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Alternative methods for control of

reproduction in small ruminants: A


focus on the needs of grazing industries
J. Alberto Delgadillo* and Graeme B. Martin†‡

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* Centro de Investigación en Reproducción Caprina (CIRCA), Universidad Autónoma Agraria Antonio Narro, Torreón, Coahuila, Mexico

The UWA Institute of Agriculture and School of Animal Biology, University of Western Australia, Crawley, WA, 6009, Australia

Nuffield Department of Obstetrics and Gynaecology, University of Oxford, Oxford OX3 9DU, UK

livestock industries to contribute to the solution of global food supply,


Implications several problems need to be addressed, including: i) consumption of hu-
man food by livestock; ii) use of livestock species and genotypes that are
• 
Extensive grazing systems need simple, low-cost tools for
poorly adapted to the local environment; iii) poor animal health and wel-
managing reproduction that improve productivity while nurturing
fare resulting in suboptimal productivity; iv) inadequate animal nutrition;
future markets. To develop such tools, we need to understand the
and v) significant environmental footprint (Eisler et al., 2014).
biology of the grazing animals in their particular environments.
Reproductive efficiency is obviously an important component of pro-
• Nutrition is the primary challenge because it is limited to grazing, ductivity in the milk and meat industries, so given the issues outlined above,
but nutrition also offers opportunities for improving animal health we need to optimize reproduction in extensive management systems. More-
and carbon emissions. over, reproductive efficiency contributes to the environmental footprint of
• N
 utritional inputs must be coordinated with reproductive events. We ruminant industries because methane emissions can be addressed by focus-
use the term “focus feeding” (optimized management of nutrition, a ing on “emissions intensity”—the amount of methane produced per unit of
concept that has much in common with “nutrient synchrony”). meat or milk. Reproductive waste is major factor in emissions intensity be-
• To coordinate nutritional inputs with reproductive events, we can cause females that fail to reproduce are effectively producing only methane.
control the timing of reproduction by restriction of male access This magnifies the consequences of delayed puberty, low fecundity, delayed
to females and then add precision with the male effect and, ulti- postpartum conception, and high postnatal mortality.
mately, ultrasound for pregnancy detection and measurement of At a superficial level, questions about the management of reproduction
potential litter size. usually lead to a discussion of modern, “high-tech” reproductive technolo-
• Maximum neonatal survival is an imperative, so management of gies. However, even the simplest high-tech tools (e.g., cycle synchronization
the birth site should be a high priority. and artificial insemination) are relevant only to the intensive livestock indus-
tries or to specialist elite breeders (if they exist) in the grazing industries. The
• T
 he power of genetics must underpin any management strategy, with
frontier technologies, such as embryo manipulation or cloning, are not even
breeding goals that take advantage of the strengths of indigenous
on the horizon for almost all managers of extensively grazed ruminants who
genotypes and matching their performance to their environment.
see it as irrelevant because of economics, physical geography, and practical-
ity. In the long term, the grazing industries should benefit from genetic im-
Key words: goats, male effect, nutritional management, sheep, ultrasound
provements that flow down to them as a result of advanced technology, but
the breeding goals have often not been planned with extensive management in
Introduction mind, and in any case, the time delay can be measured in decades. In the short
to medium term, managers of extensively grazed ruminants need tools that
Recently, Eisler et al. (2014) outlined the important role of grazing ru-
are simple, effective, low cost, and applicable on a large scale (Martin, 2014).
minant industries in global food production as we strive to cope with pro-
In addition, livestock industries need to plan with an eye on the future
jections of population growth, demand for more animal protein in devel-
markets of their products, as expressed in the concept of “clean, green,
oping nations, and losses in production capacity because of urbanization,
and ethical” (CGE) management (review: Martin, 2014). In brief: “clean”
biofuel production, climate change, lack of water, and soil degradation.
means reduced usage, if not elimination, of practices that depend on drugs,
The authors argued against headlines claiming that ruminant industries
chemicals, and exogenous hormones (e.g., for controlling reproduction);
cause, rather than solve, such problems. They pointed out, for example,
“green” means minimal damage to the environment (e.g., reducing methane
that foraging animals can, and should, consume foods that humans can-
production); and “ethical” primarily means consideration of animal welfare
not and that crop and livestock systems often complement each other. For
(e.g., surgical castration). Therefore, the development and implementation
of alternative methods for managing reproduction need to take into account
© Delgadillo and Martin. predictable changes in markets and in the regulatory environment.
doi:10.2527/af.2015-0009

Jan. 2015, Vol. 5, No. 1 57


progresses and yet need to be coordinated with seasonal changes in the
Table 1. The characteristics of the extensive grazing
feed supply. The farmers might decide to try to coordinate the period of
industries for small ruminants in southern Australia and
highest demand (lactation) with the period of maximum pasture supply
northern Mexico.
but might then be frustrated by the seasonal breeding patterns of their
Southern Northern
animals. As illustrated in Figure 1, the outcome is often a very large quan-
Australia Mexico
titative mismatch between the availability of feed, particularly energy, to
Production system Sheep Goat
grazing animals and their physiological requirements for energy (Martin
Dominant genotype Merino Indigenous
et al., 2008). Moreover, the cost of supplements is often prohibitive, so
Meat production – local consumption  
even if they are feasible, they must be used with maximum efficiency.

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Meat production – international export  
Milk production  
Fleece production   CGE Management
Extensive grazing, natural forage   of Reproduction—A Possible System
Extensive grazing, improved forage  
Industry problems Over the years, our research teams have turned to basic knowledge of
Wide seasonal variation in quality/quantity of feed   the reproductive physiology and behavior of our animals, particularly the
Reproductive seasonality   processes that the animals developed to cope with changes in their envi-
Males and females together all the year round   ronment over the millions of years of evolution before domestication. In
Technical control of reproduction Ultrasound  reviewing this knowledge base, we searched for ways to manipulate the
only environmental factors that affect the reproductive axis rather than using
High neonatal mortality   hormones, drugs, or complex technology. Given the importance of nutri-
Animal health problems GI nematodes* Brucellosis GI tion in the context of grazing animals, it was no surprise that it became
Blowfly† nematodes* the core management tool (Figure 2). We use the operational term “focus
CAEV‡ feeding” for optimized management of nutrition, a concept that has much
Pressure from community in common with the “nutrient synchrony” described by Hersom (2008) in
Pollution, urbanization, ethics, product health and   the context of intensively fed animals.
safety
The framework in Figure 2 is best viewed as an ideal because full imple-
* 
Resistance to anthelminthic drugs.
mentation of all aspects awaits more research and development, and in some
External parasite (flystrike) – prevention dependent on surgery without anaesthesia.
† 
cases, requires major changes in management. Some of the tools require
‡ 
Caprine arthritis encephalitis virus.
a new level of precision in management, and for others, we would need
to convince farmers to abandon “traditional” practices and, instead, adopt
As we shall see, the need for simple, low-cost tools can complement practices such as separation of males and females and the early mating of
the need for CGE management. In this paper, we will consider this situa- pubertal females. To ensure change of practice, we would need “fourth gen-
tion from the perspectives of goat herds in the sub-tropical north of Mexi- eration” adoption strategies because the traditional top-down approaches to
co and sheep flocks in the southern regions of Australia. There are obvious extension have largely failed livestock industries based on extensive man-
similarities in the physical geography of the two regions, an important agement (for an example, see Sneddon, 2009). Given that it not feasible
consideration because the CGE concept depends on understanding and to expect producers in either country to adopt the whole CGE package in
controlling the responses of the animals to their environment (Table 1). one step, a staged approach with progressive levels of difficulty seems logi-
On the other hand, the socio–economic environment and human geog- cal (Martin, 2014). For individual managers, the optimum series of stages
raphy differ markedly. This need not be a problem because the scientific would depend on the economic, geographical and societal environment.
basis of the CGE concept is relevant to all socio–economic environments, On the other hand, it is important to note that almost all of the interven-
from large rangeland systems in developed economies to smallholders in tions in Figure 2 are simple, “low tech,” and applicable on a large scale.
developing economies. They all need simple and inexpensive manage- Moreover, some of the tools will be familiar to many farmers because they
ment options and to target consumers with discretionary spending power have been known, perhaps even part of normal practice, for many years. A
if they are to get higher prices (Martin, 2014). Therefore, we expect this possible exception is the ultrasound for which the investment in equipment
discussion to be applicable to livestock industries in a wide variety of and skills would be beyond most farmers, and a service industry would be
socio–economic and geographical contexts. required similar to that which has developed rapidly in Australia.
In the next section of this article, we will describe each aspect of the
CGE package in relation to the Australian and Mexican systems. As can
Key Reproductive Problems
be seen from Figure 2, an essential goal is the accurate coordination of
As can be deduced from Table 1, insufficient nutrition is the domi- nutritional inputs with events in the reproductive process because it would
nant problem for both environments. This is to be expected because it is ensure that costly supplements are kept to a minimum while appropriate
the major limiting resource in all animal production systems and it exerts metabolic signals are used to manage the reproductive outcome. At the
strong and specific effects on most events in the reproductive process. Un- simplest level, farmers control timing by managing the presence of the
der extensive management, where nearly all feed comes from pasture and male among females and thus the time of fertile mating. More refined
other forages, a solution to this problem can be difficult to find because the options allow more precise control and offer more opportunities to use
nutritional requirements of the animals change as the reproductive process aspects of male reproductive biology other than gamete supply.

58 Animal Frontiers
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Figure 1. Upper panel: The annual cycle of pasture availability and its relationship with two common alternative reproductive scenarios for sheep in southern Australia.
In Scenario 1, late pregnancy and early lactation coincide with maximum energy availability, the normal situation for temperate genotypes that are strict “short-day breed-
ers.” Scenario 2: Milk production begins when energy availability is at its lowest, a situation that reflects the relationship for temperate genotypes trying to reproduce in
a “mediterranean region” such as southern Australia. Lower panel: The annual cycle of pasture availability during natural grazing and its relationship with alternative
reproductive scenarios for goats in northern Mexico. Scenario 3 is the most common—conception coincides with high food availability, but birth, nursing, and milking
occur when food availability is lowest. In addition, in this scenario, bucks and does remain together all year round. In Scenario 4, the time of conception is modified by
the male effect so that birth, nursing, and milking coincide with the period of high food availability. Scenario 5 is similar to Scenario 1, with the breeding season starting
in September. Modified after Martin et al. (2008) and Sáenz-Escárcega et al. (1991; cited by Delgadillo, 2011).

Jan. 2015, Vol. 5, No. 1 59


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Figure 2. A “Clean-Green-Ethical Package” for managing reproduction in sheep and goats: Periods of “focus feeding” to supply energy and protein are used to control the
reproductive process and improve reproductive success (modified after Martin, 2014). With the advent of “nutritional pharmacology,” forages can also be used to supply
bioactive secondary compounds that improve health and reduce emissions. To accurately synchronize the periods of feeding with specific reproductive events, mating must
be controlled and brief, or ultrasound must be used to classify the mothers based on the age of their fetuses. Finally, the survival of the newborn must be maximized by a
combination of good genetics and good management. Several issues, such as peri-conceptual programming, organ development, and carcase finishing have been covered
by Martin (2014) but not in the present paper where the focus was on control of reproduction.

Restrict the Duration of Births for separation of animals and control of service; and iii) ways to measure
milk production and the growth of kids to generate data for genetic selec-
In extensive production systems in Mexico, it is common for the males tion. The science is long established, so as mentioned above, the real need
to be with the females throughout the year, so there is no control over the is probably a “fourth generation” process to ensure adoption.
timing of mating, births, and lactation. About 70% of pregnancies occur In southern Australia, the situation is less extreme because very few
spontaneously over 4 mo of June through September, and most kids are sheep farmers would leave rams and ewes together year-round. Neverthe-
born between the end of autumn and the end of winter, the worst season less, many farmers like to offer the ewes the “maximum opportunity to
for availability of feed (Figure 1). Most young animals are sold when they conceive” and so will leave the rams in the ewe flock for up to 12 wk.
are only 4 wk old, but young females retained for herd renewal are always However, this apparently self-evident principle does not survive scrutiny
with the males, and many of them become pregnant at a young age when because the actual gains are much smaller than most appreciate (Figure 3).
they are not physiologically mature. A short mating period is the better option for several reasons:
Why do the Mexican farmers leave the males continuously with the fe- i) Selection pressure on female fertility—failure to conceive in a
males? Their reasoning seems to be that they cannot feed the males in con- short mating period should lead to culling;
finement so must graze them with the rest of the herd. It is difficult to know ii) Avoiding the effects of the seasonal declines in feed supply during
how much of this practice is due to tradition. Some farmers try to control mating—small ruminants mate naturally in autumn and, in many
reproduction by fitting the bucks with aprons that prevent copulation, but environments, pasture quality and quantity decline rapidly as au-
this method is prone to failure. Clearly, this problem needs to be addressed tumn progresses; females that fail to conceive early will inevitably
by exploring, for example, the possibility of a cooperative that retains all be attempting to conceive when they have lost body mass, leading
the males for all farmers in a community (it could also incorporate a system to lower conception rates and fecundity;
for genetic improvement). Challenges that need to be considered include iii) Increased value from ultrasound scanning—if conceptions are lim-
i) the need for a system of animal identification; ii) a facility and a process ited to one or two cycles, ultrasound scanning also becomes more

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Figure 3. Theoretical
aspects of mating out-
comes over a series of
three estrous cycles
for a sheep flock. Top:
even assuming a low
value for conception
rate at 70%, it is clear
that that gains for a
third cycle, in par-
ticular, are negligible.
Bottom: cumulative
flock conceptions
comparing 60% ewe
fertility (left) with
90% ewe fertility
(right). Modified after
Martin (2014).

cost-effective because it will allow segregation of females into groups v) Avoiding the “tail”—a cohort of offspring of an even age and size is
carrying zero, one, or two fetuses; non-pregnant females can be culled easier to manage and to market.
for profit and to improve female fertility; segregation of mothers car- To overcome the insecurity of restricted duration of mating, fertility
rying single and multiple fetuses is also the first step toward precision can be detected in individual females. The simplest way is to use a har-
management of the birth environment and “focus feeding” for colos- ness with marking crayons on the males and then change the color to
trum production (see “Maximize Neonatal Survival” below); determine whether the mated (marked) females come back into estrus.
iv) Reduced neonatal mortality—a short, concentrated period of births This low-cost technique works well for sheep and goats. Alternatively, as
allows intensive management of the birth environment to reduce we will discuss below, an ultrasound can be used to identify pregnant and
neonatal mortality, including focus feeding to maximize the pro- non-pregnant females. Both marking crayons and ultrasounds offer data
duction of colostrum (see below); on fertility that can be used for genetic improvement.

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Control the Time of Births practical solutions for extensive management can be difficult to imagine at
present. However, one area that could be resolved relatively quickly is the
Three fundamental aspects of reproduction limit the time of year for quality of male stimulus, as recently summarized in reviews by Fabre-Nys
natural mating: Puberty, seasonal breeding, and postpartum anestrus. One (2014) and Jorre de St Jorre et al. (2014):
of the major causes is the evolution in sheep and goats of brain processes i) The intensity of male behavior—sexually active males improve the
that link their reproduction control systems to changes in the length of the response (Delgadillo, 2011);
night. Despite 10,000 yr or more of domestication, these processes remain. ii) The age and experience of the males (Ungerfeld et al., 2007);
Science has been working for many decades on ways to use exogenous hor- iii) Strategies for stimulating male activity: nutritional supplementa-
mones to overcome this block, and great progress has been made, but these tion and pre-exposure to estrous females have been tested (Unger-

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treatments are not compatible with CGE management. In any case, they are feld, 2007), but night-length treatment seems to be the most effec-
too expensive for extensive management systems. Here we describe two tive, at least for goats (Delgadillo, 2011).
non-pharmacological methods that have proven to be efficient alternatives: These problems and solutions depend on interactions between geno-
type and season, so the genotype of the stimulus male is a major factor
Changing the night length (Jorre de St Jorre et al., 2014). For sheep, but not goats, the first ovulation
Controlling the night length for about 6 wk can be used to “switch on” is not accompanied by estrus so it might be feasible to use a specific geno-
the reproductive centers of the brain in goats and sheep. This is attractive type of male for the male effect and then change the genotype for mating
for farmers with flocks or herds of seasonal genotypes but is probably after ovulation is induced.
limited to the males because they are fewer in number, so implementation
would be more practical than for females. In goats, this technique allows a
reduction in the duration of contact between the sexes without decreasing
Nutrition
fertility (Bedos et al., 2010). The treatment can be used to ensure that the Feed males for fertility
male reproductive axis is working at maximum efficiency when mating
Restricting the duration of mating immediately places pressure on the
is restricted to a brief period, as described above, or for “male effect” is
males, so they need to be managed correctly—adequate numbers, maxi-
used, as described below (Delgadillo, 2011).
mum mass of testis, anatomically sound, healthy, and fit. To maximize
The ‘male effect’ testis mass and therefore sperm production, rams and bucks need to be fed
well for 8 wk before mating (review: Martin, 2014). This is especially im-
In sheep and goats of many genotypes, the sudden introduction of portant if the normal practice is to mate females that have been synchro-
novel males can induce ovulation in females that are reproductively qui- nized by the “male effect” because up to 30% of the females could be in
escent because they are pre-pubertal, out of season, or lactating (reviews: estrus on some days. In Australia and New Zealand, the number of males
Ungerfeld, 2007; Delgadillo, 2011; Jorre de St Jorre et al., 2014). This might have to be increased from 1.5 to 4.0 per 100 females.
“male effect” offers control at two levels: i) the timing of births is no
longer determined by the natural breeding season of the females; and ii) Feed the females for maximum litter size
the induced ovulations are synchronized among the females in the flock
Ovulation rate determines the upper limit of prolificacy, and thus pro-
or herd. The synchrony can be sufficient for artificial insemination (Mar-
ductivity. Ovulation rate is under primary genetic control, so it can be
tin and Scaramuzzi, 1983; Lopez-Sebastián et al., 2014) and is certainly
improved through selection, but the expression of that genetic potential is
sufficient for implementation of CGE management tools such as an ul-
greatly influenced by the nutritional regime before mating (review: Scar-
trasound, focus feeding during pregnancy, and the management of birth.
amuzzi et al., 2011). This is evident from the correlations between body
Equally as important is the simplicity and practicality—putting males in
condition and litter size, but more importantly in the context of focus feed-
the field with the females and then knowing with confidence when the
ing, there is also an acute effect—in sheep and goats, supplementation
females will ovulate, conceive, and give birth.
for only 4 to 6 d in the final stages of the estrous cycle can increase the
However, the male effect is not universally effective—it works best
frequency of twin ovulations by 20 to 30% (Viñoles et al., 2005, 2009;
with moderately seasonal genotypes, and currently, it can only be applied
Fitz-Rodríguez et al., 2009; Zabuli et al., 2010). In the absence of very
when the females are not ovulating spontaneously. Even in situations
precise control over the time of ovulation, supplementation for only 4 to
where the response is reliable, there can be wide variation among females
6 d is not realistic, and it is necessary to offer the supplement for an entire
in delay to the induced ovulation, in the incidence of short cycles follow-
cycle to ensure that all females have an opportunity to respond. If supple-
ing the first ovulation, and in the outcomes of mating (reviews: Unger-
ments are not feasible, high quality pasture or even cactus cladodes can be
feld, 2007; Delgadillo, 2011). The synchrony of ovulation declines over
used (Viñoles et al., 2009; Rekik et al., 2012).
subsequent weeks, so focus feeding for fecundity still cannot be refined
to the theoretical minimum of 4 to 6 d (see below): 7-d supplementation Nutrition—more than energy and protein
is needed in goats (Fitz-Rodríguez et al., 2009) and at least 11 d in sheep
because ovulations are dispersed by the short-cycle phenomenon (Martin Generally, the relationship between nutrition and reproductive perfor-
and Scaramuzzi, 1983; Nedelkov et al., 2013). Despite these limitations, mance is explained by energy supply (review: Martin et al., 2008), but we
there is a significant saving in feed costs over the requirements for mating need also to consider benefits that can be derived from other components
over a full cycle. of fodder plants, particularly “bioactive” secondary compounds. Interest
Some of the problems with the male effect will not be resolved without in this area was initially stimulated by restrictions on the use of growth-
significantly more research and development (review: Martin, 2014), so promoting antibiotics and by the need to combat drug-resistant gastroin-

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In northern Mexico, goats are under extensive management conditions, and males and females are usually kept together all year round (left). Animals eat only available
natural vegetation (right).

testinal parasites (review: Durmic and Blache, 2012). In Australia, for ex- in-hand with any plan to improve fecundity through genetic selection or
ample, more than 100 Australian native plant species have been assessed focus feeding. In addition, if neonates are lost because of mismanage-
on their potential to improve rumen function, gut health, nutritional value, ment, then any investment in genetic improvement has been wasted.
and biomass productivity, and many of them can also reduce methane
emissions (Revell et al., 2008; Durmic et al., 2010), so they contribute in Maximize colostrum production
several ways to the CGE credentials of the industry. This concept is obvi- In addition to the nutritional and immunological benefits, colostrum in
ously applicable outside Australia (e.g., Akkari et al., 2014) so should be the gut improves the ability of the newborn to recognize its mother and
investigated within local contexts. thus contributes to the establishment of the mother–young bond (Gour-
This concept can be extended to other environments. In Mexico, for saud and Nowak, 1999; Ramírez-Vera et al., 2012a). With the imple-
example, local forage plants are known to contain bioactive compounds mentation of restricted, brief mating periods and ultrasound assessment
that affect reproduction, as demonstrated by studies on the effects of mes- of pregnancy, focus feeding can be used in the last week of gestation to
quite in rats (Retana-Márquez et al., 2012). There are many possibilities increase greatly the amount of colostrum available at birth (review: Ban-
for research of this type that would be relevant for the extensively man- chero et al., 2006) and improve neonatal survival (Goodwin and Norton,
aged goat industry in this part of the world, and the concept needs to be 2004; Ramirez-Vera et al., 2012b).
investigated for other environments.
Manage the birth environment
Ultrasound Scanning The simplest (and often least expensive) strategy for improving neo-
natal survival is to better manage birth. The provision of a calm envi-
Ultrasound pregnancy diagnosis is now a routine procedure in small
ronment, shelter, feed, and water close to the birth site will increase the
ruminants, even for the large flocks and herds that are typical in Australia
amount of time the mother spends at the birth site and therefore improve
and New Zealand, with numerous commercial operators offering reason-
the development of the mother–young bond (Nowak, 1996).
able rates and accurate results. The rate of adoption in Australia was the
subject of a survey of 1,000 sheep farmers by Jones et al. (2014) who
found that pregnancy scanning was used by more than 40% of farmers, of Management of Puberty
which 18% scan for litter size. In Mexico, male and female goats remain together throughout the year,
Even at the simplest level, the identification of pregnant and non-pregnant so young females often mate at 7 to 10 mo of age, with apparent conse-
females, the information is valuable because it offers: i) an opportunity for re- quences being high rates of abortion and delayed growth and a decrease
mating if there has been a disaster; ii) culling for improvement of fertility; and in future productivity. In Australia, there is the opposite problem because
iii) planning of conditions for birth. If conceptions are limited to two cycles, the traditional practice is to manage the sexes in separate groups but de-
ultrasound scanning becomes more cost-effective because it will provide two lay the mating of young females until they are 18 to 24 mo old. This
extra levels of information: i) identification of single-bearing and multiple- approach might avoid the issues encountered in Mexico but has broader
bearing females, allowing segregation into separate management groups; and consequences for the industry such as high emission intensity, delays in
ii) estimation of age of fetus to increase the precision of management of birth. genetic progress, and reduction of the effective size of the breeding flock.
The ideal solution lies between these two extremes, with young females
Maximize Neonatal Survival mating at the optimum stage of their development. This is a complex
problem, with a mix of sociological and biological causes and solutions:
High rates of neonatal mortality have obvious consequences for profit-
i) Sociological: Among Australian farmers, there is a widely held belief
ability and also raise questions about the ethical credentials and the emis-
that mating young animals impairs their performance in subsequent
sion intensity of the industry. Neonatal mortality, therefore, goes hand-

Jan. 2015, Vol. 5, No. 1 63


Importantly, genetic improvement is ideal for extensive grazing indus-
tries because it is a simple, low-cost management tool and the benefits are
cumulative. Moreover, it is becoming feasible for the extensively man-
aged industries because we are on the verge of a technology-led revolu-
tion in the generation of high quality data, such as electronic identifica-
tion, DNA pedigrees, and the automatic recording of body weights and
litter size. It will soon be possible to assess large numbers of animals for
a wide variety of production traits under extensive production systems.

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Conclusions
For industries based on extensively managed, grazing small ruminants,
there is a wide range of simple, effective low-cost tools for managing re-
In Australia, about 80% of breeding ewes are the Merino genotype, with most man- production, and they can be combined into a CGE scheme that is also re-
aged in improved pastures that are nevertheless very seasonal in productivity. Most sponsive to predicted changes in markets and the regulatory environment.
of these ewes only have a single lamb, but focus feeding and genetics can improve Many of tools can be implemented now, with specific choices based on lo-
the frequency of twins. This can then precipitate other problems such as poor co-
cal geographical and socio–economic constraints. A primary focus must be
lostrum production and ewe–lamb bonding. Photo by Dr. Nui Milton, Casalana,
Western Australia. maximizing the value of feed inputs from grazing, but there is now the pos-
sibility of an extra dimension in the choice of forages because some plant
years, a position supported by the Mexican experience; however, re- species can reduce carbon emissions and improve health. This area deserves
search in New Zealand shows that this is not that case with good research in many countries because the outcomes will help address the in-
management during the first year of life (Kenyon et al., 2004); ternational problem of the environmental impact of ruminant industries.
ii) Biological: In young females, the timing of puberty is determined The second major factor is control of the timing of mating, births, and lac-
by body mass, condition score, and season (night length), with body tation in relation to the seasonal availability of fodder. At the most basic level,
mass being the primary factor and maximizing weight gain after the males and females need to be separated for all except the mating period.
birth therefore considered essential. It has long been thought that At a second level, the “male effect” offers excellent precision over the timing
the active component of body mass was fat, but recent work with and synchrony of births. The most complex modes of implementation of the
sheep suggests that genetic selection for accelerated muscle growth “male effect” are still subject to basic research and development.
can also advance puberty (Rosales-Nieto et al., 2013), offering dual For some components of the CGE scheme, the priority is to work on
advantages for the efficiency of meat production as well as reproduc- genetic improvement of the process, and for others, the priority is adop-
tion. The male effect, ultrasound, and birth-site management are also tion processes rather than the science of reproduction. The extensive graz-
obvious targets for managing puberty and the reproductive perfor- ing industries have a poor record of transformation from traditional prac-
mance of young females, as outlined above for adult females. tice to the modern, science-based management that is essential if we are to
Again, there should be positive outcomes for genetic improvement if young feed and clothe the world without destroying the planet.
females are under pressure to conceive at a reasonably early opportunity.
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In vitro evidence that the pastoral Artemisia campestris species exerts an an-
In the discussion above, most of the focus was on the physiological, thelmintic effect on Haemonchus contortus from sheep. Vet. Res. Commun.
behavioral, and managerial aspects of reproduction. The possibility of 38:249–255.
genetic improvement of female fertility has been mentioned, and geno- Banchero, G.E., R. Perez-Clariget, R. Bencini, D.R. Lindsay, J.T.B. Milton, and
type has been acknowledged as a frequent constraint to implementation G.B. Martin. 2006. Endocrine and metabolic factors involved in the effect of
nutrition on the production of colostrum in female sheep. Reprod. Nutr. Dev.
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sheep, Martin and Greeff (2011) concluded that genetic selection could be Bedos, M., J.A. Flores, G. Fitz-Rodríguez, M. Keller, B. Malpaux, P. Poindron
used to enhance the power of the male effect, increase fecundity, increase and J.A. Delgadillo. 2010. Four hours of daily contact with sexually active
fertility, increase colostrum production, and enhance mother–young bond- males is sufficient to induce fertile ovulation in anestrous goats. Horm. Behav.
ing. In addition, there needs to be a strong focus on resistance to para- 58:473-477.
Delgadillo, J.A. 2011. Environmental and social cues can be used in combination
sites (Karlsson and Greeff, 2012). These traits could be incorporated into to develop sustainable breeding techniques for goat reproduction in the subtrop-
Australia’s industry-wide program for genetic selection that is based on ics. Animal 5:74–81.
published breeding values for important traits for both sheep and goats Durmic, Z., and D. Blache. 2012. Bioactive plants and plant products: Effects on
(http://www.mla.com.au/Livestock-production/Genetics-and-breeding). animal function, health and welfare. Anim. Feed Sci. Technol. 176:150–162.
By contrast, Mexican goat production does not have this sort of plan and, Durmic, Z., P. Hutton, D.K. Revell, J. Emms, S. Hughes, and P.E. Vercoe. 2010. In
vitro fermentative traits of Australian woody perennial plant species that may
in fact, does not even have a simple system for recording basic perfor- be considered as potential sources of feed for grazing ruminants. Anim. Feed
mance measures (e.g., milk production and kid growth rate). It is impor- Sci. Technol. 160:98–109.
tant to move in this direction, perhaps beginning with the introduction of Eisler, M.C., M.R.F. Lee, J.F. Tarlton, G.B. Martin, J. Beddington, J.A.J. Dungait,
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(London) 507:32-34. About the Authors
Fabre-Nys, C. 2014. L’effet mâle chez les ovins: Resultats récents sur l’origine de
la variabilité de la réponse et les mécanismes centraux impliqués. In: Curso Dr. J. Alberto Delgadillo earned his
Internacional VII– Reproducción de Rumiantes– Innovaciones en Reproduc- Ph.D. at the Université des Sciences et
ción Animal. Programa de Ganadería, Colegio de Postgraduados, Montecillo, Techniques de Languedoc II at Montpel-
Mexico. p. 266. lier, France. He is currently Professor at
Fitz-Rodríguez, G., M.A. De Santiago-Miramontes, R.J. Scaramuzzi, B. Malpaux, the Departamento de Ciencias Médico
and J.A. Delgadillo. 2009. Nutritional supplementation improves ovulation and Veterinarias and Chair of the Centro de
pregnancy rates in female goats managed under natural grazing conditions and Investigación en Reproducción Caprina

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exposed to the male effect. Anim. Reprod. Sci. 116:86–94. (CIRCA) at the Universidad Autónoma
Goodwin, N., and B.W. Norton. 2004. Improving doe nutrition immediately prior Agraria Antonio Narro. He teaches the bi-
to kidding increases kid survival. Proc. Aust. Soc. Anim. Prod. 25:233. ology of reproduction and the control of
Goursaud, A.P., and R. Nowak. 1999. Colostrum mediates the development of reproduction in farm animals to graduate
mother preference by the new born lamb. Physiol. Behav. 67:49–56. and veterinary students. The main focus of
Hersom, M.J. 2008. Opportunities to enhance performance and efficiency through his research is the physiology of goat re-
nutrient synchrony in forage-fed ruminants. J. Anim. Sci. 86:E306–E317. production and the development of natural methods to control reproduction
Jones, A., M. Curnow, and A. van Burgel. 2014. Report on the national producers of goats managed under extensive grazing conditions in subtropical latitudes.
and service providers surveys. Cooperative Research Centre for Sheep Industry
Innovation, Armidale, Australia. Dr. Graeme B. Martin earned his Ph.D. at
Jorre de St Jorre, T., P.A.R. Hawken, and G.B. Martin. 2014. New understanding the University of Western Australia (UWA)
of an old phenomenon: Uncontrolled factors and misconceptions that cast a and then worked in France and the UK for
shadow over studies of the ‘male effect’ on reproduction in small ruminants. 5 yr. He is now Professor (Chair) at UWA
Turk. J. Vet. Anim. Sci. 38:625–636. and a member of the UWA Institute of Ag-
Karlsson, L.J.E., and J.C. Greeff. 2012. Genetic aspects of sheep parasitic diseases. riculture. His research team mostly works
Vet. Parasitol. 18:272–278. on how environmental factors influence the
Kenyon, P.R., S.T. Morris, N.R. Perkins, and D.M. West. 2004. Hogget mating use reproductive system. He is also committed
in New Zealand– a survey. Proc. N.Z. Soc. Anim. Prod. 64:217–222. to science communication and to the use of
Lopez-Sebastián, A., M. Coloma, A. Toledano, and J. Santiago-Moreno. 2014. Hor- science to transform industry. Since 2004,
mone-free protocols for the control of reproduction and artificial insemination he has pioneered the concept of “clean,
in goats. Reprod. Domest. Anim. 49 (Suppl. 4):22–29. green, and ethical management of live-
Martin, G.B. 2014. An Australasian perspective on the role of reproductive tech- stock” and promoted it nationally and inter-
nologies in world food production. Adv. Exp. Med. Biol. 752:181–197. nationally. He also chairs the UWA Future Farm 2050 project.
Martin, G.B., D. Blache, and I.H. Williams. 2008. The costs of reproduction. In: Correspondence: graeme.martin@uwa.edu.au
W.M. Rauw, editor, Resource allocation theory applied to farm animals. CABI
Publishing, Oxford, UK. p. 169.
Martin, G.B., and J.C. Greeff. 2011. Genetic frontiers in the development of ‘clean, The potential to capitalise on plant diversity for livestock health and productiv-
green and ethical’ management systems for the extensive sheep industry. Proc. ity. In: J.F. Skaife and P.E. Vercoe, editors, Harvesting knowledge, pharming
Assoc. Adv. Anim. Breed Genet. 19:143–150. opportunities. Cambridge Univ. Press, Cambridge, UK. p. 36.
Martin, G.B., and R.J. Scaramuzzi. 1983. The induction of oestrus and ovulation Rosales-Nieto, C.A., M.B. Ferguson, C.A. Macleay, J.R. Briegel, D.A. Wood, G.B.
in seasonally anovular ewes by exposure to rams. J. Steroid Biochem. 19:869– Martin, and A.N. Thompson. 2013. Ewe lambs with higher breeding values for
875. growth achieve higher reproductive performance when mated at age 8 months.
Nedelkov, K., N. Todorov, and M. Simeonov. 2013. Effect of focused flushing at Theriogenology 80:427–435.
the end of the anticipated normal luteal phase on synchronization of oestrus by Scaramuzzi, R.J., D.T. Baird, B.K. Campbell, M.A. Driancourt, J. Dupont, J.E.
introduction of ram in the flock. Bulg. J. Agric. Sci. 19:1085–1092. Fortune, R.B. Gilchrist, G.B. Martin, K.P. McNatty, A.S. McNeilly, P. Mon-
Nowak, R. 1996. Neonatal survival: contributions from behavioural studies in get, D. Monniaux, C. Viñoles, and R. Webb. 2011. Regulation of folliculogen-
sheep. Appl. Anim. Behav. Sci. 49:61-72. esis and the determination of ovulation rate in ruminants. Reprod. Fertil. Dev.
Ramírez-Vera, S., A. Terrazas, J.A. Delgadillo, N. Serafín, J.A. Flores, J.M. Elizun- 23:444–467.
dia, and H. Hernández. 2012a. Feeding corn during the last 12 days of gestation Sneddon, J.N. 2009. Identifying and exploiting opportunities for ‘clean, green and
improved colostrum production and neonatal activity in goats grazing subtropi- ethical’ animal production. Agrociencia 13:51–58.
cal semi-arid rangeland. J. Anim. Sci. 90:2362–2370. Ungerfeld, R. 2007. Socio-sexual signalling and gonadal function: Opportunities
Ramírez-Vera, S., A. Terrazas, J.A. Delgadillo, J.A. Flores, N. Serafín, J. Vielma, for reproductive management in domestic ruminants. In: J.I. Juengel, J.F. Mur-
G. Duarte, I.G. Fernández, G. Fitz-Rodríguez, and H. Hernández. 2012b. Inclu- ray, and M.F. Smith, editors, Reproduction in domestic ruminants VI. Notting-
sion of maize in the grazing diet of goats during the last 12 days of gestation ham Univ. Press, Nottingham, UK. p. 207.
reinforces the expression of maternal behaviour and selectivity during the sen- Viñoles, C., M. Forsberg, G.B. Martin, C. Cajarville, J. Repetto, and A. Meikle.
sitive period. Livest. Sci. 148:52–59. 2005. Short-term nutritional supplementation of ewes in low body condition
Rekik, M., A.M. Gonzalez-Bulnes, N. Lassoued, H. Ben Salem, A. Tounsi, and I. affects follicle development due to an increase in glucose and metabolic hor-
Ben Salem. 2012. The cactus effect: An alternative to the lupin effect for in- mones. Reproduction 129:299–309.
creasing ovulation rate in sheep reared in semi-arid regions? J. Anim. Physiol. Viñoles, C., A. Meikle, and G.B. Martin. 2009. Short-term nutritional treatments
Anim. Nutr. 96:242–249. grazing legumes or feeding concentrates increase prolificacy in Corriedale
Retana-Márquez, S., F. García Aguirre, M. Alcántara, E. García-Díaz, M. Muñoz- ewes. Anim. Reprod. Sci. 113:82–92.
Gutiérrez, M. Arteaga-Silva, G. López, C. Romero, P. Chemineau, M. Keller, Zabuli, J., T. Tanaka, W. Lu, and H. Kamomae. 2010. Intermittent nutritional stimu-
and J.A. Delgadillo. 2012. Mesquite pod extract modifies the reproductive lus by short-term treatment of high-energy diet promotes ovarian performance
physiology and behavior of the female rat. Horm. Behav. 61:549–558. together with increases in blood levels of glucose and insulin in cycling goats.
Revell, D.K., Z. Durmic M. Bennell G.C. Sweeney, and P.E. Vercoe. 2008. The in Anim. Reprod. Sci. 122:288–293.
situ use of plant mixtures including native shrubs in Australian grazing systems:

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