heritability for SCC are generally low ranging (1989) who successfully bred for enhanced footrot resistance
from 0.10 to 0.20 (Mrode and Swanson, 1996; El-Saied et al., in Merinos. Additional evidence of the feasibility of selecting
1999; Barillet et al., 2001; Rupp et al., 2003; Serrano et al., sheep for footrot resistance using phenotypic observations is
2003; Gonzalo et al., 2003; Legarra and Ugarte, 2005; Bishop given by Skerman and Moorhouse (1987), who report an
and Morris, 2007). The review by Mrode and Swanson (1996) evaluation of lines of New Zealand Corriedale ewes selected
summarized many genetic estimates, concluding that the for enhanced footrot resistance. Therefore, breeding for
heritability of mastitis incidence in dairy cattle is low (∼ 0.04), enhanced footrot resistance using phenotypic assessment
as also is the heritability of SCC (0.11 ± 0.04), but the genetic alone is possible and feasible, provided that footrot is present
correlation between the two is high at ∼0.70. Attention is in the flock (Bishop and Morris, 2007). Less work has been
now turning to the mapping of QTL for SCC in dairy ewes, in done on QTL or genetic marker tests for footrot resistance
both experimental crosses and commercial breeding pro- than on phenotypic assessment of resistance; however,
grams, as described by Barillet et al. (2005). associations between resistance and MHC markers, particu-
In spite of the low heritabilities, numbers of daughters per larly within the MHC class II region, have been published
young sire are generally large enough that breeding values for (Litchfield et al., 1993; Escayg et al., 1997). A specific
SCC can be determined accurately for dairy sires, and effective association with the DQA2 gene has been used in New
selection against SCC can then be applied. In many dairy Zealand as a marker for footrot resistance (Hickford et al.,
countries today, a selection index approach is used, combin- 2004). This test is now commercially available (Hickford,
ing various production and disease traits, including milk yield 2000) as a tool to select more tolerant or resistant animals,
and SCC (Sölkner et al., 1999; Willam et al., 2002). Analyses of without having to expose the animals to infection.
large data sets have shown that there is a small unfavorable
genetic correlation between SCC and first-lactation milk yield 3.7. Resource allocation theory (production vs. fitness traits)
(the weighted estimate from Mrode and Swanson (1996)
being 0.14 ± 0.04, as indicated above), but this correlation can Under selection within a particular environment the
be broken by appropriate genetic selection. Heringstad et al. resources used by the animal are optimally distributed
(2003) reported a 0.19 percentage point annual decrease in between the important traits for breeding and production
cases of clinical mastitis in Norway since the 1990 calf crop, within that environment (Beilharz et al., 1993) implying that
from using index selection, whereas clinical mastitis would any additional selection mediated increase in performance of
otherwise increase with positive selection for milk yield a production-related trait, without a concurrent increase in
alone. There have been questions about the ability of the resources, must lead to declines in other traits, due to
immune system to function effectively against mastitic reallocation of resources (Mignon-Grasteau et al., 2005).
pathogens if SCC is selected downwards genetically. Philips- The decrease in these traits is proportional to the heritability
son et al. (1995) used extensive Swedish industry data to of the “resource allocation factor,” defined by the proportion
investigate this subject, testing for non-linearity in the of resources devoted to production vs. fitness (van der Waaij,
relationship between clinical mastitis values and SCC for 2004; Mignon-Grasteau et al., 2005). In animal production,
sires. From the size and linearity of the genetic relationship negative correlations are observed between production and
between clinical mastitis and SCC, they concluded that cell fitness-related traits, such as fertility and health (Rauw et al.,
 T. Mirkena et al. / Livestock Science 132 (2010) 1–12 9

1998). In lactating animals, poor BCS during the period of system needs often results in the substitution of exotic cattle for
negative energy balance results in decreased fertility (Pryce indigenous breeds. This emanates from the view that most
et al., 2000). It seems that energy allocated to production indigenous livestock breeds are ‘unproductive’ when traits like
cannot be applied to other body functions, resulting in in- milk and beef are considered. This has resulted in many
creased health and fertility problems (Collard et al., 2000). misguided livestock improvement programs importing exotic
Apart from the balance between production and health and breeds which are assumed to be more productive based on their
fertility, the selection environment influences animal perfor- performances in their conducive environments of origin.
mance. For example, the effect of the negative energy balance Adaptive fitness is characterized by survival, health and
has been partly compensated for by improving the environ- reproductive related traits. The wealth of knowledge generated
ment. However, despite these actions, negatively correlated so far indicate that genetic variation for adaptive performance
responses to increased production are becoming stronger: particularly disease resistance is ubiquitous both within and
environmental sensitivity increases and is especially among breeds of livestock indicating that genetic studies on
expressed in decreased fertility. Animals tend to adapt to adaptation of farm animals can be determined at three genetic
the environment they are selected in, which may result in the levels: species, breed and unique genetic variation among
development of a G × E interaction. It is clear that selection for individual animals within a breed. In the warmer tropical areas,
production, without due consideration of the concurrent where pathogens and epidemic diseases are widespread,
undesirable effects on reproduction, leads to problems in climatic conditions are stressful, and feed and water are scarce,
health and fertility. locally adapted autochthonous breeds display far greater level
According to van der Waaij (2004), production gets first of resistance and adaptation due to their evolutionary roots as
priority in contrast to what happens under natural selection compared to imported breeds. There are three pathways of
in resource allocation; when selection is on observed genetic improvement: improvement of local breeds through
production and resource intake is limited, selection pressure purebred selection, breed substitution (by other local breeds or,
is consequently shifted toward resource intake and allocation more frequently, by exotic breeds), and systems of crossbreed-
of proportionally more resources for production and away ing (terminal crosses, rotations, formation of synthetic lines).
from fitness. An insufficient proportion of resources allocated Whichever pathway to follow, choice of the most appropriate
to fitness may result in decreased health, fertility, and energy breed or breeds to use in a given environment or production
available for maintenance, with consequences for reproduc- system should be the first step when initiating a breeding
tion rate and probability of survival. Beilharz et al. (1993) and program and due attention must be given to the adaptive
Knap and Bishop (2000) argue that when resources become performance. Major limitation is that selection for less heritable
limiting, a negative correlation between production traits and traits such as fitness-related traits results in low selection
fitness-related traits will result. Results of a modeling study response due to measurement problems and the underlying
by van der Waaij (2004) indicate that environmental sen- antagonistic biological relationships between productive per-
sitivity, indicated by the negative correlation between ob- formance and adaptive traits. The appropriate strategy for any
served production and survival probability, develops as soon breeding program would therefore be to set suitable selection
as there is metabolic stress. Kolmodin et al. (2002) also found goals, which match the production system rather than
a similar trend in environmental sensitivity in Scandinavian ambitious performance objectives that cannot be reached
dairy cattle. Apparently, the animals with the highest ob- under the prevailing environment. Area-specific approach
served production (trait under selection) tend to be the utilizing the existing resources and taking into account the
animals with poorer values for the “resource allocation prevailing constraints appears to be the only reasonable
factor,” and thus are those with increased environmental sustainable solution. Such approach would also enable in situ
sensitivity. Under artificial selection, when the weighting conservation of farm animal genetic resources, the only viable
given to production is increased, highly specialized animals and practical conservation method in less developed countries
will have difficulty in adapting to changes in their breeding compared to ex situ or cryopreservation approaches. Therefore,
conditions, as no buffer is left to respond to unexpected changes the importance of identifying the most adapted genotype
(and hence care must be taken in introducing highly specialized capable of coping with the environmental challenges posed by
breeds in to the tropical environment) as equilibrium is any particular production system has been indicated.
expected to be reached within a given environment. If the
weighting given to production is disproportionate, resources
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