Produced for:

Sheep Easy Breeding Group

Breeding easier-managed sheep

Judith Collins and Joanne Conington

SAC, West Mains Road, Edinburgh, EH9 3JG, Scotland.

Contents
1 Summary................................................................................................................................. 3
1.1 Recommendations........................................................................................................... 4
2 Introduction............................................................................................................................ 5
2.1 Potential characteristics of ‘easy care’ sheep ................................................................. 5
3 Breeding for resistance to disease......................................................................................... 6
3.1 Nematodes ...................................................................................................................... 7
3.1.1 Faecal egg count....................................................................................................... 7
3.1.2 Faecal consistency scores and dag scores ................................................................ 9
3.1.3 Phenotypic and genetic markers for nematode resistance...................................... 10
3.2 Footrot........................................................................................................................... 12
3.2.1 Genetic variation .................................................................................................... 12
3.2.2 Footrot lesion scoring............................................................................................. 13
3.2.3 Use of molecular genetic markers.......................................................................... 13
3.3 Mastitis ......................................................................................................................... 13
3.3.1 Milk Somatic Cell Counts...................................................................................... 14
3.3.2 Molecular technologies to quantify mastitis resistance ......................................... 16
3.4 Blowfly strike ............................................................................................................... 18
4 Breeding for enhanced flock fertility ................................................................................. 19
4.1 Ewe fertility .................................................................................................................. 20
4.2 Litter size ...................................................................................................................... 20
4.3 Scrotal circumference ................................................................................................... 21
4.4 Enhanced ram sexual performance ............................................................................... 22
5 Breeding for improved ewe maternal ability and lamb survival..................................... 23
5.1 Maternal ability traits.................................................................................................... 25
5.1.1 Ease of parturition.................................................................................................. 25
5.1.2 Care of the newborn ............................................................................................... 26
5.1.3 Maternal behaviour score....................................................................................... 28
5.1.4 Milk production...................................................................................................... 29
5.2 Lamb traits .................................................................................................................... 30
5.2.1 Lamb survival ........................................................................................................ 30
5.2.2 Gestation length ..................................................................................................... 31
5.2.3 Parturition............................................................................................................... 32
5.2.4 Time taken to stand and suck ................................................................................. 32
5.2.5 Ewe recognition and attachment ............................................................................ 33
5.2.6 Cold resistance ....................................................................................................... 33
6 Breeding for other traits...................................................................................................... 34
6.1 Tolerance to food shortages.......................................................................................... 34
6.2 Longevity...................................................................................................................... 35
6.2.1 Teeth and bone ....................................................................................................... 36
6.3 Body composition......................................................................................................... 36
6.4 Wool shedding .............................................................................................................. 36
6.5 Temperament ................................................................................................................ 37
7 Conclusions ........................................................................................................................... 38
7.1 Breeding for resistance to disease................................................................................. 38
7.1.1 Nematodes.............................................................................................................. 38
7.1.2 Footrot .................................................................................................................... 38
7.1.3 Mastitis................................................................................................................... 39
7.1.4 Blowfly strike......................................................................................................... 39
7.2 Breeding for enhanced flock fertility............................................................................ 39
7.2.1 Ewe prolificacy ...................................................................................................... 39

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 7.2.2 Scrotal circumference............................................................................................. 40
7.2.3 Ram serving capacity ............................................................................................. 40
7.3 Breeding for enhanced maternal ability and lamb survival .......................................... 41
7.4 Breeding for other traits................................................................................................ 42
7.5 Application to the UK sheep industry........................................................................... 43
7.6 Recommendations......................................................................................................... 44
8 References ............................................................................................................................. 45
9 Appendices............................................................................................................................ 72
9.1 Footrot scoring system.................................................................................................. 72
9.2 Molecular markers for mastitis resistance in cattle ...................................................... 72
9.3 Maternal behaviour scores at tagging ........................................................................... 75
9.4 Trait checklist ............................................................................................................... 76

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1 Summary

To maintain the financial viability of UK sheep systems in recent years it has been necessary to reduce
labour costs. Better handling and management practices enable greater numbers of sheep to be looked
after per labour unit but less time for specific tasks may lead to reduced input in areas such as
supplementary feeding, health care and supervision and inspection of stock. Furthermore, modern
farmed breeds, bred and raised under intensive conditions, may not be adequately adapted for more
extensively managed systems and may not be able to survive and thrive in situations where there are
fewer human inputs. Animal welfare could consequently be compromised due to insufficient animal
care.

It is widely recognised that some breeds are easier to look after compared to others, in particular at
lambing time. However, many farmers may be reluctant to substitute their existing breeds for easier-
managed ones entirely, to prevent the loss of other important characteristics such as carcass
conformation and growth. For this reason, new breeding tools are required, to facilitate within-breed
selection for traits that confer greater efficiency and easier management.

The purpose of this fact-finding study was to investigate the key components of ‘easy care’ breeding
strategies and the best approach for the development of practical breeding programmes to achieve
breeding for easier-managed sheep. The following groups of candidate traits that may confer better
adaptation to more extensive management systems were considered:
• Resistance to disease (nematodes, footrot, mastitis, blowfly strike and others)
• Enhanced flock fertility (ewe fertility, litter size, ram sexual capacity)
• Improved maternal ability and lamb survival
o maternal traits (lambing ease, care of the newborn, milking ability)
o lamb traits (lamb survival, gestation length, parturition, neonate behaviours, cold
resistance)
• Tolerance to food shortages
• Longevity (including teeth and bone)
• Body composition
• Wool shedding
• Temperament

Some of these traits are already used in UK sheep breeding schemes (e.g. ewe longevity is predicted in
breeding programmes for hill sheep although it is not recorded separately in practice). Other traits are in
the pipeline to be included in breeding programmes (e.g. direct lamb survival), and others are still being
researched (e.g. resistance to footrot, nematodes and mastitis).

Difficulties that may be encountered when trying to incorporate some of the traits into breeding
programmes include:
• insufficient evidence for genetic variation in the trait concerned (e.g. bone quality);
• insufficient consensus or knowledge about the ‘best’ trait to be recorded to achieve the breeding
goal (e.g. resistance to nematodes);
• traits that are difficult, time-consuming or expensive to record (e.g. behavioural traits);
• the fact that ‘hidden’ benefits of trait improvement may not immediately appeal to some breeders
because their cumulative benefits are not realised until some time in the future (e.g. longevity).

However, most of the traits reviewed could potentially be used in UK sheep breeding programmes. The
use of genetic information from related animals to predict easier-managed traits would be particularly
useful where the heritability is low or if the traits are difficult to record.

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1.1 Recommendations

Some of the traits considered in the report could be recorded immediately. These include: i) lamb
survival, ii) ewe longevity, iii) ewe fat levels, iv) ‘gestation’ length, v) lambing ease and vi) ram scrotal
circumference. The incorporation of these into indexes would require varying degrees of programming
effort including full genetic analyses of traits iv), v) and vi) and the identification of associations with
other traits that are part of the breeding objective. Faecal Egg Count could also be recorded now but as
the application of the use of this indicator trait for nematode resistance in the UK is yet to be established
(e.g. when to sample, how many sampling times, recommended levels of pasture contamination before
sampling etc.), in order to make the best use of this technology it may be preferable to wait until this
knowledge becomes available.

For flocks that have problems with mastitis, the use of Somatic Cell Count (SCC) could be instrumental
in improving resistance to the disease. Further research is needed to quantify the genetic variation in
this trait and to develop a protocol for its inclusion into breeding programmes.

More work is required (which is currently underway) before molecular tests for resistance to disease
(footrot, nematodes, mastitis) can be used in practice.

A promising trait for easy care systems is ram libido, as defined by ram serving capacity. The
verification of existing tests from other research groups should be done using UK sheep breeds to
establish a useful protocol that could be used in the field. Further research work is needed before the
commencement of recording.

Further research is also needed for most of the behavioural traits, including the use of either physical or
physiological proxy traits.

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2 Introduction

To maintain the financial viability of UK sheep systems in recent years it has been necessary to reduce
labour costs. Better handling and management practices enable greater numbers of sheep to be looked
after per labour unit (it is now common for each shepherd to be responsible for more than 1000 sheep)
but less time for specific tasks may lead to reduced input in areas such as supplementary feeding, health
care and supervision and inspection of stock. Furthermore, modern farmed breeds, bred and raised
under intensive conditions, may not be adequately adapted for more extensively managed systems and
may not be able to survive and thrive in situations where there are fewer human inputs. Animal welfare
could consequently be compromised due to insufficient animal care.

It is widely recognised that some breeds are easier to look after compared to others, in particular at
lambing time. For example Blackface ewes require less intervention at lambing compared to Suffolks,
and lambs take a shorter time to stand after birth (Dwyer and Lawrence, 2005). However, many farmers
are reluctant to substitute their existing breeds for easier-managed ones entirely, to prevent the loss of
other important characteristics such as carcass conformation and growth. For this reason, new breeding
tools are required, to facilitate within-breed selection for traits that confer greater efficiency and easier
management.

To work towards this goal, the ‘Sheep Easy’ breeding group was set up in 2004, comprising breeders of
Llyen, Blackface, Easicare, Romney, Texel and Suffolk sheep. The aim of this group is to share
common interests in selective breeding for ‘easy care’ traits, and to work towards the development of
appropriate breeding strategies for easier-managed sheep.

There exists a dearth of genetic information on this subject in UK sheep breeds. In addition, several of
the traits involve some degree of subjective assessment. Therefore, the development of robust scoring
methods is crucial before any genetic analyses can be adequately implemented or interpreted, prior to
their inclusion in breeding programmes. The purpose of this fact-finding study is to investigate the key
components of ‘easy care’ breeding strategies and the best approach for the development of practical
breeding programmes to achieve breeding for easier-managed sheep. It is envisaged that this initial
study will lead to the development of a further collaboration with the industry partners which in turn
will lead to the development of new selection indices and breeding recommendations for the sheep
industry. It is also hoped that this project may stimulate breeders to record a wider set of traits than is
currently undertaken with a view to their ultimate inclusion in selection indices for easier-managed,
more efficient sheep.

2.1 Potential characteristics of ‘easy care’ sheep

The trend in Europe towards more extensive rearing conditions in areas of intensive production and
towards the maintenance of farming activity in less favourable regions will lead to considerable changes
in rearing management.

Greater emphasis will be placed on the adaptation of animals to their environment and on their
behavioural response to different stresses. It will be particularly important to achieve harmonious
establishment of mother-young relationships. Animals will have to be accustomed to the presence of
human beings even if the contact periods are only short, because adverse responses can lead to stress.
Reduced levels of supervision may mean that disease, injury and parasitism, for example, go undetected
and untreated.

Genetic selection for certain traits could result in the development of breeds and strains of sheep better
able to survive and thrive in such systems. For example, the ‘Marshall easy care’ Romney from New
Zealand. Development of this breed began in the mid 1930s and, at least initially, the ewes were not
shepherded at all in the hope that sheep would be selected which could look after themselves in difficult
terrain. The result is a strain with a reputedly high survival rate, excellent growth rate, good mobility,
good wool production and the ability to thrive in harsh conditions. In a study of maternal ability, this
‘easy care’ strain had a significantly higher maternal behaviour score than control Romneys, and
required less intervention by the shepherd. In a 5 year study, lamb survival in the Marshall Romney

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averaged 92%, compared with 88%, 86% and 73% in the Dorset-Romney, Perendale and Romney
respectively, a result which may be partly due to the smaller litter size of Marshall Romney ewes
(Kilgour and de Langen, 1980).

Another example is the Easycare breed developed by Iolo Owen in Wales. The aim was to produce a
breed of sheep which would require minimal shepherding and veterinary care and yet offer good meat
yields and lambing ratios. This breed sheds its wool, reducing or eliminating labour costs associated
with shearing and dagging lambs prior to slaughter.

Starting in the 1960s, the breed was derived from the Nelson Welsh Mountain (Welsh Mountain x
Cheviot) which were crossed twice to the Wiltshire Horn, a wool shedding breed, and selected for wool
shedding, no horns and easy care traits. Typically they are not housed and are fed minimum amounts of
concentrates. Where there is no threat from sheep scab, no dipping is required as there are no incidences
of blowfly on non-soiled parts of the wool. Easycares carry a reasonable fleece of up to 1 - 2 inches in
length through the winter which they cast in the spring. The breed appears to have good conformation
and genuine easy care features. Lambs at birth appear to have sufficient wool, are easily born and
quickly gain their feet and start suckling. The breed has the potential to significantly reduce labour input
and is recommended where shearing is expensive and difficult to organise. Other hair breeds like the
Dorper could make good terminal sires or early breeding sheep (Vipond, 2006).

This report will consider the following groups of candidate traits that may confer better adaptation to
more extensive management systems:
• Resistance to disease (nematodes, footrot, mastitis, blowfly strike and others)
• Enhanced flock fertility (ewe fertility, litter size, ram sexual capacity)
• Improved maternal ability and lamb survival
o maternal traits (lambing ease, care of the newborn, milking ability)
o lamb traits (lamb survival, gestation length, parturition, neonate behaviours, cold
resistance)
• Tolerance to food shortages
• Longevity (including teeth and bone)
• Body composition
• Wool shedding
• Temperament

Each section will include discussions on reported heritability estimates for the various traits and other
relevant genetic parameters, where available.

The report will conclude with a summary of the different traits and their current and future potential for
inclusion in breeding programmes in the UK.

3 Breeding for resistance to disease

Animal health affects production economics, animal welfare, and food safety. Prevention of diseases
will usually be preferable to curing them, and in cases where there is genetic variation in resistance to a
disease, selection may be a useful preventative measure. This approach may be particularly useful in
extensive systems where the animals cannot be inspected as regularly as those in more intensive
systems.

Variation in disease resistance (or susceptibility) has been found between species, between breeds and
also between individuals within breeds. Much research into genetic resistance to disease in sheep is
focused on gastrointestinal parasites, footrot and blowfly strike because they impose severe economic
constraints on sheep grazing systems. In addition, the increasing resistance of the causative organisms
to commercially available chemical solutions, and public concern over chemical residues in meat,
brings into question the long term effectiveness of disease control through chemotherapy. A fresh
approach to control the incidence of mastitis in sheep is also required due to the importance of the
disease affecting sheep breeds in the UK, and the costly, ineffective treatments used to control it.

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3.1 Nematodes

Gastro-intestinal nematode infection threatens the health and welfare of livestock, compromises the
efficiency of production and is possibly the major disease challenge facing ruminants (Perry and
Randolph 1999), costing the UK sheep industry an estimated £84M per annum (Nieuwhof and Bishop,
2005).

The increasing problems of attaining effective control of nematode parasites through the use of
anthelmintics due to the evolution of drug resistance in parasite populations is well documented (e.g.
Jackson and Coop, 2000; Bartley et al., 2003), and this threatens sustainable sheep production
throughout the world. The decreasing efficacy of anthelmintics, coupled with the desire to move
towards production systems less reliant upon chemical interventions, has stimulated the search for
alternative sustainable control methods. The selection of sheep, particularly growing lambs, with
enhanced resistance to nematode parasites is often advocated as a control measure that may
complement other strategies and, in the longer term, lead to a reduction in the requirements for
anthelmintics.

Between- and within-breed genetic variation in resistance to nematodes has been demonstrated in many
countries and production environments. Well known examples include the Merino (e.g. Woolaston and
Piper, 1996; Woolaston and Windon, 2001), Romney (e.g. Morris et al., 1997 and 2000a) and Scottish
Blackface (Bishop et al., 1996; Stear et al., 1997) as well as feral Soay sheep (Smith et al., 1999).
Genetic differences between host animals in nematode parasite resistance have also been observed for a
variety of parasite species including Haemonchus contortus, Trichostrongylus colubriformis,
Teladorsagia circumcincta and various Nematodirus species. In the UK, the major gastro-intestinal
nematode of sheep is Teladorsagia circumcincta, and the mechanisms by which infections caused by
this parasite reduce growth have recently been reviewed by Stear et al. (2003).

Lambs are generally susceptible to infection until about 1 year of age and become increasingly less
susceptible as they grow older (Courtney et al., 1985; Gamble and Zajac, 1992; Kambara et al., 1993).
In contrast, adult ewes are relatively resistant to infection except during late pregnancy and lactation
(Courtney et al., 1984). Female lambs have been reported to be more resistant to infection and have
lower Faecal Egg Counts (FEC) than males after puberty, although there appears to be no differences
between sexes before puberty (Courtney et al., 1985; Barger, 1993; Woolaston and Piper, 1996).

If selective breeding for nematode resistance is to be implemented then it is necessary to be able to

quantify resistance. Faecal egg count, the number of eggs per gram of faeces, is the indicator trait
commonly used to assess the level of infection. FECs indicate the product of the adult nematode
numbers and the mean fecundity of the resident parasite population. Other potential indicator traits
include phenotypic physiological markers such as plasma IgA activity, pepsinogenaemia, fructosamine
concentrations in the plasma and eosinophilia. Potential genetic markers include the major
histocompatibility complex and the interferon gamma region.

3.1.1 Faecal egg count

Selective breeding using faecal egg count (FEC) as an indicator trait for animals that have greater
resistance to nematodes has been shown to be effective under New Zealand and Australian conditions
(Morris et al., 1997 and 2000a; Woolaston and Piper, 1996; Woolaston and Windon, 2001). In these
countries, the exploitation of host genetic variation in resistance using FEC in commercial sheep
breeding programmes is now well-established (e.g. the ‘WormFEC’ scheme in New Zealand and the
‘Nemesis’ scheme in Australia) leading to reduced dependency on drug usage. Studies have also
demonstrated that it would be feasible, in principle, to select sheep for resistance to gastrointestinal
nematode parasites under typical commercial sheep conditions in the UK where sheep face a natural
parasite challenge, using the same indicator trait (Bishop et al., 1996; Bishop et al., 2004).

Heritability estimates for FECs lie between 0.2 and 0.4 (reviewed in Safari and Fogarty, 2003; see also
Pollot and Greef, 2004a), i.e. moderately high and similar to most performance traits in lambs.
However, egg counts are generally a lot more variable than performance traits, giving considerable

7
opportunities for genetic progress if they are used as a measured trait in a breeding programme (Bishop
et al., 2004). Heritablities for Nematodirus egg counts tend to be slightly higher than corresponding
heritabilities for Strongyle egg counts (McEwan et al., 1992 and 1995; Bishop et al., 2004).

Research in the UK has shown that resistance to different species of nematodes tends to be related, with
genetic correlations between the FEC values arising from different species or genera of parasites
generally being close to 0.5 (e.g. Bishop et al., 2004). Studies in New Zealand, looking at the genetic
correlation of FEC for Trichostrongylus and Nematodirus spp. (Morris et al., 2004), found the
correlation to be approximately 0.43. This means that in practice, it might not be necessary for the
presence of all genera to benefit from selection strategies to improve host genetic resistance.

Vanimisetti et al (2004), in agreement with Courtney et al. (1986), found that selection for indicators of
parasite resistance in nonlactating ewes would have little correlated effect on resistance in lambs,
suggesting that response to infection with H. contortus is mediated by different mechanisms in lambs
and nonlactating ewes. Therefore they recommend that because the economic impact of parasitism is
much larger in lambs than in ewes (and because of the apparent lack of correlation between parasite
resistance in ewes and lambs) selection should focus on reducing parasite susceptibility in lambs.

Other studies indicate that in the periparturient ewe, FEC is a moderately heritable trait (Bishop and
Stear, 2001, Morris et al., 1998; Watson et al., 1995; Woolaston, 1992) and that selection of lambs for
increased resistance to parasitic infection also confers a degree of resistance in periparturient ewes
(Woolaston, 1992; Morris et al., 1998).

The direct effects of selection for reduced FEC on lamb growth and other performance traits appear to
vary according to production environment, the level and species of parasite challenge and the trait of
interest (Morris et al., 2000a; Bisset et al., 2001). For relationships between Stongyle egg counts and
lamb growth traits, published values from European studies have tended to be negative and strong, e.g. -
0.8 between egg counts and live weight in Scottish Blackface lambs (Bishop et al., 1996) and -0.6 for
the same traits in Polish lambs (Bouix et al., 1998). However, Bishop et al. (2004) reported a
correlation of only -0.13 between egg counts and live weight in UK Texel lambs. Under New Zealand
and Australian conditions, strong correlations are seldom seen, with values generally being between 0
and -0.3 (eg Bisset et al., 1992; Douch et al., 1995; Eady et al., 1998; Pollot et al., 2004), however
slightly positive, i.e. unfavourable, correlations have also been reported (e.g. McEwan et al., 1992 and
1995; Greef and Karlsson, 1998 and Morris et al., 2000a).

Correlations of Nematodirus egg counts with lamb growth are not well documented, although McEwan
et al (1992) reported genetic correlations that were more negative than the corresponding correlations
with Strongyle egg counts. In contrast, Bishop et al. (2004) found correlations of performance traits
with Nematodirus egg counts tended to be neutral or slightly positive.

Genotype xenvironment interaction is a term used to describe the phenomenon that occurs when a set of
genotypes change their relative performance in different environments (Falconer, 1981). Such
genotype x environment interactions potentially have implications for the efficacy of selection
strategies to improve host resistance. Pollot and Greeff (2004) working with Merinos in Australia
found that FEC had a high heritability at the extremes of the environmental range, but a moderate level
in the middle. This implies that in low FEC environments, the genetic variation for FEC was high and
therefore some rams have the genetic predisposition to have high egg counts even when the challenge is
relatively low (ie they have no resistance to the parasites at all). Under a moderate challenge regimen,
animals are more similar in their genetic control of FEC; presumably the animals with lower resistance
in good environments increase their FEC values towards the animals with no resistance. At high
challenges, some animals have the ability to resist the parasites more than others. The way heritability
varies across the range of trait environments may have some implications for selection programmes. For
example, selection schemes could be designed to select against the animals that have high FEC in poor
environments, as well as selecting those that have a low FEC in all environments.

Experimental results from New Zealand and France, along with theoretical considerations, show that
added benefit is obtained from genetically reducing FEC, and hence pasture contamination, as sheep

8
grazing this pasture subsequently face a lower level of challenge and hence perform better. These are
epidemiological effects that give greater apparent responses to selection than predicted by conventional
genetic theory, both in terms of reduced FEC (Bishop and Stear, 1997) and increased performance
(Bishop and Stear, 1999). Experimental verifications of these concepts are provided by Gruner et al.
(2002) and Leathwick et al. (2002). In both cases, the grazing of fields by ‘susceptible’ sheep led to
considerably heavier pasture contamination, increased FEC and reduced performance, than grazing by
‘resistant’ sheep. This means that direct benefits of selection will accrue from decreased anthelmintic
requirements of selected animals and indirect, flock-wide, benefits on health and performance will
follow from reduced pasture contamination and hence decreased larval challenge (Bishop and Stear,
2003).

The cost effectiveness of including parasite resistance in the breeding goal using a selection index
approach depends on the identification of a reliable and repeatable measure of resistance, the
relationships between resistance and other production traits, and the specificity of selection to resistance
in terms of other parasite species and non-parasite pathogens. The weighting attributed to parasite
resistance in the breeding goal will be dependent on the degree of parasitism in the production
environment and the cost-effectiveness of breeding for resistance relative to other control measures.
Independent studies looking at various selection index models indicate considerable value from
improving resistance to nematode infection, and that these benefits may be obtained without sacrificing
much direct gain in liveweight (Woolaston, 1994; Amer et al., 1999; Bishop et al., 2004). Some
benefit is predicted to come directly due to the moderate but favourable genetic correlation between live
weight and Stongyle egg counts, however, greater benefit may accrue from the indirect benefits of
reducing pasture larval contamination and reduced use of anthelmintics. Bishop et al. (2004) suggest
that extra benefit will be obtained from basing selection on both Strongyle and Nematodirus egg counts
because of the higher heritability of the latter and the strong genetic correlation between them.

3.1.2 Faecal consistency scores and dag scores

The accumulated faecal matter in the wool of the breech area is referred to as ‘dags’, which can be
subjectively scored. Dag scores (DS) and faecal consistency scores (FCS) have been investigated as
possible indirect indicator traits to breed for decreased worm-related diarrhoea or scouring (Greeff and
Karlsson, 1997; McEwan et al., 1997).

Dag score seems to be a moderately heritable trait (between 0.25 and 0.35; reviewed in Pollot et al.,
2004) with higher values of heritability at the yearling age than either weaning or hogget age. Many
studies have found dag score to be unfavourably correlated with FEC, suggesting that selection for
decreased FEC will result in an increase in DS (Watson et al., 1986, Baker et al., 1991,; Pollot et al.,
2004). However, others have reported a positive correlation between the two traits (Bisset et al., 1992,
1996; McEwan et al., 1992).

There are various reports of FCS having a low heritability (Bisset et al., 1994; Bisset et al., 1996; Pollot
et al., 2004), although a similar number have estimated higher values (Karlsson and Greeff, 1996;
McEwan et al., 1992). McEwan et al (1997) and Greeff and Karlsson (1997) found negative genetic
correlations between FCS and FEC. However, in a larger study using a flock unselected for FEC,
Greeff and Karlsson (1999) found a small positive genetic relationship of 0.1 between FCS and FEC
and a value of 0.23 was reported by McEwan et al. (1992). Pollot et al (2004) found the correlations
between FEC and FCS to be effectively zero.

Dag score and FCS have been found to be highly correlated in all reports indicating that these two traits
are controlled by the same genes and suggesting that FCS could potentially be an indicator trait for DS.
This could be useful given that wool is normally removed from around the breech area to make the
animals less susceptible to blowflies, a practice which removes the variation in dags between animals.

The genetic correlations between liveweight and DS and liveweight and FCS have been found to be
generally negative (Watson et al., 1986; McEwan et al., 1992; Bisset et al., 1996) with the exception of
Pollot et al. (2004) who reported positive genetic correlations between the traits.

9
Dag score and faecal consistency score are clearly not the same trait as faecal egg count. Thus, although
these traits are generally used as indicators of host resistance to parasites, their poor relationship with
faecal egg count implies that they would be ineffective to improve host resistance to parasites.
However, selection for these traits in their own right could be important to decrease the occurrence of
scouring in sheep, where it is a problem (Pollot et al., 2004).

3.1.3 Phenotypic and genetic markers for nematode resistance

The use of FEC as the only indicator of host susceptibility or resistance to nematodes is time-
consuming and costly. Traits other than FEC, particularly various peripheral blood measurements, may
be used to assess resistance to nematodes or host response to infection. If the blood sample taken is then
also used for further genetic and physiological screening (e.g. for resistance to scrapie, and potentially,
for resistance to footrot), then multiple testing of blood samples may well prove to be an economically
efficient route to assess genetic resistance to several diseases. Furthermore, for T. circumcincta
infections, FECs appear to be relatively insensitive to changes in infection intensity (Bishop and Stear,
2000), and additional indicator traits may aid the identification of resistant and susceptible sheep.

There are several phenotypic and genetic markers for nematode resistance in sheep naturally infected
with nematodes that could potentially assist responses to selection. The phenotypic physiological
markers include immunoglobulin A (IgA) activity (Strain et al. 2002), pepsinogenaemia (Stear et al.
1999), fructosamine concentrations in the plasma (Stear et al. 2001) and eosinophilia (Stear et al. 2002).
The genetic markers include the major histocompatibility complex (Schwaiger et al. 1995) (Stear, et al.,
2005) and the interferon gamma region (Coltman et al. 2001).

Phenotypic markers
Ideally, a useful phenotypic marker, in addition to being strongly correlated with nematode resistance,
would be easy to sample and its assay would be inexpensive and able to be automated. Potential
phenotypic markers are described below (Box 1).

Box 1: Potential phenotypic markers

• IgA is a secreted antibody which is part of the acquired immune response. It has a major role in gut
infections and appears to regulate worm fecundity (Smith et al., 1985; Stear et al., 1995).
• Pepsinogen is a precursor of the digestive enzyme pepsin. An increase in pepsinogen activity
therefore indicates the presence of parasites and resultant damage to the gut of the host animal
(McKellar et al., 1986; Fox et al., 1989).
• Fructosamine concentration reflects average glucose and protein concentrations as well as the rate of
protein turnover. T circumcincta can cause a relative protein deficiency as well as an increase in
protein turn-over. Heath and Connan (1991) observed a decrease in fructosamine concentration
following deliberate gastro-intestinal infection, while Stear et al. (2001) reported that naturally
infected animals with low fructosamine concentrations subsequently acquired more nematodes of all
species and had shorter, less fecund, female T. circumcincta.
• Eosinophils are a type of white blood cell and are part of the immune response. Changes in
eosinophil counts have been associated with resistance to parasitic infection, and they may interact
with IgA to regulate nematode growth (Stevenson et al., 1994; Doligalska et al., 1999; Stear et al.,
2002).

Taken from Davies et al., 2005

As a first step towards developing useful phenotypic markers, various studies have looked directly at
the genetic control of parasitic traits describing the properties of the infection in lambs. The parasite
development traits - mean worm length and mean number of eggs in utero in adult female worms -
appear to be considerably more heritable than the numbers of larvae or adult worms present in the gut
(Table 1; Stear et al., 1997). These results were confirmed by Davies et al. (2005). In contrast, Gauly
et al (2002) reported a heritability value of 0.54 for worm burden yet found worm length to be not
heritable, in a small study of Rhoen sheep deliberately infected with Haemonchus contortus.

10
Table 1 Necropsy trait heritabilities
Trait h2 s.e.
Worm length 0.53 0.17
Worm burden 0.13 0.10
No. eggs in utero 0.50 0.16
No. of adult females 0.08 0.09
No. of adult males 0.12 0.10
No. of L4 larvae 0.06 0.09
No. of L5 larvae 0.12 0.09
Total no. of worms 0.12 0.10
Stear et al., 1997

Antibody responses to helminth infection have been investigated on a number of occasions. For
example, Douch et al. (1995) reported heritabilities of 0.18 for immunoglobulin G (IgG) specific to T.
colubriformis and 0.31 for antibodies specific to T. circumcincta. However, Gauly et al. (2002) found
IgG activity not to be heritable. Davies et al (2005), working with Scottish Blackface sheep, reported
strong and consistent heritabilities for IgA specific to T. circumcincta (0.57 at a mean age of 22 weeks).

Davies et al (2005) also observed strong negative genetic correlations between both IgA activity and
eosinophil counts and the worm development traits. This suggests that families with high levels of IgA
or eosinophil counts will have shorter, less fecund worms. This result is in agreement with previous
work that associated IgA activity with the regulation of worm fecundity (Smith et al., 1985; Stear et al.,
1995). Thus IgA activity and eosinophil count may be useful traits for selection purposes, with the aim
being to increase values of these traits in order to decrease worm development and fecundity.
Confirming this hypothesis further, FEC was found to be positively genetically correlated with worm
burden, and negatively correlated with eosinophil and IgA activity (Davies et al, 2005).

Genetic correlations between worm development traits and fructosamine were positive and moderate to
strong (Davies et al., 2005). This indicates that families with high fructosamine concentration have
long, fecund worms and therefore it may be possible to use fructosamine concentration to indicate
infection status. In general, the indicator traits discussed above are moderately to strongly heritable
and favourably genetically correlated with both FEC and worm size and fecundity, suggesting that they
do indeed reflect the animal’s ability to respond to infection (Davies et al., 2005). By using these
indicator traits within a selection scheme it would potentially be possible to influence worm
development traits to a greater extent than by using FEC alone. However, selection on these traits
would need to be implemented as part of a structured breeding programme and many factors, including
the cost and logistical implications, would need to be investigated first.

Genetic markers
Studies to detect quantitative trait loci (QTL) for resistance or detect associations with candidates are
now well advanced in New Zealand, Australia, Kenya, US and Europe - including the UK, France, Italy
and Spain - although results are not always readily available in the public domain. Full or partial
genome scans have revealed QTL for FEC on chromosome 1 (for T. colubriformis (Beh et al., 2002,
Diez Tascon et al, 2005) and H. contortus (Cockett et al, 2005)), chromosome 2 (for Nematodirus
(Davies et al., 2006)), chromosome 3 (for T. colubriformis (Beh et al., 2002), general Strongyles
challenge (Paterson et al, 1999, Davies et al., 2005) and Nematodirus (Davies et al., 2006)),
chromosome 6 (for T. colubriformis (Beh et al., 2002)) and H. contortus (Cockett et al., 2005)),
chromosome 14 (Nematodirus (Davies et al., 2006)), chromosome 19 (H. contortus (Cockett et al,
2005)) and chromosome 20 (general Strongyles challenge (Davies et al., 2006)). Further, Marshall et al.
(2005) report several significant QTL for H. contortus FEC, in sheep from the ‘Golden Ram’ flock
within which a major gene for resistance is believed to be segregating. The most consistently
significant region is that containing the interferon gamma locus on chromosome 3, although current
evidence suggests that the causative mutation is not within the interferon gamma gene.

QTL for IgA produced in response to Strongyles challenge have been reported by Davies et al. (2006)
on chromosomes 3, in the interferon gamma region, and 20, in the MHC region. The study by Cockett

11
et al (2005) also found a QTL for packed red blood cell volume (PCV) after H. contortus challenge in
the same location on chromosome 1 as the FEC QTL.

Several studies have also looked at associations between specific genes or markers and FEC. In
particular, Coltman et al. (2001) found significant associations with a microsatellite within the
interferon gamma gene (IFNG) in feral sheep, and various associations with microsatellites in or near
the major histocompatability complex (MHC) have been observed (Schwaiger et al., 1995, Janssen et
al., 2002). In addition, research in New Zealand has also implicated some QTLs for dagginess (J.
McEwan, pers. Comm).

3.2 Footrot

Footrot of sheep is an infectious and highly contagious disease, easily transmitted from sheep to sheep
via pasture, bedding or handling pens, and even spread by sheep that show no clinical signs of disease.
The disease results from invasion of epidermal tissue of the hooves by a mixed group of bacteria
(Egerton et al., 1969; Roberts and Egerton, 1969), the essential component of this mixture being
Dichelobacter nodosus, formerly Bacteroides nodosus (Dewhirst et al., 1990). Footrot-affected sheep
frequently experience pain, discomfort and lameness that affects their ability to compete for feed
(Abbot and Lewis, 2005). Affected sheep are also more susceptible to other diseases because of their
weakened condition.

Footrot is widespread and is the major cause of lameness in sheep in the UK (Grogono-Thomas and
Johnston, 1997), costing the UK sheep industry an estimated £24M per annum (Nieuwhof and Bishop,
2005). In a recent survey of farmers’ practices and attitudes towards footrot (Wassink and Green, 2001;
Hosie, 2003), more than 90% of sheep farmers had seen footrot in their sheep in the past year and 31%
considered that 6% or more of their flock were affected with footrot.

Control of footrot has largely focused on elimination of virulent isolates of D. nodosus from flocks,
prophylactic control of hosts through vaccination, and therapy of affected sheep through the use of
antibiotics and chemicals such as zinc oxide and formalin. None of these measures offers a long-term
easy-care approach to sheep management as they are expensive, not fully effective and will become
even less effective if the causative organism becomes resistant to chemotherapy.

It is likely that successful selection of sheep with improved genetic resistance to footrot through the
incorporation of footrot resistance into structured breeding programmes would be cost-effective,
reducing the current dependency on chemical solutions to control the disease, and would contribute to
increased sustainability by improving animal health, welfare and productivity.

3.2.1 Genetic variation

Breed differences in susceptibility to footrot have long been recognised and confirmed when animals of
different commercial breeds were uniformly exposed to infection by experimental and field challenge
(Skerman et al., 1982; Emery et al., 1984; Stewart et al., 1985; Cumlivski, 1988; Lewis et al., 1988).

There are also strong indications that within breeds, contradisposition to footrot infection is responsive
to selection. In the USA, Parker et al. (1983) reported significantly improved footrot resistance in
progeny of Targhee sheep which had failed to succumb to preliminary artificial challenge. In New
Zealand, similar advantages have been demonstrated in the Broomfield strain of Corridale sheep
developed from animals that have endured intensive field exposure to the disease (Skerman and
Moorhouse, 1987).

Estimates for the heritability of footrot liability vary. From an investigation of straightbred Romney
and Perendale ewes and their Booroola Merino crosses, Baker et al. (1986) reported estimates of 0.34
for liability to footrot and 0.03 for liability to severe footrot. Heritabilites of 0.53 and 0.23 respectively
were reported by Alwan (1983) for the same traits in Perendale sheep, and for the Romney breed,
Skerman et al (1988) reported heritabilities of 0.28 and 0.17. In a genetic study following challenge
and subsequent vaccination, Raadsma et al (1994), working with Merino sheep, found heritability

12
estimates of liability to footrot to range between 0.09 and 0.41, depending on the time after challenge
when the inspections were made. Half-sib heritability estimates of resistance to footrot were low to
moderate for single observations recorded pre-vaccination (0.07-0.22) and slightly lower for inspections
made after vaccination (0.07-0.15).

3.2.2 Footrot lesion scoring

Using Merino sheep in Australia, a footrot lesion scoring method has been developed, with severity
scored on a scale of 0 to 4 (Egerton and Roberts, 1971). This was further developed with sub-classes
that separated clinical signs into 8 categories (appendix 8.1; Raadsma et al., 1994). Successful breeding
for enhanced footrot resistance in Merinos has been described using this approach (Patterson and
Patterson, 1989), and Skerman (1985) and Skerman and Moorhouse (1987) have reported the
development of lines resistant to footrot in both the Romney Marsh and Corridale breeds using similar
scoring protocols.

3.2.3 Use of molecular genetic markers

Although selection using footrot lesion scoring has been shown to be successful, the development and
use of a molecular genetic test for footrot resistance potentially has enormous advantages. This is
because of the practical difficulty of objectively scoring feet lesions well, and the difficulty of
classifying them objectively and repeatably. In addition, with genetic markers, animals that are
candidates for selection do not have to be exposed to infection to determine whether they are
genetically susceptible or not. This method can also shorten the generation interval.

A group of genes important for controlling immune response lie within the Major Histocompatibility
Complex (MHC). These genes show great allelic diversity between individuals and it is thought that
some alleles are more efficient at initiating an immune response to specific pathogens than others. Data
exist, which suggest an association between genetic polymorphisms within the MHC Class II region
and response to footrot infection (Litchfield et al., 1993; Escayg et al., 1997). Indeed, genetic variation
within the ovine MHC loci in the class II region, specifically at the DQA2 gene, has subsequently been
used by Lincoln University, New Zealand to develop genetic markers for footrot resistance (Hickford et
al., 2004). Importantly, no links were found between the variation of D. nodosus strain (of which there
are at least 9 serogroups covering 18 subsidiary serotypes; Claxton, 1989) and resistance to footrot, so
selection of resistant sheep would not be compromised by inter-strain variation.

A footrot test for the New Zealand (NZ) sheep industry, based on their DQA2 alleles, is now
commercially available (Hickford, 2000) to select more tolerant or resistant animals, without having to
expose the animals to infection. Even though for IP reasons the data associating DQA2 alleles with
footrot have not been published, to date, more than 28,000 sheep have been genotyped in New Zealand
for 258 ram breeder clients. It is estimated that over 1 million sheep have already been born to rams that
have been screened using the NZ test, and the number of tests carried out have been growing by 30%
per annum. Recent survey results indicate that on properties that have adopted the technology, zinc
sulphate use has been reduced by 26% on Merino farms and 55% on Mid-Micron farms, formalin use
by 68% and 77% respectively, the number of doses of vaccine by 67 and 55%, and the numbers of
doses of antibiotics by 66 and 99%. Overall, footrot prevention and control costs have been reduced by
between 50 and 70% (Greer, 2005).

Current research underway in the UK aims to determine the ‘best’ way to breed footrot-resistant sheep,
which includes the further development of the NZ footrot gene test to include greater coverage of the
genome and it’s efficacy in UK sheep breeds.

3.3 Mastitis

Mastitis is considered to be one of the most important health problems in dairy cattle and sheep
(Heringstad et al., 2005; Leitner et al., 2004). Mastitis in sheep costs the UK industry millions of
pounds per annum in lost production and premature culling of affected animals. Conington et al. (2006)
predicted that the disease results in losses of approximately £11 per ewe.

13
Clinical mastitis can be defined as an inflammation of the mammary gland resulting from the
introduction and multiplication of pathogenic microorganisms. The main causative bacteria include
Staphylococcus aureus and Streptococcus agalactiae, (both of which are contagious) and coliforms,
streptococci and enterococci. All of these pathogens are found in animals’ environment (bedding,
manure and soil). These major pathogens can cause clinical mastitis, which can lead to swelling or pain
in the udder, changes in milk composition and appearance, increased rectal temperature, lethargy,
anorexia and even death.

Other, minor pathogens are also responsible for a rise in Somatic Cell Count (SCC). Sub-clinical
mastitis does not lead to visible changes in milk or the udder, although it is characterised by reduced
milk yield, altered milk composition and the presence of inflammatory components and bacteria in
milk. Both forms of mastitis can have serious economic consequences, due to loss of production and
premature culling of affected animals.

Most of the information available on mastitis in the literature relates to dairy cattle, with well-
documented evidence of the causative bacteria, prevalence and incidence of disease, economic costs
and control measures, including quantitative and molecular methods of breeding for resistance to
mastitis. Much less information on mastitis is reported for sheep, which in turn is dominated by the
dairy sheep sector in Mediterranean countries. There is almost no reported data on mastitis in meat
sheep. However, a full report on the possibilities of including mastitis in meat sheep breeding
programmes is the subject of a previous SPARK project by the same author (J.Conington) and is the
subject of a review paper (in press).

Due to the importance of mastitis affecting sheep, and the costly, ineffective treatments used to control
it, a fresh approach is required, potentially through the use of selective breeding for mastitis resistance.
Selective breeding for mastitis resistance requires at least a suitable selection trait or molecular genetic
marker, and that the additive genetic variance for this trait is of sufficient magnitude. In addition,
effective breeding requires knowledge of its genetic relationship with other traits of economic
importance.

3.3.1 Milk Somatic Cell Counts

Cattle
Using breeding as a measure to combat mastitis in dairy cattle has been the subject of considerable
research effort over the past 20 years or so. In the last decade, genetic evaluations for somatic cell count
(SCC) in dairy cattle have become available in most countries. Selecting for SCC rather than for clinical
mastitis (CM) has several advantages: SCC is routinely recorded in most dairy cattle recording systems,
contrary to CM, and SCC has a higher heritability than CM (0.15 vs. 0.02; reported in Barillet et al.,
2001). As SCC is genetically correlated with CM (a correlation of around 0.7), selection for reduced
SCC will lower the incidence of mastitis.

Many studies have shown that unfavourable genetic correlations exist between resistance to mastitis and
milk production traits in cattle. Beilharz et al.(1993) and subsequently Rauw et al. (1998) and Bakken
et al.(1998) used resource allocation theory to explain the negative genetic correlations that many, but
not all, researchers have observed between performance traits and those important for fitness and
health. Several countries have included mastitis into breeding programmes alongside conventional traits
to stem the deterioration in genetic susceptibility as a result of selection for increased productivity.
Mark et al. (2002) list 12 such countries. Veerkamp et al. (1998) estimated that selection to decrease
mastitis or limit the rate of increase of mastitis, with a cumulative impact of 1% per year and a national
penetration of 50% would result in a national benefit of £0.9m/year, and Conington et al. (2003)
reported that increases in mastitis incidence would be halted by the inclusion of SCC into dairy cow
genetic evaluations.

Sheep
Genetic literature is limited for dairy sheep and not always in agreement with dairy cattle results.
Intramammary infections in dairy sheep mainly differ from bovine infections by their etiology and by a
lower incidence of clinical mastitis versus subclinical mastitis (Emanuelson et al., 1988). In both

14
species, coagulase-negative staphylococci are usually the most frequently isolated germs in subclinical
mastitis. They are, however, considered minor pathogens in cattle whereas in dairy sheep they are
responsible for most cases of mastitis caught in milking parlours and consequently appear to be major
pathogens in this species (Barillet et al., 2001).

Selection for resistance to mastitis in sheep is currently underway in the French dairy sheep breed
Lacaune using SCC as a proxy trait for mastitis and incorporating it into sheep dairy breeding
programmes (Rupp et al., 2002). The Latxa Breeders’ Associations’ Confederation in Spain
(CONFELAC) also began recording milk composition traits including SCC in 2001 (Legarra and
Ugarte, 2005).

Genetic parameters
There are several heritability estimates for somatic cell count and other milk traits in dairy sheep (Table
2.1). The h2 values for SCC are relatively low and range from 0.04 to 0.24, depending on the breed and
stage of lactation when they were estimated, tending to increase with days in milk (Barillet et al., 2001).
Genetic correlations with other production traits are shown in Table 2.2.

From this table it is clear that there is no consensus among the literature estimates for the genetic
relationships between milk yield and SCC, unlike those for dairy cattle. For example, Legarra and
Ugarte (2005), working with Spanish Laxta dairy sheep, found that selection for milk yield would have
a negative effect on udder depth and teat placement (with implications for milking) but would decrease
SCC. This negative (favourable) genetic relationship between milk yield and SCC is in agreement with
estimates for other Spanish breeds and Greek Chios sheep (see table 2.2) but not with the positive
estimates for the French Lacaune breed (Barillet et al., 2001; Rupp et al., 2003).
Differences among reported genetic correlations may be related to models, level of production (Spanish
breeds show similar levels of production whereas Lacaune shows a higher level), or data collecting. It
may be that genetic improvement in sheep dairy breeds has not been implemented to the same degree as
that for dairy cattle and therefore any susceptibility among high-producing sheep to mastitis is not yet
apparent. It could also mean that there simply are no antagonisms between selection for increased milk
production and SCC in sheep. What these estimates do imply, is that any attempt to introduce milk
sampling as a way to detect mastitis in sheep will need robust estimates of genetic parameters in the
relevant population, in particular in relation to key production traits.

Table 2.1: Heritability estimates of milk production traits in sheep

Traits Heritability Population (breed) References

SCC 0.15 (0.04-0.12*)
Milk yield 0.34
French Lacaune dairy sheep Barillet et al., 2001
Milk fat 0.50
Milk protein 0.63
SCC 0.12
Milk yield 0.24 Spanish Churra ewes El Saied et al., 1999
Protein percentage 0.17
SCC 0.13 French Lacaune dairy sheep Rupp et al., 2003
SCC 0.11 Churra ewes Othmane et al., 2002
SCC (1st lactation) 0.12 Manchega ewes (Serrano et al., 2005)
SCC (2nd lactation) 0.19
SCC (3rd lactation) 0.24
SCC (all lactations) 0.04
SCC 0.14 Greek Chios dairy sheep Ligda et al., 2002
Milk yield 0.35
Milk fat percentage 0.21
Milk protein 0.31
percentage
SCC** 0.13 Spanish Laxta dairy sheep Legarra and Ugarte,
Milk yield 0.21 2005

15
 SCC 0.04 Spanish Churra ewes Baro et al., 1994
Milk yield 0.34
Milk protein % 0.13
* increasing with days in milk
** Lactational Somatic Cell Score

Table 2.2: Genetic correlation between mastitis (or SCC) and other production traits in sheep

Traits Genetic Population References

correlation
SCC-milk yield 0.15 French Lacaune Barillet et al., 2001
dairy sheep
SCC-milk yield -0.15 Spanish Churra
El Saied et al., 1999
SCC-protein % -0.47 ewes
SCC-milk yield 0.18
French Lacaune
SCC-fat content 0.04 Rupp et al., 2003
Dairy sheep
SCC-protein content 0.03
SCC-milk yield (ml/d) -0.36
SCC-Fat (g/L) 0.04
SCC-Protein (g/L) 0.13
Churra ewes Othmane et al., 2002
SCC-Casein (g/L) 0.09
SCC-Serum protein (g/L) 0.20
SCC-cheese yield (kg/100 L) 0.33
SCC-120d Milk Yield (1st lactation) -0.12 Manchega ewes Serrano et al., 2005)
SCC-120d Protein % (1st lactation) 0.23
SCC-120d %Dry Matter (1st lactation) -0,04
SCC-120d Milk Yield (2nd lactation) -0.14
SCC-120d %Protein (2nd lactation) 0.09
SCC-120d %Dry matter (2nd lactation) 0.02
SCC-120d Milk Yield (3rd lactation) -0.15
LSCS-120d %Protein (3rd lactation) 0.08
SCC-120d %Dry Matter (3rd lactation) -0.00
SCC-120d Milk Yield (1st, 2nd and 3rd -0.16
lactation)
SCC-120d %Protein (1st, 2nd and 3rd 0.22
lactation)
SCC-120d %Dry Matter (1st, 2nd and 3rd 0.04
lactation)
SCC – Milk yield -0.11 Greek Chios Ligda et al., 2002
SCC – Fat % -0.05 dairy sheep
SCC – Protein % 0.12
SCC* - Milk yield -0.30 Spanish Laxta Legarra and Ugarte,
dairy sheep 2005
SCC – Milk yield -0.37 Spanish Churra Baro et al., 1994
SCC – Protein % 0.37 ewes
* Lactational Somatic Cell Score

3.3.2 Molecular technologies to quantify mastitis resistance

Molecular genetic markers

As the continuous measurement of phenotypic indicator traits for mastitis resistance or susceptibility is
time and labour intensive, the use of suitable molecular genetic markers could be an attractive
proposition, particularly for meat sheep breeds as it does not require milk sampling. However, all the
literature published to date on molecular markers refers to dairy cattle, and only one (unpublished)
study is relevant to dairy sheep. A summary of the literature is presented in appendix 8.2.

16
Even though there are markers reported for 22/30 cattle chromosomes, the greatest number (6) are on
chromosome 23, where the genes responsible for the major histocompatibility complex are located.
These genes are responsible for the induction and regulation of immune response and are the focus for
several other studies related to disease resistance.

In a review on the genetics of mastitis in cattle (Rupp and Boichard, 2003) the immune mechanisms
underlying mastitis resistance pointed towards better functionality of neutrophils (white blood cells),
although the associations with them and clinical mastitis were not straightforward and require further
investigation. This review gives a comprehensive summary of the major markers and candidate genes
that have significant relationships with mastitis resistance or susceptibility. Interestingly, in one study,
the markers associated with resistance to clinical mastitis were not the same as those responsible for
low SCC indicating that the use of SCC data in QTL studies aimed at reducing the incidence of mastitis
should be carefully evaluated. This highlights the usefulness of markers for traits that are essentially
polygenic (i.e. expressed by many different genes) and it is notable that to date, no published reports
exist of using molecular techniques for resistance to mastitis in dairy breeding programmes, although
most such knowledge stems from cattle.

The hunt for molecular genetic markers in cattle and sheep

In cattle, many Quantitative Trait Loci (QTL) for clinical mastitis and SCC were found by using whole
genome scanning. For example, the QTL for clinical mastitis were localised to Chr. 9, 10, 11,18 and 25
(Holmberg & Andersson-Eklund, 2004b; Schulman et al., 2004). QTL for SCC were mapped to Chr. 3,
5, 7, 9, 11, 15, 18, 23, 26 and 29 (Boichard et al., 2003; Holmberg & Andersson-Eklund, 2004b;
Schrooten et al., 2000; Schulman et al., 2004; Kuhn et al., 2003; Ashwell et al., 2004), especially the
QTL on Chromosomes 11, 15 and 18 reached the genome-wise significance level. Hence, chromosomes
9, 11 and 18 should be the candidate chromosomes for mastitis resistance in cattle as QTL for clinical
mastitis and SCC are localised on the same chromosome. More markers should be selected on these
chromosomes to detect the fine location of QTL or genes of clinical mastitis and SCC until the real
gene that controls the susceptibility of mastitis is known so that it can be used inbreeding for mastitis
resistance.

The best route to identify potential molecular genetic markers in sheep would be to align the ovine
genome with bovine genome and find the homozygous chromosomes or regions that are similar (9, 11
and 18) by using comparative genome mapping. The next stage would be to select sufficient markers,
especially microsatellite markers, on the alignments with other species and detect the QTL for clinical
mastitis or SCC, which then could be used in marker-assisted selection (MAS). Whole genome
scanning also could also be used to detect QTL for clinical mastitis and SCC.

Detection of candidate genes

Detection of candidate genes for clinical mastitis or SCC is another way to improve the mastitis
resistance in sheep. BoLA (bovine leucocyte antigen)-DBR3 gene of the bovine major
histocompatibility complex (MHC), IgG2 gene and CD18 gene were studied as the candidate genes of
clinical mastitis and SCC in cattle (Kelm et al., 1997). BoLA-DRB3.2*16 was significantly associated
with lower SCS in Holsteins (P< or =0.05) (Sharif et al., 1998). There was a significant association
between BoLA-DRB3.2*23 and occurrence of severe mastitis (P< or =0.05) (Sharif et al., 1998). To
date, the corresponding studies in sheep haven’t been reported. So, it is worthwhile to study the
association between these genes and clinical mastitis in sheep. Single nucleotide polymorphisms (SNPs)
can be detected by using PCR-RFLP or PCR-SSCP, and then the association between the genotype and
phenotype needs to be analysed to finally select the favourable genotypes for mastitis resistance and
their use in marker assisted selection.

17
Candidate genes for mastitis resistance in dairy cows

The most extensively studied genes having significant associations with different indicators of mastitis
are the MHC class II DRB3 alleles. The general consensus from the majority of the published literature
is that these genes may have potential usefulness as genetic markers of higher or lower risk of disease
occurrence in cows. Specifically, the DRB3.2*23 has been associated with severe mastitis from which
coliforms were the most commonly isolated bacteria (Sharif et al., 1998). The presence of allele
DRB3.2*16 was associated with a higher Estimated Breeding Value (EBV) for SCC, and allele
DRB3.2*8 was associated with increased EBV for clinical mastitis, as was the IgG2b allele and the
normal CD18 allele. Alleles DRB3.2*11, *23, IgG2a, and the recessive allele for bovine leukocyte
adhesion deficiency were associated with decreased clinical mastitis. A positive genetic association was
found between allele DRB3.2*24 and EBV for intra mammary infections (IMI) by major pathogens and
between DRB3.2*3 and IMI by minor pathogens. Several correlations between EBV for immunological
assays and EBV for mastitis measures were significantly different from 0. Cows with low EBV for SCS
tended to have neutrophils that had greater functional ability at maximal immunosuppression, low
serum IgG1, and high numbers of circulating mononuclear cells. Immunological parameters, including
physiological and molecular markers, are useful aids to understand the genetics of resistance to mastitis
(Kelm et al., 1997). It is also possible that the number of DQ genes that a cow actually has, and the ratio
of certain T-cell subsets (CD4:CD8) affects their susceptibility or resistance to mastitis. Susceptibility
to mastitis was associated with MHC haplotypes that have a single set of DQ genes, and animals with
CD4:CD8 ratio of 0.42 compared to 3.2 for mastitis resistant animals (Park et al., 2004).

Despite the evidence supporting the involvement of the DRB3 locus, there are inconsistent reports of
whether or not they confer increased or reduced resistance to mastitis. In a review by Rupp and
Boichard (2003), three authors showed significant association of allele DRB3.2*24 with susceptibility
to mastitis, more intra mammary infections with major pathogens, more clinical mastitis and higher
SCC. However, from other cited studies, allele DRB3.2*16 was associated with either higher or lower
cell count. Similar inconsistent trends were reported for DRB3.2*23 and DRB3.2*8. Several
explanations could be given to explain such trends. First, alleles may be related to resistance or
susceptibility according to environmental conditions (present pathogens), which may be different in the
five studied populations. More likely, the studied polymorphisms were not causal but linked to other
MHC loci involved in mastitis resistance, which would lead to different associations according to
families. Thus, analysis of effect of MHC haplotypes rather than single locus should be preferred to get
a better handle on the links between genotype and resistance to mastitis.

3.4 Blowfly strike

Blowfly strike is an important disease of sheep, causing the deaths of an estimated 12,000 animals each
year in the UK (Sargison, 2004). In a survey of organic farmers in Scotland, it was the only disease
specifically mentioned as a health problem in organic farming (Halliday et al., 1991).

Blowfly strike is caused by the invasion of living tissue by the larvae of dipteran flies, primarily the
sheep blowfly Lucilia sericata (MacLeod, 1992; Morris and Titchener, 1997). The maggots feed
directly on the skin of the infested sheep, creating serious welfare and economic problems. Affected
animals are restless, dull and reluctant to graze, and kick at the struck area. Secondary bacterial
infection often occurs and the animal may die of septicaemia or the absorption of toxins from liquefied
body proteins.

Clipped sheep and young lambs with short fleeces are not usually attacked, but as the length of the
fleece increases so does the risk of strike (French et al., 1996). The struck area is usually soiled or
damaged in some way to attract flies. The breech is the most commonly infested area. Soiling with
faeces or mycotic infection of the fleece, as a result of high humidity, can trigger an attack (French et
al., 1995). Clipping wounds, footrot lesions and headfly lesions may also become infested.

Effective prevention of flystrike remains problematic. Blowflies can travel for several miles, so unlike
lice and scab mites, they can not be eradicated from a farm. Furthermore, while modern insecticides are

18
extremely effective, in practice correct application of these drugs to achieve satisfactory residual
activity is difficult. There is also evidence for blowfly resistance to insecticidal dips in New Zealand
(Litherland et al., 1992) which is probably also the case in other countries.

Differences exist both between and within breeds in susceptibility to fly strike. Litherland et al. (1992)
reported a fly strike rate of 0.33, 0.10, 0.10 and 0.00 in Merinos, Romneys, Wiltshires and feral sheep
respectively. These differences appear to be mainly due to the short wool and different fleece structure
associated with summer moulting in the feral and Wiltshire sheep presenting an environment not
conducive to strike (see also section 5.4 of this report on selection for wool shedding). The feral and
Wiltshire sheep had the lowest dag scores over the treatment period. Blowfly strike incidence was far
higher in Merinos than Wiltshire Horn - Merino crosses, but these crosses were inferior to the pure bred
Merinos in terms of wool traits (Rathie, 1994).

There exists considerable evidence for large differences between Merino flocks in their susceptibility to
flystrike (Atkins and McGuirk, 1979, Dunlop and Hayman, 1958; Raadsma et al., 1989). Atkins and
McGuirk (1979) presented two heritability estimates for the prevalence of flystrike of 0.25 and 0.53.
Gilmour and Raadsma (1986) reported an estimate of 0.37 and Raadsma (1991) reported estimates of
0.58, 0.53 and 0.10 from three small data sets in which the prevalence of blowfly strike was very high.
These data support the view that the predisposition of sheep to flystrike is in part genetically
determined, with potential scope for within-flock genetic improvement, at least within the Merino
breed.

An alternative approach to the problem is to introduce new genetic material into the population in an
endeavour to make sheep less susceptible to blowfly strike. Of interest in this respect is the Wiltshire
Horn breed in which fleece wool is shed annually (Slee and Carter, 1961, 1962; Ryder, 1969).

4 Breeding for enhanced flock fertility

Number of lambs weaned per breeding ewe has a greater influence on productivity and profitability of
most sheep enterprises than any other trait (Matos et al., 1992). Net reproductive rate is determined by
several components, with fertility (number of ewes lambing per ewe joined), prolificacy (litter size;
number of lambs born per ewe lambing) and lamb survival (see section 4) having the greatest influence
(Wang and Dickerson, 1991).

In the male, testicular size has long been considered one of the most likely criterion from the
physiological, genetic and practical perspective to improve reproductive performance of related females
(Land, 1973; Bindon and Piper, 1976; Walkley and Smith, 1980). The value of testicular size as an
indirect selection criterion for improvement of female reproduction is dependent on the heritability of
testicular size and the genetic correlation between testicular size and female reproductive traits.

Sexual performance of rams is highly variable (Terrill, 1937; Price, 1987) and strongly influences flock
fertility (Matos and Thomas, 1992). Rams that exhibit relatively rapid ejaculation rates are capable of
inseminating a greater number of ewes per unit of time than males with poorer libido or mating
technique (Kilgour and Whale, 1980; Perkins et al., 1992). Tests for ranking rams on sexual
performance are repeatable (Snowder et al., 2002) and reliable predictors of sexual performance under
field conditions (Ibarra et al., 2000), potentially allowing breeders to evaluate the mating competence of
individual males before they are employed in a breeding programme.

In the U.S., number of lambs born in a given interval is the only reproductive trait currently genetically
evaluated. However, other reproductive trait measures are being studied, particularly those dealing with
accelerated lambing or lambing more than once per year, for possible incorporation at a later time. In
Australia and New Zealand Estimated Breeding Values are produced for litter size born, and Sheep
Genetics Australia currently offers Lambplan and Merinoselect customers Australian Sheep Breeding
Values for scrotal circumference at post-weaning, yearling and hogget ages.

19
4.1 Ewe fertility

Fertility does not usually receive direct emphasis in selection programmes, since it is subject to
continuous natural selection, and reported heritability estimates average less than 0.10 (Fogarty, 1995;
Safari and Fogarty, 2003). However, heritability estimates are on average positive, and the importance
of the trait indicates it merits attention, at least in flocks/environments where mean performance is low
(less than 90%; Bradford, 2002).

Ewes dry in any year can be culled, or at least not used to produce replacements, and rams should be
consistently selected from dams that have lambed every year.

4.2 Litter size

There is an abundance of evidence for differences between and within sheep breeds in prolificacy (eg
Turner et al., 1962; Owen, 1971; Fahmy, 1996) which is mostly attributable to ovulation rate
(Hanrahan, 1980). Mean values range from about 1.5 to 4.0, excluding the involvement of major genes.
Hanrahan (2002) reported that divergent selection for ovulation rate in the Finn breed resulted in the
High line (selected for high ovulation rate) having a mean ovulation rate 2.2 times that of the Low line
(selected for low ovulation rate) confirming the availability of a considerable amount of genetic
variation. Analysis of the variation within the lines did not yield any evidence that a gene with large
effect was involved. It is likely that a similar degree of genetic variation is available for exploitation in
any population of sheep but the rate of selection response will depend on the actual mean value.

The heritability of ovulation rate is generally greater than that for litter size as a result of the influence
of embryo wastage (Hanrahan, 1982). Weighted mean heritability estimates based on numerous studies
for number of lambs born per ewe lambing (litter size) and ovulation rate are low at 0.13 and 0.15
respectively (Safari et al., 2005). A previous review (Fogarty, 1995) reported mean estimates for the
same traits of 0.10 and 0.21 respectively, although estimates of the heritability of ovulation rate for the
prolific Finn and Romanov breeds were considerably higher at 0.50 and 0.39 respectively.

In a recent review, Safari et al (2005) reported that the number of lambs born per ewe joined was highly
genetically correlated with both its components: litter size (0.89) and fertility (0.73). These correlations
were higher than those for number of lambs weaned per ewe joined with litter size (0.62) and fertility
(0.73). Ewe fertility was moderately genetically correlated with both ewe rearing ability (0.44) and
litter size (0.44). On the other hand, litter size had a small negative genetic correlation with ewe rearing
ability (-0.14).

Bradford (1985), in a review of previous literature, concluded that an annual increase of 1-2% in
average litter size could be achieved by selection on this trait. There is also the possibility of ‘jump
starting’ the process by an initial screening of exceptionally prolific ewes from a larger population to
establish the foundation flock (Clarke, 1972; Hanrahan, 1982; Sakul et al., 1999).

Waldron and Thomas (1992) estimated that adding information on ovulation rate would increase rate of
genetic change in litter size by 23% compared to use of litter size data only. An experienced operator
can measure ovulation rate rapidly and accurately by means of laparoscopy but the extra cost of
obtaining the information may be justified only in breeding flocks with an effective marketing
programme for improved breeding stock (Bradford, 2002).

However, an undesirable side effect of increased average litter size is an increase in the incidence of
triplet and higher order multiple births (Bradford, 1985). Mortality amongst multiples is usually higher
than amongst single-born lambs, for reasons including a lower body weight (Smith, 1977; Hinch et al.,
1983; Elving et al., 1986; Gama et al., 1991; Fogarty et al., 2000) and an increased risk of poor maternal
behaviour (Dwyer and Lawrence, 2005). Furthermore, multiple-bearing ewes (and their lambs) will be
at greater risk from nutritional deprivation than their single-bearing counterparts, both pre and post-
partum, unless management is tailored to the litter size of individual ewes. In a hill environment the
nutritional quality of the grazing may make it unlikely that the ewe will succeed in carrying more than
one lamb through pregnancy without some constraint on the viability of the lambs at birth (e.g. low

20
birthweight; Robinson et al., 1999). Her ability to produce sufficient milk to support adequate growth in
her lambs is also dependent on good nutrition (O’Doherty and Crosby, 1996; Bizelis et al., 2000). In an
extensive environment, therefore, a large litter size may be associated with impaired lamb survival, and
the lower litter size often seen in breeds traditionally managed in these environments may be
attributable to some degree of natural selection (Deag, 1996; Dwyer and Lawrence, 2005).

There may also be a genetic link between production traits and litter size. Selection for lean tissue in
Blackface and Coopworth breeds resulted in increased mean litter size in the lean lines compared to
divergently selected or control lines (Conington et al., 1998; Dwyer et al., 2001; McEwan et al., 2001).
A similar increase in litter size with selection for reduced backfat has been seen in mice (Armbrust and
Eisen, 1994). Ap Dewi et al. (2002) reported a small negative genetic correlation between litter size
and ultrasonic fat depth (-0.01) and a positive correlation between litter size and ultrasonic muscle
depth (0.35) in Welsh Mountain sheep. Genetic correlations between litter size and weaning, post-
weaning and adult weights are generally positive and moderate in magnitude (Safari et al., 2005).
Taken together, these results indicate that the increased litter size in intensively vs. extensively
managed animals may be a consequence of selection for production traits rather than (solely) a
consequence of extensive management. Confirming this further, a comparison of Scottish Blackface
(hill) and Suffolk (lowland) ewes following similar nutritional management systems showed that the
Suffolk ewes still had a higher average litter size than the Blackface ewes (Dwyer and Lawrence, 1998).

Single gene effects

The discovery that the high prolificacy of Boorola Merino sheep was due to a gene with a large effect
on litter size (Davies et al., 1982; Piper and Bindon, 1982) provided a new perspective on the genetics
of fecundity in sheep and this new paradigm quickly led to evidence for major genes for prolificacy in
other sheep populations (eg Cambridge (Hanrahan and Owen, 1985); Icelandic (Jonmundsson and
Adalsteinsson, 1985); Javanese (Bradford et al., 1986); Belclare (Hanrahan, 1991) and Romney (called
the Inverdale gene; Davies et al., 1991)). Studies of ovulation rate using molecular genetic techniques
in these and other breeds have since identified the major genes involved (see Hanrahan, 2003, for a full
description).

4.3 Scrotal circumference

Particularly since Land (1973) drew upon common hormonal mechanisms governing reproductive
development in males and females to postulate that selection to improve female reproductive traits
could be based upon measures of testis size and growth rate in related males, considerable research has
been directed at studying testicular traits. Several authors have indicated that males with larger testes
have either greater sperm production or higher daily sperm output (Cameron et al., 1984; Purvis et al.,
1984; Mukasa-Mugerwa and Ezaz, 1992). In the live ram, scrotal circumference is highly correlated
with, and is a reliable indicator of, testis weight (Notter et al., 1981), and testis weight correlates well
with daily sperm production (Lino, 1972). Selection of rams for greater potential to produce increased
numbers of spermatozoa on a daily basis will enhance flock pregnancy rate and lambing percentage
(Memon, 1983). Póti et al. (1999) reported a positive correlation between scrotal circumference and
libido of 0.56, but no significant correlation between scrotal circumference and sperm quality.
Mickelsen et al. (1981) suggested that there is no direct relationship between ram fertility and scrotal
circumference.

Scrotal circumference increases with age (Ruttle and Southward, 1988; Moore and Sanford, 1985) and
shows seasonal variation (Mickelsen et al., 1981; Moore and Sanford, 1985).

In contrast to low heritabilities for female reproductive traits, moderate heritabilities have been
summarised for scrotal circumference in sheep (means of 0.24 and 0.21 reported by Fogarty (1995) and
Safari and Fogarty (2003) respectively). Scrotal size has also been correlated with fertility, ovulation
rate, litter size, and age at puberty in several species. In beef cattle, strong relationships have been
reported between testis size (measured as scrotal circumference) at about 1 year of age and measures of
age at puberty and yearling fertility in females (Brinks et al., 1978; King et al., 1983; Toelle and

21
Robison, 1985; Smith et al., 1989). Also, divergent selection for cow fertility in tropical conditions led
to a significant positive correlated response in scrotal circumference between 9 and 18 months of age
(Mackinnon et al., 1990). After 22 generations of selection for increased litter size in mice, a realised
genetic correlation between testis weight and litter size of 0.42 was reported (Eisen and Johnson, 1981).

Results in sheep, summarised by Matos and Thomas (1992), are variable, but suggest a generally
positive relationship of testis size with ovulation rate and a somewhat smaller positive relationship with
realised prolificacy. Waldron and Thomas (1992) reported a positive genetic correlation between
scrotal circumference and ovulation rate of 0.20, but a correlation of -0.25 between scrotal
circumference and litter size. Haley et al (1990) reported that selection for testis diameter in crossbred
sheep did not change ovulation rate or litter size but increased the number of lambs born per ewe mated,
apparently through an increase in fertility. Duguma et al. (2002) found that rams with larger scrotal
circumference induced a significantly higher fertility and general productivity in their ewe mates, and
suggested that the use of rams with larger testis measurements would allow a reduction in the number of
rams required for breeding each year and increase the overall reproductive efficiency of the flock. Al-
Shorepy & Notter (1996) reported genetic correlations between scrotal circumference at 90 days (SC90)
and spring fertility and fall litter size of 0.29 and 0.36 respectively. Fossceco & Notter (1995) reported
a genetic correlation between SC90 and ewe fertility of 0.20.

Walkley and Smith (1980) indicated that greater genetic gain in selection for a ewe reproductive
measure could be achieved if direct selection for a ewe reproductive trait was complemented with
indirect selection based on a male trait with an approximate heritability of 0.35 and genetic correlation
with the female reproductive trait of >0.30. Scrotal circumference may meet these minimum criteria.

However, although various reports indicate that scrotal circumference is moderately to strongly
positively correlated with liveweight at different ages (Brash et al., 1994a,b; Fossceco and Notter, 1995;
Al-Shorepy and Notter, 1996; Duguma et al., 2002), others have reported a negative correlation
between the two traits (Land, 1982; Burfening and Davis, 1998; Haley et al., 1990) raising doubts about
the value of including information on scrotal circumference in a selection programme to improve litter
size.

If selection for testicular size is to be practised, scrotal circumference growth from 90 to 180d (with
some adjustment for liveweight) appears to be the trait of choice because it generally has the highest
heritability (Matos et al., 1992; Fossceco & Notter, 1995; Al-Shorepy & Notter, 1996). In addition, this
trait does not need to be adjusted for ram type of birth (single or multiple; Matos et al., 1992).

4.4 Enhanced ram sexual performance

Sexual performance is highly variable among populations of rams (Terrill, 1937; Price, 1987) which
means that a relatively rapid response to selection is very likely. Most mature rams readily court,
mount and mate oestrual ewes, whereas the intensity of sexual behaviour varies from asexuality to high
sexual activity. This fact has led to the development of procedures to rank rams for breeding soundness
based on their sexual behaviour (Wiggins et al., 1953; Blockey, 1976; Kilgour and Whale, 1980;
Perkins and Fitzgerald, 1992). Such tests, which assess the rate at which rams attain successful matings
when housed with oestrous females, have been shown to accurately predict the breeding performance of
high and low performance, female-oriented rams (Stellflug et al., 2006). These tests are highly
repeatable (Snowder et al., 2002) and are reliable predictors of sexual performance under field
conditions (Ibarra et al., 2000). They can also identify rams with high sexual performance at an early
age (14 months; Snowder et al., 2002).

High sexual performance rams (ie males that exhibit relatively rapid ejaculation rates) are capable of
servicing more ewes per unit of time and therefore produce more lambs than males with poorer libido
(Perkins et al., 1992; Stellflug et al., 2006). High-performing rams are also less likely than low-
performing rams to repeatedly mate with the same females (Price et al., 1996). This is an important
consideration when choosing rams for sheep breeding programmes in which the female:male ratio is
relatively high, as in many extensive range breeding situations.

22
Bench et al (2001) found that sire sexual performance classification influenced the proportion of high to
low performing ram lambs produced. 82% of high-performing ram lambs were sired by high
performing sires, whereas 60% of low performing ram lambs were sired by low performing sires.
Furthermore, the average number of ejaculations attained was significantly higher in sons born to high-
performing sires versus sons born to low performing sires. The greater breeding success of high
performance rams compared with low performance rams is probably related to their greater sexual
motivation (more mounts and ejaculations), rather than being attributable to general dominance (Erhard
et al., 1998) or differences in ability to efficiently perform the motor patterns associated with mounting
and copulation (Bench et al., 2001).

These results suggest that there is an important genetic component to sexual performance and libido in
sheep. Heritability estimates of 0.33 (Kilgour, 1985) and 0.22 (Snowder et al., 2002) for the serving
capacity of rams support this conclusion. Response to selection for serving capacity should therefore be
favourable, with selection resulting in rams capable of mating with more ewes, improving the
reproductive efficiency of the flock and reducing the number of rams needed.

Wilkins and Kilgour (1978) found that lambing success was significantly greater for the daughters of
high-performing rams but the number of lambs born to ewes that lambed was very similar for the two
groups, suggesting no difference in the ovulation rate. This result is supported by Bench et al. (2001)
who found no significant difference in ovulation rate or the number of days between first and second
behavioural oestrus periods in daughters of high and low performing sires. However, daughters of high
performing sires were significantly younger at first behavioural oestrus than daughters of low
performing sires. This is important because early maturing females may attain more oestrus cycles
before being bred, thus improving their conception rate as ewe lambs (Price, 1985). Breeding animals
early in life can shorten the generation interval for genetic selection and may increase the lifetime
number of offspring produced (Hulet et al., 1969). Snowder et al. (2004) found correlations of sexual
performance of rams with number of lambs born and weaned to be nil to low, depending on the breed.

5 Breeding for improved ewe maternal ability and lamb survival

Lamb mortality is a major constraint to efficient sheep production (Alexander, 1988; Haughey, 1991)
with the vast majority of lamb deaths occurring within 1-3 days of birth (Nowack et al, 2000; Southey
et al., 2004)1. The first critical stage is the birth process itself, dystocia representing one of the major
causes of mortality (Kerslake et al., 2005). After birth, survival of the newborn will depend largely
upon the quality of the interactions with the mother and her ability to provide the neonate with an
assured source of nutrition, protection and guidance (Haughey, 1993).

High lamb mortality could be reduced substantially by a short period of labour and by maternal
behaviours which protect the nutritional and thermal state of the lambs (Alexander, 1988; O’Connor
and Lawrence, 1992)2. This is particularly important in more extensive management systems where
ewes must be able to conceive, carry, give birth to and rear their young with little (or substantially
reduced) human intervention.

Selective breeding has been advocated as a means of improving lamb survival and ewe rearing ability
within breeds (Lindsay et al., 1990; Haughey, 1991). However, lamb survival to weaning has a low
heritability (0.02 – 0.13) suggesting that the scope for genetic selection to improve this trait is limited

1
A recent study (Southey et al., 2004) where specific causes of mortality were grouped into ‘dam-related’ (eg
dystocia and starvation), ‘pneumonia’, ‘other diseases’ and ‘other’ categories found that mortality in the dam-
related category was highest in the first week (3.2%), but it dropped below 0.3% in the second week and was
virtually non-existent after the forth week of age. In contrast, mortality in the pneumonia category was almost
constant for the whole of the 7 week study period, ranging between 0.5 and 0.8%. Overall, mortality dropped
from 9.8% in the first week to 1.7% in the second weeks and gradually decreased to 1% by weaning.
2
Maternal behavioural traits expressed at birth associated with lamb survival include: maternal licking and
grooming, low-pitched bleating, absence of aggression and lamb desertion, co-operation with lamb sucking
attempts, ewe selectivity and lamb recognition, and maintenance of close contact between ewe and lamb
(Reviewed in Dwyer and Lawrence, 2005).

23
(Hall et al., 1995; Lopez-Villalobos and Garrick., 1999; Morris et al., 2000b; Fadilli and Leroy, 2001;
Cloete et al., 2002) and that lamb survival is controlled mainly by non-genetic factors3. Nevertheless,
recent work has shown that when lamb mortality was analysed as component traits - ‘disease’, ‘dam
related’ (including dystocia and starvation), ‘pneumonia’ and ‘other’ - the heritability of the individual
traits was slightly higher (0.09, 0.16, 0.19 and 0,14 respectively; Southey et al., 2004) supporting the
suggestion of Cundiff et al (1982) that selection on components of mortality is likely to be more
effective than selection solely on mortality, regardless of cause. Mortality differences in sheep breeds
associated with different causes have also been reported by Nash et al. (1997), Mukasa-Mugerwa et al.
(2000) and Nguti et al. (2003), among others.

Based on a number of studies, the weighted mean heritability for ewe lamb-rearing ability (number of
lambs weaned/number of lambs born) was very low at 0.06 (Safari et al., 2005). However, Purser and
Young (1983) reported that lamb-rearing ability was a repeatable trait, and that the more lambs reared,
the better the subsequent performance appeared to be. If the lamb was not reared from the ewe’s first
parity, lamb mortality at age three and parity two was 26.8% but if the previous lamb was reared, the
mortality was only 13.5%. Morris et al (2000b) found that environmental variances due to permanent
maternal effects (for example uterine capacity, pelvic width, milking and maternal ability) were found
to contribute mostly to the repeatability of ewe lamb-rearing performance, providing evidence that
improved lamb survival has to be seen mainly as a successful partnership between mother and offspring
through pregnancy, parturition and lactation (Everett-Hincks et al., 2005).

Various authors (eg Alexander et al., 1990b; Cloete & Scholtz, 1998 and Kuchel & Lindsay, 1999) have
reported breed and line differences in lamb survival traits (eg length of parturition/ease of birth and
neonatal progress of lambs). Such differences may have arisen through relaxation of selection pressure
for a particular trait or traits, particularly if traits have a negative genetic correlation with production
traits. For example, hill and upland sheep breeds are more adapted to harsh environments and minimal
human intervention than more intensively managed lowland sheep breeds that have been selected and
managed for greater production of meat, wool or milk (reviewed in Dwyer and Lawrence, 2005).

It appears, however, that there are several candidate behaviours and traits, of both ewe and lamb, which
would confer a survival advantage on the new-born lamb, regardless of whether they are transmitted via
a genetic or experiential route. Identification of these adaptations is an important first step if breeds that
currently appear unsuitable for less intensive systems are ever to be managed under more extensive
systems (Dwyer and Lawrence, 2005).

Conington et al. (2001, 2002) predict that using multi-trait selection indexes, improvements in maternal
characteristics can be achieved alongside increasing lamb weaning and carcase weights, with little
change in subjective lamb carcase quality traits. Key potential index traits include lamb loss from birth
to weaning (including lambs born dead), litter size at weaning (own lambs only) and average weight of
lambs weaned (including weight of lamb fostered on). Including lamb survival as a trait of the lamb as
well as lamb losses in the index is predicted to reduce lamb ‘wastage’ between birth and weaning
(Conington et al., 2002).

In the U.S., evaluation of maternal performance is achieved by recording number of lambs born and
calculation of a ewe productivity index: pounds of lamb weaned per ewe per lambing. Ewe
productivity is calculated only for ewes that wean at least one lamb. Weights of weaned lambs are
adjusted for lamb age, lamb sex, and ewe age (but not for type of birth and rearing) and are summed to
derive ewe productivity. Maternal effects on lamb weaning weight are predicted in the current within-
flock analyses, but maternal weaning weight Expected Progeny Differences (EPD) are not currently
reported.

In Australia and New Zealand, EBVs are produced for number of lambs weaned.

24
5.1 Maternal ability traits

5.1.1 Ease of parturition

A prolonged labour increases the possibility of brain trauma and hypoxia in the neonate (Haughey,
1993) and impairs sucking, locomotor activity and thermoregulation in lambs (Haughey, 1980; Eales
and Small, 1980; Bellows and Lammoglia, 2000; Dwyer, 2003), increasing the chances of death when
subjected to cold stress or mismothering.

Differences in length of parturition between breeds (Alexander et al, 1990b; Fahmy et al., 1997; Cloete
et al 1998) and lines (Cloete and Scholtz, 1998) have been reported suggesting that there is some
genetic variation for this trait. However, heritability estimates were below 0.05 for both SA Mutton
Merino and Dormer breeds (Cloete et al., 2002) and 0.17 for Merino ewes with a small service sire
effect amounting to 0.03 (Cloete et al., 2003). This trait was, however, subject to significant maternal
effects (m2) - computed variance ratios amounting to 0.14 in Dormers and 0.15 in SA Mutton Merinos –
indicating that the inheritance of ease of parturition is likely to be mainly maternal (Cloete et al., 2002).

However, embryo transfer between Blackface and Suffolk ewes revealed that the lamb may not be
passive in the birth process but can actively contribute to its ease of delivery (Dwyer et al., 1996). Lamb
characteristics such as birth weight and sex both contribute to the incidence of birth difficulty (Smith,
1977; Dwyer, 2003). Likewise, singleton and triplet lambs experience more birth difficulty than twin
lambs (see section 4.2.3 below), an effect seen more strongly in Suffolk ewes than Blackfaces (Dwyer,
2003).

Lambe et al. (2006) reported no significant change in the incidence of ewes requiring assistance at
lambing as a result of selecting Scottish Blackface ewes using a breeding index designed to improve
both carcass and maternal traits (Conington et al., 2006), compared to a control line or to a line selected
by normal commercial (visual) methods. However, lambing difficulties were significantly affected by
ewe sire and lamb sire (suggesting a genetic component), year, ewe age (highest at 2-years-old), lamb
sex (male>female), still births (higher) and ewe pre-mating weight (higher weights = less assistance).

Further studies on genetic relationships between lambing difficulties and other traits would be useful to
help make future breeding decisions to improve productivity whilst maintaining high standards of
animal welfare in extensively managed flocks.

Pelvic dimensions
An important component of ease of delivery is the pelvic dimension of the ewe (McSporran and
Fielden, 1979; Haughey et al., 1985; Cloete et al., 1998; Bilbe et al., 2005). In Merino flocks selected
to improve the ratio of lambs weaned to lambs born (‘lamb-rearing ability’) the main outcome of the
selection process has been an increase in the speed and ease of parturition of the selected line over
control lines (Cloete and Scholtz, 1998; Cloete et al., 2003). Similar responses have been seen with
‘easy-care’ Romney ewes (Knight et al., 1988; Kilgour and de Langen, 1980). With both breeds,
selection for lamb-rearing ability is associated with an increase in ewe pelvic dimensions (Knight et al.,
1988; Kilgour and Haughey, 1993).

It is possible that assisting ewes at parturition may have reduced natural selection for ease of birth and
large pelvic dimensions in ewes of more intensively managed breeds. Comparisons of Blackface or
Romanov ewes with the Suffolk breed have shown that both Blackface and Romanov ewes tend to have
a quicker and easier birth process than Suffolks (Dwyer et al., 1996; Fahmy et al., 1997; Dwyer and
Lawrence, 1998; Dwyer, 2003). Comparative studies of Soays and Suffolks have demonstrated that the
relative pelvic dimensions in Soays (scaled for differences in body weight) were over twice as large as
Suffolks (Silva and Noakes, 1984).

Bilbe et al. (2005), using CT scanning to measure pelvic capacity in Scottish Blackface ewes, found
differences in mean pelvic dimensions of daughters from different sires indicating that genetic variation
exists within the breed.

25
Larger trials are necessary to quantify genetic properties of dystocia and pelvic measurements and their
relationship with other production traits before selective breeding for large pelvic capacity can be
recommended.

5.1.2 Care of the newborn

Fundamental to the survival of the lamb is the formation of a close and exclusive attachment or bond
between the ewe and her lambs to ensure their early sucking and colostrum intake (Nowak et al., 2000).
Specific behaviours of the ewe (licking and grooming, low-pitched bleating, absence of aggression and
lamb desertion, co-operation with lamb sucking behaviours) promote ewe–lamb recognition and a close
association between the ewe and lamb (Alexander, 1988; Nowak et al., 1997, 2000). Suppression of
olfactory cues is detrimental for maternal acceptance (Poindron and Lévy, 1990). Ewe behaviour
around lambing time has a large effect on lamb survival, particularly in extensive situations (Nowak,
1996) and may also affect weaning weight of lambs and thus ewe productivity (O’Connor et al., 1985).

Isolation
Isolation from the rest of the flock during and after parturition is an important preliminary step in the
formation of the mother-young bond as it protects the ewe from disturbances, prevents separation of
newborn lambs from their dams and reduces interference or lamb stealing by other ewes (Gonyou and
Stookey, 1983, 1985).

The amount of time that the ewe spends isolated from the flock, and her propensity to seek isolation, is
affected by breed and by parity. Whereas isolation seeking appears to be common in hill ewes (Hewson
and Wilson, 1979), less than 50% of Lacaune and Border Leicester ewes (Lécrivain and Janeau, 1987;
Alexander et al., 1990a) and less than 2% of Merinos (Stevens et al., 1981) are reported to prefer
isolation at lambing. Multiparous ewes tend to show greater isolation seeking behaviour than
primiparous ewes (Gonyou and Stookey, 1983; Alexander et al., 1990a), which may be related to
decreased fearfulness at separation from flockmates with the more experienced ewes (Viérin and
Bouissou, 2002). Characteristics of the birth site per se appear to be less important to lamb survival
than the ewe remaining undisturbed with her lambs for at least 6 h (Murphy et al., 1994).

In one study the period that ewes remained on or near the birth site was found to be heritable (h2 = 0.20;
Cloete et al., 2003). Szantar-Coddington (1994) found that fertility-flock Merino ewes remained on
their birth sites for 266 minutes compared to 251 minutes for control ewes. Knight et al., (1989) found
that Marshall Romney ewes (selected for rearing ability) grazed from their birth sites 46 minutes after
birth, while control Romneys did so after 27 minutes. Therefore, the general pattern from all these
studies was that ewes selected for lamb-rearing ability tended to remain longer on or near their birth
sites than control line contemporaries

Birth site selection

Wet, windy and cold weather are important factors in the deaths of lambs from hypothermia
(Alexander, 1962; Obst and Ellis, 1977; Alexander et al., 1980; McCutcheon et al., 1981), thus
selection by the ewe of an appropriate site in which to lamb may influence lamb survival. In studies
with Merino ewes, provision of shelter can reduce lamb mortality in poor weather by up to 50%
(Alexander and Lynch, 1976; Lynch et al., 1980; Alexander et al., 1980). Expression of appropriate
shelter-seeking behaviour would therefore be an important ewe behaviour that promotes lamb survival.

Adult ewes in full fleece seek shelter only when they are outside their own thermoneutral zone (Lynch
and Alexander, 1976; Alexander et al., 1979; Duncan et al., 2001). Shorn ewes make greater use of
shelter than full-fleeced ewes (Alexander and Lynch, 1976) as do breeds, such as the Lacaune, which
have relatively thin fleeces (Lécrivain and Janeau, 1987). However, neither Merino or Corriedale ewes
(both woolly breeds) seek shelter unless wind speeds exceed 32 km/h with rain, although mortality of
newborn lambs increases at wind speeds above 18 km/h (Obst and Ellis, 1977). Thus, ewes tend to seek
shelter for their own thermal comfort rather than to protect their newborn lambs.

26
Lactating ewes appear to seek shelter more frequently than at other times (Pollard et al., 1999).
However, this may reflect the use of shelter by lambs during inclement weather, and the desire of ewes
to remain with their lambs, rather than shelter-seeking behaviour of the ewe, particularly as shelter use
on the day of parturition was lower than during lactation (Pollard et al., 1999).

There are few breed comparisons of sheltering behaviour between intensive and extensively managed
sheep and, as described above, sheltering behaviour may be more influenced by ewe fleece
characteristics than other aspects of management. Other experiments suggest that use of shelter is
related to familiarity with the environment, which may again be a more important influence than breed
(Lynch et al., 1980).

There are, however, breed differences in selection of birth sites related to topographical and physical
features of the environment (Alexander et al., 1990a). On level pastures lambing sites for Merinos were
randomly distributed, but on pastures with slopes ewes preferred to lamb at elevated sites. By contrast,
both hill and upland (Welsh Mountain, Scottish Blackface, Cheviot) and lowland (Suffolk) ewes chose
lambing sites at the edges of the pasture rather than elevated sites (Alexander et al., 1990a).

Taken from Dwyer and Lawrence, 2005

Ewe-lamb recognition and bonding

The ewe forms a memory for her own lambs that allows her to restrict maternal care exclusively to her
own offspring (‘selectivity’) (Poindron et al., 1984a; Lévy et al., 1995). As ewes are selective for their
own offspring, a lamb that fails to form an attachment with its dam will not be cared for by any other
ewe and will not survive. Likewise, the offspring of a non-selective ewe will not thrive, as it is unlikely
that the ewe will produce sufficient milk to feed several lambs. The behaviours that promote selectivity
include grooming and ewe–lamb contact (Baldwin and Shillito, 1974; Poindron et al., 1980; Alexander
et al., 1986; Poindron et al., 1988; Lévy et al., 1991; Hernandez et al., 2001).

Experimental results suggest that lowland ewe breeds managed intensively tend to show poorer
maternal care at birth than hill breeds normally managed more extensively. For example, breed
comparisons of grooming behaviour have shown that, in French breeds, Romanov and Prealpes du Sud
ewes (hill breeds) spend more time licking their lambs than Ile-de- France (lowland meat breed) or
Lacaune (intensive dairy) ewes (Le Neindre et al., 1998; Poindron et al., 1984b). Likewise in the British
breeds, Blackface ewes show more grooming behaviour than Suffolk ewes and make more low-pitched
bleats immediately after birth (Dwyer and Lawrence, 1998, 1999a, 2000; Dwyer et al., 1998). Suffolk
ewes, particularly primiparous ewes, show more aggression towards their lambs, are more likely to
desert a lamb, and are less co-operative with the sucking attempts of their lambs than Blackface ewes
(Dwyer and Lawrence, 1998; Pickup and Dwyer, 2002). The differences between Blackface and
Suffolk ewes in the onset of maternal behaviour also appear to be related to maternal attraction to her
lambs in the immediate postpartum period. In tests where ewes are offered a choice between their own
and an alien lamb of similar appearance, Blackface ewes are quicker to approach and spend more time
with their lamb than Suffolks, (Pickup and Dwyer, 2002) although both breeds clearly recognised their
own lambs. Similarly, Dalesbred (hill) ewes are more attracted to their young lambs than are Jacob
ewes (Walser et al., 1983).

When managed as a single flock Blackface ewes maintain consistently closer spatial relationships with
their lambs than Suffolk ewes throughout lactation (Dwyer and Lawrence, 1999b, 2000; Pickup and
Dwyer, 2002). Blackface ewes are also more active than Suffolk ewes (standing frequency, Dwyer and
Lawrence, 2000) and Blackface ewes made significantly more vigilance postures (the ‘head-up’
posture, Pickup and Dwyer, 2002). Breed differences in lamb sucking behaviour also exist with Suffolk
ewes receiving a higher frequency of sucking attempts than Blackface ewes in the first few days after
birth (Dwyer and Lawrence, 2000). However, Suffolk ewes terminate a greater proportion of shorter
suckles than Blackface ewes (Dwyer and Lawrence, 2000), such that Blackface ewes receive a higher
proportion of successful sucks throughout lactation (Pickup, 2003).

27
These breed differences may have arisen through a relaxation of selection pressure on maternal
behaviour in intensive systems with human interventions (e.g. confinement of ewes and lambs in
individual pens for the first day of life) such that variation in maternal care has less influence on the
survival of the lamb. Breed differences persist in ewes reared under identical conditions, and sire effects
on maternal behaviour can be seen in cross-bred ewes (Pickup, 2003). This suggests that the effects on
maternal behaviour are primarily genetic, although nongenetic, experiential transmission of maternal
care (through maternal grooming, for example, as has been reported in rats, (Francis et al., 1999)
remains a possibility.

Maternal cooperation with suckling

The maternal environment to facilitate suckling, as provided by the dam, has been found to play a role
in the neonatal progress of lambs (Kuchel and Lindsay, 1999; Cloete and Scholtz, 1998; Cloete et al.,
2002) and has a small genetic component (heritability of 0.11; Cloete et al., 2003). For example, the
interval from standing to apparently suckling was shorter in a Merino line that was selected for multiple
rearing ability than in a parallel line that was divergently selected against multiple rearing ability. The
line difference was partially, but not wholly, accounted for by the inclusion of maternal co-operation
with the first suckling attempts of the neonate in the model of analysis (Cloete and Scholtz, 1998).
Ewes selected for rearing ability were less likely to circle and back than control line ewes and they were
more likely to adopt a posture to facilitate suckling.

5.1.3 Maternal behaviour score

One method of measuring maternal behaviour of sheep is to use a scoring system, developed by
O’Connor et al. (1985), based on the proximity of the ewe to her lamb as it is handled (appendix 8.3).
Maternal behaviour scores (MBS) are attributed to ewes based on their response to the handling of their
lambs at tagging, within 24 h of birth. A five-point scale is usually used, with higher scores awarded to
ewes which remain closer to their lambs as they are tagged in the field.

The score has been shown to be related to both postnatal lamb survival and weaning weight (O’Connor
et al., 1985; O’Connor, 1996; Lambe et al., 2001; Everett-Hinks et al., 2005; Sawalha et al., 2006), and
varies with ewe genotype (O’Connor et al., 1985; Alexander et al., 1990b). In general, Merino ewes
show a higher level of lamb desertion at handling than Romney, Border Leicester or Perendale ewes,
and Cheviot ewes are intermediate. A modified version of the score used with Blackface and Suffolk
ewes suggested there were no breed differences at 24h old but that Blackface ewes approached closer to
their handled lambs at 72h than Suffolks (Dwyer and Lawrence, 1998).

O’Connor et al. (1985) found a significant effect of ewe age on MBS. For each year increase in ewe
age, MBS increased by 0.26 units, in ewes of 6 genotypes. Lambe et al. (2001) found MBS to increase
with parity with differences in MS between ewes of different ages following a less predictable pattern
indicating that maternal experience of the ewe is an important factor. This is in agreement with the
findings of Cloete et al (1998) and O’Connor and Lawrence (1992) working with Merino and Scottish
Blackface ewes respectively.

Estimates of heritability for MBS vary but are consistently low (0.13, Lambe et al., 2001; 0.09, Everett-
Hincks et al., 2005). Lambe et al (2001) estimated a moderate, positive genetic correlation (0.4)
between MBS and the average weight gained by lambs from birth to marking although the genetic
correlation between MBS and weight gained to weaning was close to zero (0.02).

It should be noted that the MBS is not just a measure of the care and attachment of the dam to her
newborns but is also a measure of temperament of the ewe and its reaction to the presence of humans
(Dwyer and Lawrence, 2005; Sawalha et al., 2006). The Romanov ewe, for example, is considered to
be a better mother in terms of her licking and grooming behaviour and attachment to the lamb in
comparison to the Lacaune (Le Neindre et al., 1998). However, Romanov ewes have a greater flight
from humans and stood further from their handled lambs than Lacaune (behaviours that would have
earned them a lesser Maternal Behaviour Score). These responses are considered to be due to the
greater emotivity of the Romanov and not to a poorer quality of maternal care. In studies where Merino
ewes were selected for temperament by measuring their responses to a variety of tests (such as

28
behaviour within an enclosed box, see section 5.5 of this report) the ‘calm’ ewes spent longer grooming
their lambs than ‘nervous’ ewes, and bleated more frequently to their lambs (Murphy et al., 1998).
Lamb mortality in these lines was also lower in the ‘calm’ ewes in comparison to the ‘nervous’ animals.
Ewes previously selected for their ability to rear lambs also show behavioural differences in an
approach avoidance test indicative of increased ‘calmness’ (Kilgour and Szantar-Coddington, 1995).

Everett-Hincks et al (2005), working with Coopworth sheep in New Zealand, suggest that if
management practices and environmental conditions are conducive to high levels of litter survival, then
the incorporation of the MBS into the animal selection programme is likely to be of little benefit as
genetic variation and consequently heritability for MBS and litter survival are small. However, if
management practices and environmental conditions have not been integrated in such a way to enhance
litter survival, then the MBS may have a place. However, a new measure of maternal behaviour as it
relates to lamb rearing success is needed for flocks with higher litter survival rates if litter survival is to
increase further.

5.1.4 Milk production

Increased milk production of range ewes is associated with increased lamb survival and growth (Burris
and Baugus, 1955; Boyazoglu and Treacher, 1978; Torres-Hernandez and Hohenboken, 1980). The
time period when milk production most significantly influences lamb growth occurs before 6 weeks
postpartum (Slen et al., 1963; Snowder and Glimp, 1991), as may be expected due to the natural
decrease of lamb dependence on milk and increased consumption of forage and/or creep feed.
Improving milking performance and lamb production of range ewes could result in significant
economic returns to most sheep production systems (Snowder et al., 2001b).

Poor milk production is often associated with short-term gestations in ewes less physiologically
prepared for lactation and that exhibit less mammary development than full-term gestating ewes
(Rattray et al., 1974). Heavier body weight has been associated with ewes of better condition and
higher milking capability (Peart, 1968; Gibb and Treacher, 1982) and the level of nutrition can also
affect ewe milk production (Treacher, 1983). Sakul and Boylan (1992), in a study that included many
sheep breeds, concluded that substantial variation for milk production exists among and within U.S
sheep breeds indicating that there may be some genetic control of this trait.

Milk score is a subjective measure of milk production of ewes that can be qualitatively determined by
palpating the ewe’s udder and observing her lambs’ fill within a few hours of lambing. Snowder et al
(2001a,b) working with four breeds - Columbia, Polypay, Ramboullet and Targhee - investigated the
usefulness of a subjective milk score as an alternative to directly quantifying milk yield of nursing
range ewes.

For all breeds, ewes with low milk scores gave birth to lambs with lighter birth weights than ewes with
average or high milk scores (Snowder et al., 2001a). Birth weights of lambs born to ewes with average
milk score were intermediate. The heaviest birth weights tended to be associated with ewes with high
milk scores. Percentage of live-born lambs reflected the same trend as birth weight and was clearly
associated with milk score. These data suggest that that a contributing cause to higher mortality rates
among lambs with lighter birth weights may be an association with the dam’s low milking performance.

Support for the economic importance of milk score was the strong association between ewe milk score
and individual lamb weaning weight within each ewe age group. In all ages and breeds, ewes with high
milk scores reared heavier lambs than did ewes with low milk scores. With increasing lactations from 1
to 3 years of age there was an increase in individual lamb weaning weights within all milk score
classifications.

Snowder et al (2001b) reported that observed heritability estimates for milk score at first parity were
moderate and similar across breeds, ranging from 0.18 to 0.32. Heritability estimates adjusted for a
binomial distribution of milk scores at first parity were high (Colombia, 0.43; Polypay, 0.35;
Rambouillet, 0.50; Targhee, 0.84). Estimates of observed heritability for second-parity milk score were
moderate to high, ranging from 0.23 to 0.46. Estimates of heritability for lifetime records for milk score

29
ranged from 0.16 to 0.26 across breeds. Sawalha et al (2005), working with the same four breeds, found
estimates for heritability for milk score to be in the range of 0.05 to 0.18 for first, 0.01 to 0.27 for
second, 0.05 to 0.10 for mature and 0.08 to 0.13 for all lifetime parity groups. Legarra and Ugarte
(2001) reported an estimate of heritability of 0.20 for Laxta dairy sheep using 120d repeated lactation
records. Baro et al. (1994) and El Saied et al. (1999) estimated heritability to be 0.34 and 0.18 for
multiple-lactation, test-day milk yield of Spanish Churra milking sheep, respectively.

Snowder et al (2001b) found that milk score and litter weight weaned were genetically correlated at first
or second parity in Rambouillet (1.0) and Targhee breeds (1.0 and 0.61 respectively), but not in the
Colombia and Polypay breeds. Estimates of genetic correlations of annual lifetime milk score records
with litter weight weaned were high (Colombia, 1.0; Polypay, 0.81, Rambouillet, 1.0; and Targhee,
0.77). Sawalha et al (2005) reported genetic correlations at first parity between milk score and litter
weight to be high (1.00) for Rambouillet and Polypay, and near zero for Colombia and Targhee. At
second parity, estimates were positive and moderate for Rambouillet and Polypay but more variable for
Colombia and Targhee. Although estimates are variable, the average of the estimates of the genetic
correlation suggests that litter weight, at least in some breeds, could be improved by selecting ewes for
favourable milk score. Selection for lifetime production for litter weight weaned in Targhee ewes
increased milk yield of a 112-d lactation period by 13% compared to randomly bred control ewes (Head
et al., 1995).

Milk score at first or second parity was genetically correlated with milk score records at maturity (third
parity and greater) with estimates ranging from 0.69 to 1.00 (Snowder et al., 2001b; Sawalha et al.,
2005), suggesting that additive genetic value for milking ability at maturity could be evaluated as early
as at first parity.

Milk production of a ewe has an important effect on the preweaning survival and growth of her lambs.
Experimental results suggest that selection for milk score at early parities or based on lifetime records
would result in favourable improvement in ewe milk production and litter weight weaned. In
commercial flocks for which records for litter weight weaned are not commonly available, producers
may improve litter weight weaned by selecting and culling commercial replacement ewes based on milk
score. Even in purebred flocks where records on litter weight weaned are available, selection for milk
score may result in greater response for litter weight weaned than direct selection on this trait alone
(Snowder et al, 2001a,b).

5.2 Lamb traits

5.2.1 Lamb survival

Despite its importance, lamb survival as a trait of the lamb has not been widely investigated. Most
studies investigating the trait have used a type of analysis where animals are classified as alive or dead
at the end of a predetermined period (Burfening, 1993; Olivier et al., 1998; Matos et al., 2000). The
estimates of heritability of lamb survival from studies using this approach are mostly small and within
the range of 0.00 to 0.09.

However, the use of actual survival times through survival analyses enables discrimination between
lambs dying early or late in the period. Analysis of actual survival time also allows for the use of
records of lambs that were culled or died within the considered period due to reasons not related to their
viability such as slaughter or accidental death, which is relatively common under extensive production
systems (Southey et al., 2001; Carlén et al., 2005; Sawalha et al., 2006).

Using records of Scottish Blackface lamb viability at birth4, and survival from 1-120 days, 1-14 days,
15-120 days and 121-180 days, Sawalha et al. (2006) found significant differences in hazard of
mortality between gender and birth type. Female lambs had a better chance of survival during all
periods compared with males. Twin born lambs had about 50% greater hazard of mortality than singles

4
Viability at birth was defined as a binary trait (all or none) where lambs that survived for at least 24 hours after
birth were coded with 0, and lambs born dead or that died within 24 hours of birth were coded with 1

30
for the early postnatal period (1-120 days) and the hazard of mortality for triplets was about twice as
large as that for twin born lambs for the same period. The effect of type of birth on hazard of mortality
decreased with lamb age.

At birth, the probability of mortality was greater for triplets and singletons than for twin born lambs and
greater by 25% for males compared with females. These findings are in agreement with many other
authors (eg Smith, 1977; Hall et al., 1995; Morris et al., 2000b; Holst et al., 2002; Christley et al., 2003;
Dwyer, 2003). Birth weight has a quadratic relationship with hazard of mortality with both lighter and
heavier lambs at greater risk (eg of hypothermia and dystocia respectively, among other factors).
However, as mentioned above, higher birth weight lambs have a better chance of survival during the
early postnatal period. Selection for optimal rather that maximum birth weight should therefore be
practiced when lamb viability at birth and birth weight are to be improved simultaneously (Sawalha et
al., 2006).

Sawalha et al. (2006) reported estimates of direct and maternal heritability in the range of 0.06 to 0.13
and 0.10 to 0.14 respectively, for viability at birth and postnatal survival until weaning. These are in
the range of most reported heritability estimates in the literature for Scottish Blackface (Riggio et al.,
2005) and other breeds (Fogarty, 1995; Cloete et al., 2001; Southey et al., 2001; Matika et al., 2003). In
general, maternal genetic effects appear to be more important than the direct genetic effect for lamb
viability at birth and preweaning survival (Burfening, 1993; Morris et al., 2000b; Southey et al., 2001;
Sawalha et al., 2006) although survival of lambs is clearly influenced by the genetic merit of both the
lambs and the dams. It has been suggested that to achieve a higher selection gain for lamb survival,
selection indices should include lamb survival as a trait of the lamb and the dam together (Sawalha,
2006).

Conington et al. (2002) predicted that the inclusion of lamb survival as a trait of the lamb in a multi-trait
selection index for hill sheep would increase lamb survival compared to indices with no inclusion of
this trait (see also Conington et al., 2001). This index work has shown that by including lamb losses,
lamb survival and longevity in the breeding goal, gains in productivity can be made without
compromising either ewe or lamb survival.

Favourable estimates of correlations between live body weights and lamb survival pre and post weaning
suggest that selection for improved postnatal survival of lambs is possible without sacrificing genetic
gain in growth performance traits (Abegaz and van Wyk, 2002; Sawalha et al., 2006).

5.2.2 Gestation length

Gestation length is influenced by several factors including sire breed (Fogarty et al., 2005), dam age,
litter size and litter weight at birth. Older dams have significantly longer gestation lengths (Vatankhah
et al., 2000; Koyuncu et al., 2001), as do ewes carrying single lambs compared to those carrying
multiples (Osinowo et al., 1994; Vatankhah et al., 2000; Koyuncu et al., 2001; Dwyer, 2003; Fogarty et
al., 2005). Gestation length increases with total litter birth weight (Knight et al., 1988; Osinowo et al.,
1994; Vatankhah et al., 2000; Fogarty et al., 2005) and may be longer when the dam is carrying a male
lamb compared to a female (Koyuncu et al., 2001; Vatankhah et al., 2000; Fogarty et al., 2005).

Dwyer et al. (1996), in an embryo transfer study to examine the effects of maternal and lamb genotype
on characters of the dam and progeny, found that regardless of ewe breed, gestation length was longer
for Suffolk than for Scottish Blackface lambs. This agrees with the findings of Bradford et al. (1972)
that the genotype of the lamb is more important than the dam in determining gestation length. Fogerty
et al (2005) reported 2-3 days variation in gestation length due to sire breed.

Few estimates of heritability for gestation length are reported in the literature but those that do exist
tend to be moderate to high (0.20, Osinowo et al., 1994; 0.29, Vatankhah et al., 2000). Genetic
correlations between gestation length and litter size of -0.29 and -0.75 (Osinowo et al., 1994 and
Vatankhah et al., 2000 respectively) indicate that selection for a reduction in gestation length may
indirectly increase prolificacy. Osinowo et al. (1994) reported a high positive correlation (0.93)
between gestation length and litter weight at birth.

31
5.2.3 Parturition

The inheritance of ease of parturition is likely to be mainly maternal (Cloete et al., 2002), although the
genotype of the lamb has been shown to affect both the incidence of dystocia and assistance required at
birth (Scales et al., 2000; Fogarty et al., 2005), as well as the early post natal behaviour of the lamb
(Dwyer et al., 1996; Dwyer, 2003).

Embryo transfer between Blackface and Suffolk ewes revealed that the lamb can actively contribute to
its ease of delivery (Dwyer et al., 1996) with lamb characteristics such as birth weight and sex both
contributing to the incidence of birth difficulty (Smith, 1977; Cloete et al., 2002; Dwyer, 2003). Birth
weight is positively correlated with length of parturition (eg SA Mutton Merino 0.14; Dormer 0.19;
Cloete et al., 2002) and ram lambs are generally heavier and have longer parturitions than ewe lambs.
Likewise, singleton and triplet lambs experience more birth difficulty than twin lambs, an effect seen
more strongly in Suffolk ewes than Blackfaces (Dwyer, 2003).

5.2.4 Time taken to stand and suck

Neonate survival is dependent on the co-ordinated expression of appropriate behaviours from both
mother and young to ensure that the young is adequately fed and nurtured. In precocious species, such
as the sheep, the role of neonate behaviour in ensuring survival becomes increasingly important and
may be at least as important as that of the mother (Nowak et al., 1997; Dwyer and Lawrence, 1999a).
The lamb is also an important source of sensory stimuli to the ewe to ensure that maternal behaviour
continues to be expressed towards the lamb (Poindron et al., 1980). Although ewes will show maternal
behaviour towards stillborn lambs, their interest rapidly wanes in the absence of behavioural responses
from the lamb.

Lambs are born with limited tissue reserves so must suck soon after birth to survive. To suck
successfully, the lamb must be able to stand and show appropriate udderseeking behaviour and several
studies have shown lamb survival is enhanced in lambs that stand and suck quickly after birth
(Alexander, 1958; Owens et al., 1985; Cloete, 1993; Dwyer et al., 2001).

The time taken by lambs to stand and to suck has been examined in several breeds and exhibits some
genetic variation (Slee and Springbett, 1986; Alexander et al., 1990b; Cloete et al., 1998). In general,
the lambs of lowland breeds take considerably longer than hill lambs to first stand and take longer to
seek the udder. Some lowland animals, e.g. Southdown, stand relatively quickly after birth but are slow
or only poorly successful in finding the udder (reviewed in Dwyer and Lawrence, 2005). Similar
responses have also been shown in mule (crossbred Blackface and Bluefaced Leicester) lambs
(O’Connor and Lawrence, 1992). Extensively reared breeds are likely to experience a degree of natural
selection for lambs that both stand and suck quickly, which explains the generally good performance of
the hill lambs. The lowland breeds have been subjected to various degrees of selection pressure and
intensive husbandry practices that have kept lambs within the breeding population which may otherwise
have died.

Cloete et al (2002) estimated heritabilities for the interval from birth to standing and from standing to
apparently suckling to be 0.10 and 0.08 respectively in SA Mutton Merinos and 0.22 and 0.12 in
Dormers. Their studies revealed that maternal effects were generally not significant for these traits but
the maternal environment to facilitate suckling, as provided by the dam, was found to play a role in the
neonatal progress of lambs, in agreement with other reports (Kuchel and Lindsay, 1999; Cloete and
Scholtz, 1998). For example, the interval from standing to apparently suckling was shorter in a Merino
line that was selected for multiple rearing ability than in a parallel line that was divergently selected
against multiple rearing ability. The line difference was partially, but not wholly, accounted for by the
inclusion of maternal co-operation with the first suckling attempts of the neonate in the model of
analysis (Cloete and Scholtz, 1998).

32
5.2.5 Ewe recognition and attachment

Sheep are a ‘follower’ species (Lent, 1974), thus the lamb needs to keep up with its dam soon after
birth. This requires that the lamb not only shows appropriate locomotor competence, but that it can
discriminate its own dam from other ewes and is attracted to remain with her. Studies in Merinos
suggested that separation between ewe and lamb was a major contributory factor to lamb mortality
(Stevens et al., 1982; Alexander et al., 1983).

Data from Merino and Blackface crossbred animals compared to purebreds suggest that lamb factors
are implicated in ewe–lamb separation (Stevens et al., 1984; O’Connor and Lawrence, 1992). In
specific tests, singleton Merino lambs that were able to recognise their mothers 12 h after birth had
better survival than lambs that could not (Nowak and Lindsay, 1992). Thus, recognition of its mother
and attraction to her are important lamb behaviour traits for survival.

Lambs are initially attracted to any large objects, but can recognise their mothers at close quarters as
early as 12 h after birth (Nowak et al., 1987), and at a distance by 24 h old (Shillito and Alexander,
1975; Nowak, 1991). Their discriminative ability allows them to avoid potentially aggressive alien
dams and to maintain close contact with their own nursing mothers (Nowak et al., 2000). In lambs,
both visual and auditory cues are involved in mother discrimination whether at close contact or at a
distance (Nowak, 1991). Ewes and lambs answer each other’s bleats and respond to the recorded
vocalisations of their partners (Shillito-Walser et al., 1981; Shillito-Walser et al., 1982) suggesting that
the behaviour of both ewe and lamb may play an important role in the ability of the lamb to recognise
its dam (Nowak, 1991;Terrazas et al., 2002).

Breed comparisons have shown that, in general, extensively bred hill lambs (Dalesbred, Scottish
Blackface) are better at ewe recognition than the more intensively managed lowland breeds (Clun
Forest, Suffolk; Shillito-Walser, 1980; Pickup, 2003). However, whether this is due to better
discriminatory capabilities of the lambs or to maternal behaviour characteristics, particularly ewe vocal
behaviour (Terrazas et al., 2002), is not clear.

Lamb recognition of the ewe is related to lamb sucking activity and colostrum ingestion (Nowak et al.,
1997; Goursaud and Nowak, 1999). The post-ingestive internal state is believed to enhance awareness
of the characteristics of the maternal body, or facilitate the integration of different cues used for
orientation and reunion with the dam. Thus, differences between breeds may also be related to the
quality of sucking interactions.

5.2.6 Cold resistance

Cold resistance as an indicator of exposure is another important determinant of lamb survival in many
environments. Although sheep are particularly resistant to cold weather, and even new-born lambs are
able to maintain body temperature in air temperatures of well below freezing provided the lamb is dry
(McCutcheon et al., 1983), hypothermia of wet neonatal lambs may account for nearly half of all
perinatal deaths (Houston and Maddox, 1974). Hypothermia also causes a decrease in lamb sucking
activity (Alexander and Williams, 1966), so accelerating death from starvation as the limited reserves of
the neonate lamb are rapidly diminished.

Breed differences in mean rectal temperatures of lambs when measured 1 h after birth show that there is
considerable variation in the ability of lambs to maintain homeothermy after birth (Sykes et al., 1976;
Samson and Slee, 1981; Slee and Springbett, 1986; Dwyer, 2002). In general, the hill and feral breeds
of lamb maintained higher rectal temperatures over the first hour of postnatal life than the lowland
breeds. When adjusted for differences in body weight, hill and feral breeds had the highest weight
specific cold resistance (Samson and Slee, 1981).

Experiments with cold resistance using water bath tests have reported heritability estimates of 0.3 (Slee
and Stott, 1986); 0.36 (Wolff et al., 1987) and 0.7 (Slee el al., 1991). Slee and Stott (1986) reported a
realised heritability of 0.27 for increased cold resistance but a realised heritability of zero for decreased

33
cold resistance. In addition, a major gene associated with cold resistance has been identified (Slee and
Simpson, 1991; Simpson & Slee, 1998).

Breed differences in maternal factors, such as maternal grooming behaviour, might contribute to the
lamb’s ability to maintain temperature. However, an un-published comparison of rectal temperatures of
Suffolk x Blackface lambs with pure Suffolk lambs (where there was no difference in maternal
grooming behaviour by the Suffolk dams) suggests that these differences are predominantly due to the
genetics of the lamb (median rectal temperatures (8C) of crossbred lambs = 39.9, Suffolk lambs = 39.2,
P < 0.001; cited in Dwyer & Lawrence, 2005). As discussed above for neonatal lamb behaviours,
intensive management strategies may help to keep lambs with poor cold resistance within the breeding
population when they may have died under more extensive management.

Birth coat depth and skin thickness both show significant genetic correlations with cold resistance (Slee
et al., 1991), suggesting they are important components of the maintenance of homeothermy in lambs
by providing insulation. Skin thickness and coat depth are greatest in the hill breeds in comparison to
lowland breeds, probably due to natural selection for improved cold resistance in the latter (Samson and
Slee, 1981).

Although birth coat and body weight differences play a role in cold resistance, significant breed
differences still remain once these components have been accounted for (Samson and Slee, 1981). This
difference is presumably related to the ability of different breeds to generate endogenous heat.
Research indicates that the lowland intensively managed breeds have a lower ability to generate heat
through non-shivering thermogenesis compared to hill breeds (reviewed in Dwyer & Lawrence, 2005).

The composition of maternal colostrum also contributes to the available lipids for neonatal metabolism.
Although variation in lamb intake probably contributes most to variation in colostral energy supply,
unpublished data show that colostrum from Blackface ewes contains a relatively greater percentage of
lipid than that of Suffolk ewes, thus Suffolk lambs may struggle to obtain sufficient energy to maintain
homeothermy both from their reduced sucking ability and the lower level of lipid in the colostrum they
ingest when compared to hill lambs (Dwyer & Lawrence, 2005).

6 Breeding for other traits

6.1 Tolerance to food shortages

Webster (1993) suggests that selection should include traits which increase tolerance to long term
consequences of prolonged food shortage; not just shortage of energy, but also of N, phosphorus,
copper and selenium. Woolliams et al. (1986) showed that lambs from two genetic lines selected for
low or high copper status differed substantially in mortality and resistance to infection.

Differences in grazing behaviour between and within breeds may indicate that breeds or individuals
within breeds are better adapted to extensive conditions. There are reports of differences between sheep
breeds in the time spent grazing. For example, of six breeds studied by Dudzinski and Arnold (1979)
Suffolks had the longest grazing times, followed by Border Leicester, Dorset Horn and Romney sheep,
whilst Cheviot and Southdown sheep had the shortest grazing times. In a study of Merino ewes, the
individuals with the longer grazing times ended the morning bout later and started the afternoon bout
earlier than those individuals with the shorter grazing times (Arnold and Dudzinski, 1978). Similarly,
Cheviot and Suffolk sheep commenced grazing earlier in both the morning and the afternoon than
Dorset Horn, Southdown and Border Leicester sheep. Romney sheep started grazing later in both the
morning and afternoon, but also finished both bouts later (Arnold and Dudzinski, 1978). It is thought
that the differences in grazing times are mainly accounted for by increased grazing activity during the
grazing bouts.

Certain breeds of sheep which live on hill land, such as the Scottish Blackface and Cheviot sheep, form
home ranges, where the daily patterns of movement, at least initially, are learned from the preceding
generation (Lawrence and Wood-Gush, 1988; Lynch et al., 1992). Other breeds, such as the Merino, do
not establish home ranges, possibly because the same area of land is not used by successive generations.

34
The distance walked by sheep in a day also varies with breed. Cheviot sheep have been shown to walk
further than Romney sheep on both flat and hill land, although the difference between the two breeds
was greater on the hill pasture. Distances walked by individual sheep were similar in successive weeks
(Cresswell, 1960). The topography of the land, type of vegetation and frequency of drinking will all
affect the distance travelled.

The distance between grazing animals can vary from 4 m for Merinos to 19 m for the British hill breeds
(Arnold and Dudzinski, 1978). Kilgour et al. (1975) found that some sheep, Dorset-Romneys in
particular, did not disperse even when herbage availability was very low, whilst Merino sheep have
been reported to disperse only under such conditions (Fraser and Broom, 1990). It is thought that breeds
which remain close together may not be suited to areas where favoured herbage patches are some
distance apart (Hunter, 1960), such as in extensive grazing systems.

Provided that sheep are given sufficient area and choice, they will be selective in their diet (Hafez,
1975; Fraser and Broom, 1990). Sheep may select between different plant species, individual plants,
and parts of the plants available (Lynch et al., 1992). However, there is considerable variation between
individuals of the same breed (Arnold, 1981; Lynch et al., 1992). Arnold (1981) showed that sheep
from the Australian rangelands preferred different pasture plants to sheep kept on sown pastures, and
that differences are strongly influenced by the previous nutritional history of the animal.

Similarly, Key and MacIver (1980) found that when Clun Forest lambs were cross-fostered on to Welsh
Mountain ewes they preferred the tussock and heather vegetation eaten by the ewes, whilst Welsh
Mountain lambs fostered on to Clun Forest ewes preferred the improved pastures grazed by their foster
mothers.

In future, more emphasis on research on within breed genetic variation in grazing behaviour, and
associations with production and animal welfare would be valuable in helping to formulate breeding
goals for extensive systems.

6.2 Longevity

Improving ewe longevity has been documented as being important for livestock profitability
(Conington et al., 2001) and evaluations of this trait are currently used in breeding programmes in the
UK for dairy cows (Veerkamp et al., 1995; Brotherstone et al., 1997).

However estimates for heritability of longevity (defined as days or years in the flock) are low (0.06 for
Australian Dorset sheep, Brash et al.,1994c; 0.08 for Scottish Blackface, Conington et al., 2001),
suggesting that only moderate gains could be made from selection on this trait.

Brash et al. (1994c) reported negative genetic correlations between longevity and number of lambs
born/ewe joined (-0.15), litter size (-0.23) and lamb survival (-1.0). The same authors found longevity
to be positively correlated with ewe fertility (ewes lambing/ewe joined; 0.29) and negatively correlated
with number of lambs weaned per ewe joined (-0.11). In contrast, Conington et al. (2001) reported a
positive correlation between longevity and number of lambs reared (0.35). Conington et al. (2001) also
reported positive genetic correlations between longevity and mature size (0.24), fleece weight (0.26),
average weaning weight (0.21), fat depth (0.20) and muscle depth (0.31). They reported a moderate
negative correlation between number of lambs lost and longevity (-0.35).

Conington et al. (2001) suggest that improving the longevity of the flock within the context of a multi-
trait selection index should ensure that higher flock productivity will not be achieved at the expense of
shorter ewe lifespan. Recording ewe longevity in breed improvement programmes is inexpensive
relative to the benefits gained through genetic improvement.

Longevity in rams is important for commercial lamb producers but is less important in pedigree flocks
and studs where rapid replacement of both rams and ewes will reduce the generation interval and
enhance the rate of genetic progress while limiting the accumulation of inbreeding (Brash et al., 1994c).

35
6.2.1 Teeth and bone

Good bone quality in breeding ewes is important for the mineralisation of foetal skeletons and to sustain
maternal dentition, as tooth loss is one of the main reasons for culling sheep in the UK (Herrtage et al.,
1974; Aitchison and Spence, 1984). Among other functions, bone is a storage depot for calcium and
other key minerals that are mobilised to meet major demands such as during lactation.

As studies in humans and poultry have shown, there is substantial genetic variation for bone properties
(h2 between 0.5 and 0.8), suggesting a similar situation in ewes. These properties, e.g. bone density, are
key to successful production and nurturing of healthy lambs, which can be used in selective breeding
strategies to extend breeding ewes’ productive lives.

CT has been shown to be a useful method of assessing bone properties in sheep (Rubin et al., 2001).
Conington et al (2004) using CT to quantify the main bone types in Scottish Blackface ewes and
investigate environmental factors affecting bone quality found that bone depletion followed a similar
pattern to that of fat and muscle between mating and pre-tupping (Lambe et al., 2003). Age of dam and
heft were not key factors affecting bone properties but the cumulative number of lambs born was. Bone
properties showed weak relationships with body composition or live weight. Variation between sire
progeny groups indicated that there is a significant genetic component to bone properties.

More data are required to allow the estimation of genetic parameters for bone mobilisation and bone
characteristics, and to assess further the key factors affecting bone properties Conington et al. (2004).

6.3 Body composition

The ability of the ewe to maintain her own body composition throughout the reproductive cycle is an
important component for the economic efficiency of the flock. The use of CT in the prediction of
internal and carcass fat levels in hill ewes has shown that the depletion and repletion of fat and muscle
in these depots is under genetic control (Lambe et al., 2005). The use of this information to select
animals that are better able to withstand harsh environmental conditions, yet produce and rear their
lambs with no supplementary feeding, should be an important goal for easy-care sheep breeding
systems. Current research at SAC is investigating the incorporation of this trait into new selection
indices for the hill sheep sector (J. Conington, pers. comm.).

6.4 Wool shedding

It has long been recognised that feral and certain breeds of sheep, such as the Wiltshire Horn, tend to
shed their own wool, reducing the need for shepherding tasks such as shearing and dagging (Tierney,
1978; Rathie et al., 1994). Wool shedding also makes these breeds considerably less susceptible to
blowfly strike of the breech and prepuce region than non-wool shedding breeds and crosses (Tierney,
1978; Litherland et al., 1992; see section 2.4 of this report). Environmental factors such as stress and ill
health cause premature wool loss in many breeds although the degree to which fleece loss occurs in
some breeds (e.g. North Country Cheviot) is under genetic control (Conington et al., 1990).

It has been suggested that in the UK, sheep have more wool than they need, as they need very little
depth for insulation. Winter shearing of inwintered lambs reduces heat stress and therefore respiration
rate and increases the gestation length of ewes, resulting in heavier lambs at birth. Ewe lambs shorn
before mating produce 15% more lambs. Prior to summer shearing ewes are slow to move and graze
less actively (Vipond, 2006).

The Easycare breed developed by Iolo Owen was derived from the Nelson Welsh Mountain (Welsh
Mountain x Cheviot) which were crossed twice to the Wiltshire Horn, a wool shedding breed, and
selected for wool shedding, no horns and easy care traits. Easycares carry a reasonable fleece of up to 1
- 2 inches in length through the winter which they cast in the spring. Typically they are not housed and
are fed minimum amounts of concentrates. Where there is no threat from sheep scab, no dipping is
required as there is a substantially reduced risk of blowfly on non-soiled parts of the wool.

36
Lambs at birth appear to have sufficient wool and, being a leggy breed, quickly get to their feet and start
suckling. The breed has the potential to significantly reduce labour input and is recommended where
shearing is expensive and difficult to organise (Vipond, 2006). From observation, it is clear that there is
a major gene influencing the fleece shedding in the Easycare breed and it would therefore be an ideal
candidate for QTL studies.

6.5 Temperament

The adoption of extensive rearing systems will involve profound changes in animal husbandry. Certain
environmental factors or animal characteristics that were previously of little importance may emerge as
obstacles to the successful implementation of extensive management or prove to be prejudicial to
animal welfare. Of these different factors, animal-human relationships are of prime importance (Le
Neindre et al, 1996).

Studies performed on extensively farmed animals have indicated that behavioural response to humans
(or ‘temperament’, defined as the manner in which individuals react to novel or challenging situations;
Mason, 1984; Wilson et al., 1994) affects production. It has been shown to influence reproductive
performance in heifers, for example, in which ‘emotional’ females have more silent ovulations
(ovulation without sexual behaviour) than ‘unemotional females’ (Gauthier and Thimonier, 1982). In
cattle reared under extensive conditions, reaction to handling seems to determine, at least in part, the
incidence of bruising at the slaughter house (Fordyce et al., 1985). In sheep, Murphy et al (1998) and
O’Connor et al. (1985) have shown that ‘calm’ ewes exhibit better maternal behaviour after parturition
than ‘nervous’ ewes, affecting lamb survival (see section 4.1.3 of this report). Gelez et al (2003) found
that temperament affects ewe sexual behaviour. Calm ewes were more proceptive5 than nervous ones
and tended to be more receptive6 to the ram. In bighorn sheep, Réale et al. (2000) observed that ‘bold’
ewes started reproducing earlier and had a significantly higher weaning success than ‘shy’ ewes.

The reaction of animals to human presence depends on their previous experience and their genetic
characteristics (Le Neindre et al., 1996).

Murphy et al., (1994) established a method for assessment of temperament by combining two
behavioural tests: (1) the ‘arena test’, a motivational choice test that measures the approach and
avoidance behaviour of sheep, similar to tests used to measure reactivity in cattle (Fell et al., 1999).;
and (2) the ‘box test’, similar to an isolation test used in sheep and cattle to measure the degree of
anxiety (Cockram et al., 1994; Burrow, 1997). For sheep, the heritability of these temperament
parameters is around 0.23 and the repeatability from weaning to 3-year old ranges from 0.54 to 0.82
(Martin et al., 2004). For over 10 years, researchers in Australia have used these measures to select two
experimental lines of sheep, a ‘calm’ line and a ‘nervous’ line, to create the ‘Allandale Flock’ that is
now used to study the relationship between temperament and production characters (Martin et al.,
2004).

The main outcome has been a clear demonstration that calm ewes are better mothers than nervous ewes
(Murphy et al., 1994). The calm ewes spend more time with their lambs, have a shorter flight distance
when disturbed and return to their lambs faster than nervous ewes. Consequently, lamb mortality from
birth to weaning for calm ewes was about half that of nervous ewes. The poor mothering ability of the
nervous ewes was the main factor associated with lamb mortality.

Effectively, the selection process produces animals that show less intense stress responses as they
interact with their environment. In addition to postnatal lamb survival, stress disrupts many other
aspects of the reproductive process, all of which might be improved by genetic selection for ‘calm
temperament’. These include the length of the oestrous cycle (Braden and Moule, 1964; Przekop et al.,
1984), ovulation rate (Doney et al., 1976), the proportion of ewes mated (McMillan and Knight, 1982),
embryo survival (Van Niekerk et al., 1968) and sexual behaviour (Gelez et al., 2003). Finally, in other
species, temperament seems to improve other aspects of production such as growth rate (Voisinet et al.,

5
Proceptivity = time spent in close contact with males
6
Receptivity = percentage of immobilizations in response to nudges by the male

37
1997; Burrow, 1998; Fell et al., 1999), immune function (Fell et al., 1999), milk yield (Lawstuen et al.,
1988) and meat quality (Jones and Hocking, 1999; Reverter et al., 2003). All of these outcomes would
improve productivity without compromising animal welfare or ethics.

Sheep Genetics Australia is planning to offer Australian Sheep Breeding Values for temperament in the
near future to Lambplan and Merinoselect customers.

7 Conclusions

7.1 Breeding for resistance to disease

7.1.1 Nematodes

Genetic variation in many aspects of host resistance to nematodes is well documented. Including GI
parasite resistance in breeding goals may benefit sheep production enterprises in a number of ways, for
example, (i) direct increases in productivity resulting from favourable genetic correlations between
resistance and performance, (ii) decreased pasture contamination, leading indirectly to improved
performance, (iii) decreased treatment costs and (iv) less tangible benefits such as improved health and
welfare (Bishop et al., 2004). The genetic correlation between resistance and performance will
influence the nature of the benefits of improving resistance. As the correlation becomes progressively
less favourable, the benefits will change from direct improvements in productivity to less easily
quantifiable benefits via decreased pasture contamination, improved health and welfare, and decreased
treatment costs.

If selective breeding for nematode resistance is to be implemented then it is necessary to be able to

quantify host susceptibility or resistance to infection. Faecal egg count (FEC) is the indicator trait
commonly used and the exploitation of host genetic variation in resistance using FEC in commercial
sheep breeding programmes is now well-established in New Zealand and Australia (Morris et al., 1997,
2000b; Woolaston and Piper, 1996; Woolaston and Windon, 2001), leading to reduced dependency on
drug usage. Studies have also demonstrated that it would be feasible, in principle, to use FEC to select
sheep for resistance to gastrointestinal nematode parasites under typical commercial sheep conditions in
the UK where sheep face a natural parasite challenge (Bishop et al., 1996; Bishop et al., 2004).

However, the use of FEC as the only indicator trait is time-consuming and costly. Traits other than FEC
may be used to assess resistance to nematodes or host response to infection:
• Dag score and faecal consistency score have a poor relationship with faecal egg count implying that
they would be ineffective to improve host resistance to parasites.
• Selection for increased IgA activity and eosinophil count might be useful to decrease worm
development and fecundity.
• Families with high fructosamine concentration have long, fecund worms and therefore it may be
possible to use fructosamine concentration to indicate infection status.
• There has been some success in QTL detection, but generally the number of significant QTL
reported is probably less than expected given the degree of international research effort into this
area. Current research aims to fine map QTL, for example using dense SNP markers, and to study
the functional significance of genes that may underlie host responses to infection. Early results
suggest that microarray studies do have the ability to detect genes differentially expressed between
'resistant' and 'susceptible' sheep, with pathways implicated in these differences including the
development of acquired resistance and the structure of the intestinal smooth muscle (Diez-Tascón et
al., 2005).

7.1.2 Footrot

Although selection of sheep with enhanced resistance to the disease using footrot lesion scoring
(appendix 8.1) has been shown to be successful (Skerman, 1985; Skerman and Moorhouse, 1987;
Patterson and Patterson, 1989), genetic markers potentially offer a practical alternative to laborious
scoring and protocols that require exposure to infection.

38
A footrot gene-marker test is now commercially available in New Zealand (Hickford, 2000). Recent
survey results indicate that on properties that have adopted the technology, the use of chemical
solutions to control the disease have been substantially reduced, as have the number of doses of vaccine
and antibiotics. Overall, footrot prevention and control costs have been reduced by between 50 and
70% (Greer, 2005).

While it is possible that the NZ information will be relevant to sheep populations in the UK, the
associations among the molecular genetic markers, footrot resistance and footrot-causing bacteria
strains need to be demonstrated in UK sheep populations. Together with phenotypic data describing
footrot severity and additional information contributed from the exploration of other parts of the Major
Histocompatibility Complex region, it would then be possible to determine whether or not breeding for
enhanced footrot resistance using molecular markers could be a practical and feasible option for the UK
sheep industry.

7.1.3 Mastitis

Unexpectedly from its economic impact there is a dearth of information about the genetics, incidence
and economic consequences of mastitis in meat sheep breeds, and the potential to include mastitis
resistance in meat sheep breeding programmes. Therefore the majority of all relevant literature reported
is on dairy cattle, with some on dairy sheep. The implications of this are that fundamental research on
mastitis in meat sheep breeds is urgently required. This needs to be done at the population level to
understand the major contributing factors to mastitis, the timing, prevalence and ultimately genetics of
the disease. This is important knowledge when choosing the right model for genetic evaluations for
mastitis resistance and will enable the development of robust recording protocols so that unbiased and
objective records of familial susceptibility to mastitis can be understood, and appropriate breeding
programmes instigated to breed mastitis-resistant sheep.

Selection for resistance to mastitis in dairy sheep is currently underway in the French breed Lacaune
using SCC as a proxy trait for mastitis and incorporating it into sheep dairy breeding programmes
(Rupp et al., 2002). However, the continuous measurement of phenotypic indicator traits for mastitis
resistance is time and labour intensive, and the future use of suitable molecular genetic markers could
be an attractive proposition, particularly for meat sheep breeds as it does not require milk sampling.
More research is needed in this area.

7.1.4 Blowfly strike

Blowfly strike is an important disease of sheep, causing the deaths of an estimated 12,000 animals each
year in the UK (Sargison, 2004). Clipped sheep and young lambs with short fleeces are not usually
attacked, but as the length of the fleece increases so does the risk of strike (French et al., 1996).

Differences exist both between and within breeds in susceptibility to fly strike indicating that the
predisposition of sheep to flystrike is in part genetically determined, with potential scope for within-
flock genetic improvement.
An alternative approach to the problem is to introduce new genetic material into the population in an
endeavour to make sheep less susceptible to blowfly strike. Of interest in this respect is the Wiltshire
Horn breed in which fleece wool is shed annually (Slee and Carter, 1961, 1962; Ryder, 1969).

7.2 Breeding for enhanced flock fertility

7.2.1 Ewe prolificacy

Many studies indicate that an increase in ewe prolificacy could potentially be achieved through genetic
selection on litter size, ovulation rate or ram scrotal circumference, or a combination of the three.
However, increasing litter size may also contribute to an increase in lamb mortality, particularly in more
extensive systems, i.e. the optimum is not necessarily the maximum achievable. A target mean value
appropriate to the particular management system, feed resources and lambing season for the flock(s) in
question should be set before a selection programme is initiated. Optimum litter size at birth is

39
influenced by survival rates of singles, twins and higher multiples, and by the growth rate of survivors
in a specific set of management conditions.

Conington et al. (2001) suggest that, particularly in extensive systems, it is more cost effective to farm
ewes with the ability to rear more of their lambs (see sections 4 and 6.3 of this report), rather than
increasing prolificacy per se. This is partly because of the costs involved in supplementary feeding
ewes that are carrying higher litter sizes during winter. This is in agreement with the findings reported
by Ercanbrack and Knight (1998) and Snowder (2002) that selecting directly for total weight of lamb
weaned per ewe rather than litter size or fertility will lead to more improvement. Total litter weight
weaned represents a ‘biological index’ which incorporates variation in fertility, litter size, viability and
growth rate, and no doubt other components not normally recorded such as ewe lamb-rearing ability
(Bradford, 2002).

In the U.S., Australia and New Zealand, litter size born is currently genetically evaluated.

7.2.2 Scrotal circumference

Scrotal circumference may be a useful indirect selection criterion for improvement of female
reproduction. Although results are variable, testis size appears to have a positive relationship with
ovulation rate and possibly also litter size (Matos and Thomas, 1992). The use of rams with larger
testis measurements may allow a reduction in the number of rams required for breeding each year and
increase the overall reproductive efficiency of the flock (Duguma et al., 2002).

However, although various reports indicate that scrotal circumference is moderately to strongly
positively correlated with liveweight at different ages (Brash et al., 1994a,b; Fossceco and Notter, 1995;
Al-Shorepy and Notter, 1996; Duguma et al., 2002), others have reported a negative correlation
between the two traits (Land, 1982; Burfening and Davis, 1998; Haley et al., 1990) raising doubts about
the value of including information on scrotal circumference in a selection programme to improve litter
size.

If selection for testicular size is to be practiced, scrotal circumference growth from 90 to 180d (with
some adjustment for liveweight) appears to be the trait of choice because it generally has the highest
heritability (Matos et al., 1992; Fossceco & Notter, 1995; Al-Shorepy & Notter, 1996).

Sheep Genetics Australia currently offers breeders Australian Sheep Breeding Values for scrotal
circumference at post-weaning, yearling and hogget ages.

7.2.3 Ram serving capacity

Response to selection for serving capacity should be favourable, with selection resulting in rams
capable of mating with more ewes, improving the reproductive efficiency of the flock and reducing the
number of rams needed. Tests for serving capacity7 have been shown to accurately predict the breeding
performance of high and low performance rams (Ibarra et al., 2000; Snowder et al., 2002; Stellflug et
al., 2006) and can identify rams with high sexual performance at an early age (14 months; Snowder et
al., 2002).

Sire sexual performance influences the proportion of high to low performing ram lambs produced, with
sons born to high-performing sires attaining a significantly higher number of ejaculations than sons
born to low performing sires (Bench et al., 2001). Furthermore, daughters of high performing sires are
significantly younger at first behavioural oestrus than daughters of low performing sires (Bench et al.,
2001). However, the general trend for low genetic correlations between sexual performance of rams
and ewe reproductive traits suggests that breeding rams with high sexual performance will have little, if
any, positive effect on genetic improvement of number of lambs born or number of lambs weaned.
Direct selection on the ewe is therefore likely to be more effective for improving lamb production
(Snowder et al., 2004).

7
Serving capacity tests assess the rate at which rams attain successful matings when housed with oestrous females

40
Rams with highly desirable production traits and high serving capacity will leave more offspring for
future generations compared to rams with similar desirable production traits but a low serving capacity,
thus increasing the rate of genetic improvement for both serving capacity and production traits
(Snowder et al., 2002).

7.3 Breeding for enhanced maternal ability and lamb survival

Lamb mortality is a major constraint to efficient sheep production (Alexander, 1988; Haughey, 1991)
with the vast majority of lamb deaths occurring within 1-3 days of birth (Nowack et al, 2000; Southey
et al., 2004). High lamb mortality could be reduced substantially by a short period of labour and by
maternal behaviours which protect the nutritional and thermal state of the lambs (Alexander, 1988;
O’Connor and Lawrence, 1992). This is particularly important in more extensive management systems
where ewes must be able to conceive, carry, give birth to and rear their young with little (or
substantially reduced) human intervention.

In general, maternal genetic effects appear to be more important than the direct genetic effect for lamb
viability at birth and preweaning survival (Burfening, 1993; Morris et al., 2000b; Southey et al., 2001;
Sawalha et al., 2006), although survival of lambs is clearly influenced by the genetic merit of both the
lambs and the dams. Selective breeding has been advocated as a means of improving lamb survival and
ewe rearing ability within breeds, despite the low heritability of these traits.

Conington et al. (2001, 2002) predict that using multi-trait selection indexes, improvements in maternal
characteristics can be achieved alongside increasing lamb weaning and carcase weights. Key potential
index traits include lamb loss from birth to weaning, litter size at weaning and average weight of lambs
weaned. ). Including lamb survival as a trait of the lamb as well as lamb losses in the index is predicted
to reduce lamb ‘wastage’ between birth and weaning (Conington et al., 2002).

The U.S. National Sheep Improvement Programme uses weight of lamb weaned per ewe exposed over a
given time interval as an indicator of ewe production efficiency. This is a complex trait that includes
fertility, prolificacy, maternal ability, lamb survival, and growth and is calculated only for ewes that
wean at least one lamb (Wilson and Morrical., 1991; Notter, 1998).

Maternal ability traits

• An important component of ease of delivery is the pelvic dimension of the ewe. Ewes selected for
lamb-rearing ability have increased pelvic dimensions and therefore experience fewer lambing
difficulties (Kilgour and Haughey, 1993). CT scanning may prove to be a useful way to measure
pelvic capacity in ewes to aid selection for lambing ease but larger trials are necessary to quantify
genetic properties of dystocia and pelvic measurements and their relationship with other production
traits before selective breeding for large pelvic capacity can be recommended (Bilbe et al., 2005).

• Ewe behaviour around lambing time has a large effect on lamb survival, particularly in extensive
systems (Nowak, 1996).
o Isolation from the rest of the flock during and after parturition is an important preliminary step
in the formation of the mother-young bond. Ewes selected for lamb-rearing ability tend to
remain longer on or near their birth sites than control line contemporaries (Knight et al., 1989).
o The ewe forms a memory for her own lambs that allows her to restrict maternal care
exclusively to her own offspring (Poindron et al., 1984a). Lowland ewe breeds managed
intensively tend to show poorer maternal care at birth than breeds managed more extensively
suggesting a relaxation of selection pressure in intensive systems such that variation in maternal
care has less influence on the survival of the lamb (Dwyer and Lawrence, 2005).
o Maternal co-operation shortens the interval from standing to apparently suckling. Ewes
selected for rearing ability are less likely to circle and back than control line ewes and are more
likely to adopt a posture to facilitate suckling (Cloete and Scholtz, 1998).

• Maternal Behaviour Score, based on the proximity of the ewe to her lamb as it is handled, (appendix
8.3) may be a useful tool to measure maternal behaviour. The score has been shown to be related to
both postnatal lamb survival and weaning weight (O’Connor et al., 1985; O’Connor, 1996; Lambe et

41
 al., 2001; Everett-Hinks et al., 2005; Sawalha et al., 2006). The score may reflect the ewes’
underlying temperament as much as it measures their maternal behaviours (Dwyer and Lawrence,
2005).

• Increased milk production of range ewes is associated with increased lamb survival and growth
(Torres-Hernandez and Hohenboken, 1980). Selection on Milk Score8 at early parities or based on
lifetime records may result in favourable improvements in ewe milk production and litter weight
weaned (Snowder et al., 2001 a,b).

Lamb traits
• Birth weight has a quadratic relationship with hazard of mortality with both lighter and heavier
lambs at greater risk (eg Smith, 1977; Hall et al., 1995; Morris et al., 2000b; Holst et al., 2002;
Christle et al., 2003; Dwyer, 2003). Selection for optimal rather than maximum birth weight should
therefore be practiced (Sawalha et al., 2006).

• Favourable estimates of correlations between live body weights and lamb survival pre and post
weaning suggest that selection for improved postnatal survival of lambs is possible without
sacrificing genetic gain in growth performance traits (Abagaz and van Wyk, 2002; Sawalha et al.,
2006).

• The inclusion of lamb survival as a trait of the lamb in a multi-trait selection index for hill sheep
would increase lamb survival compared to indices with no inclusion of this trait (Conington et al.,
2002). To achieve a higher selection gain for lamb survival, selection indices should include lamb
survival as a trait of the lamb and the dam together (Sawalha et al., 2006).

• The genotype of the lamb is more important than the dam in determining gestation length (Bradford
et al., 1972; Dwyer et al., 1996). Selection for a reduction in gestation length may indirectly
increase prolificacy (Osinowoet al., 1994; Vatankhah et al., 2000).

• The inheritance of ease of parturition is likely to be mainly maternal (Cloete et al., 2002), although
lamb characteristics such as genotype, birth weight, sex and birth type (singles/multiples) have been
shown to affect the incidence of dystocia (Scales et al., 2000; Fogarty et al., 2005; Cloete et al.,
2002), as well as the early post natal behaviour of the lamb (Dwyer et al., 1996; Dwyer, 2003).

• The time taken by lambs to stand and suck exhibits some genetic variation (Alexander et al., 1990b;
Cloete et al., 1998) and therefore could be a possible candidate trait for inclusion in a selection
programme.

• Recognition of its mother and attraction to her are important lamb behaviour traits for survival
(Nowak and Lindsay, 1992). Both visual and auditory cues are involved (Nowack, 1991) as well as
sucking activity and colostrum ingestion (Goursaud and Nowak, 1999).

• Breed differences in mean rectal temperatures of lambs show that there is considerable variation in
the ability of lambs to maintain homeothermy after birth (Samson and Slee, 1981; Dwyer, 2002).
Birth coat depth and skin thickness are also correlated with cold resistance (Slee et al., 1991).

7.4 Breeding for other traits

It has been suggested that selection should include traits which increase tolerance to long term food
shortage (Webster, 1993). More research on within breed genetic variation in grazing behaviour and
associations with production and animal welfare would be valuable in helping to formulate breeding
goals for extensive systems.

8
Milk score is a subjective measure of milk production of ewes that can be qualitatively determined by palpating
the ewe’s udder and observing her lambs’ fill within a few hours of lambing (Snowder et al., 2001a,b).

42
Improving ewe longevity has been documented as being important for livestock profitability and,
despite its low heritability, inclusion of the trait in a multi-trait selection index should ensure that higher
flock productivity will not be achieved at the expense of shorter ewe lifespan (Conington et al., 2001).
Recording ewe longevity in breed improvement programmes is inexpensive relative to the benefits
gained through genetic improvement.

Good bone quality in breeding ewes is important for the mineralization of foetal skeletons and to
sustain maternal dentition. CT has been shown to be a useful method of assessing bone properties in
sheep (Conington et al., 2004) but more data are required to allow the estimation of genetic parameters
for bone characteristics.

The ability of the ewe to maintain her own body composition throughout the reproductive cycle is an
important component for the economic efficiency of the flock. The use of CT in the prediction of
internal and carcass fat levels in hill ewes has shown that the depletion and repletion of fat and muscle
in these depots is under genetic control (Lambe et al., 2005). The use of this information to select
animals that are better able to withstand harsh environmental conditions, yet produce and rear their
lambs with no supplementary feeding, should be an important goal for easy-care sheep breeding
systems. Current research at SAC is investigating the incorporation of this trait into new selection
indices for the hill sheep sector (J. Conington, pers. comm.).

Selection for wool shedding may result in sheep requiring less shepherding input (e.g. shearing and
dagging) and that are less susceptible to blowfly strike (Tierney, 1978; Rathie et al., 1994). More data
are required to allow the estimation of genetic parameters for this trait.

Selection for animals with a calm temperament produces animals that show less intense stress responses
as they interact with their environment, including stockpeople (Martin et al., 2004). This could result in
improvements in postnatal lamb survival and many other aspects of reproductive and productive
efficiency. Behavioural tests have been developed which allow the effective assessment of
temperament (Murphy et al., 1994).9 Sheep Genetics Australia is planning to offer Australian Sheep
Breeding Values for temperament in the near future to Lambplan and Merinoselect customers.

7.5 Application to the UK sheep industry

Some of the traits included in the report are already used in UK sheep breeding schemes (e.g. ewe
longevity is predicted in breeding programmes for hill sheep although it is not recorded separately in
practice). Other traits are in the pipeline to be included in breeding programmes (e.g. direct lamb
survival) and others are still being researched (e.g. resistance to footrot, nematodes and mastitis).

Difficulties that may be encountered when trying to incorporate some of the traits into breeding
programmes include:
• insufficient evidence for genetic variation in the trait concerned (e.g. bone quality);
• insufficient consensus or knowledge about the ‘best’ trait to be recorded to achieve the breeding
goal (e.g. resistance to nematodes);
• traits that are difficult, time-consuming or expensive to record (e.g. behavioural traits);
• the fact that ‘hidden’ benefits of trait improvement may not immediately appeal to some breeders
because their cumulative benefits are not realised until some time in the future (e.g. longevity).

However, most of the traits reviewed could potentially be used in UK sheep breeding programmes. The
use of genetic information from related animals would be particularly useful where the heritability of
the traits is low or if they are difficult to record.

9
Behavioural tests for the assessment of temperament in sheep include: (1) the ‘arena test’, a motivational choice
test that measures the approach and avoidance behaviour of sheep, similar to tests used to measure reactivity in
cattle (Fell et al., 1999).; and (2) the ‘box test’, similar to an isolation test used in sheep and cattle to measure the
degree of anxiety (Cockram et al., 1994; Burrow, 1997); or a combination of the two (Murphy at al., 1994).

43
7.6 Recommendations

Some of the traits considered in the report could be recorded immediately. These include: i) lamb
survival, ii) ewe longevity, iii) ewe fat levels, iv) ‘gestation’ length, v) lambing ease and vi) ram scrotal
circumference. The incorporation of these into indexes would require varying degrees of programming
effort including full genetic analyses of traits iv), v) and vi) and the identification of associations with
other traits that are part of the breeding objective. Faecal Egg Count could also be recorded now but as
the application of the use of this indicator trait for nematode resistance in the UK is yet to be established
(e.g. when to sample, how many sampling times, recommended levels of pasture contamination before
sampling etc.), in order to make the best use of this technology it may be preferable to wait until this
knowledge becomes available.

For flocks that have problems with mastitis, the use of Somatic Cell Count (SCC) could be instrumental
in improving resistance to this disease. Further research is needed to quantify the genetic variation in
this trait and to develop a protocol for its inclusion into breeding programmes.

More work is required (which is currently underway) before molecular tests for resistance to disease
(footrot, nematodes, mastitis) can be used in practice.

A promising trait for easy care systems is ram libido, as defined by ram serving capacity. The
verification of existing tests from other research groups should be done using UK sheep breeds to
establish a useful protocol that could be used in the field. Further research work is needed before the
commencement of recording.

Further research is also needed for most of the behavioural traits, including the use of either physical or
physiological proxy traits.

44
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9 Appendices

9.1 Footrot scoring system

8-point footrot scoring system developed by Raadsma et al., 1993

Trait Range Derivation

Affected - At least one foot affected, adjusted for healing
Underrun - At least one foot underrun, adjusted for healing
No. of feet 0-4
affected
No. of feet 0-4
underrun
Overall grade 0-5 Overall grade
Healing - At least one affected foot showing a reduction in the inflammation of the
lesions, with signs of dry footrot lesions.
Healed, partial - At least one affected foot showing complete healing of footrot lesions,
with no signs of inflammation or exudation in the footrot lesion.
Healed, total - All affected feet showing complete healing of footrot lesions, with no
signs of inflammation or exudation in any of the footrot lesions.

9.2 Molecular markers for mastitis resistance in cattle

Chromo Trait Closest P(%) Location Reference

marker confidence
(position interval
cM) (cM).2/
3 CM Klungland et al., 2001
(putative)
4 CM Klungland et al., 2001
(putative)
6 CM Klungland et al., 2001

9 CM TGLA73 ***(chromosome Holmberg &

) Andersson-Eklund,
†(genome) 2004a
10 CM 0.02 1 Schulman et al., 2004
11 CM INRA177 *(chromosome) Holmberg &
NS Andersson-Eklund,
(genome) 2004a
11 CM <0.01 1 Schulman et al., 2004

14 CM Klungland et al., 2001

(putative)
18 CM TGLA227 0.02(P-value at 120 Schulman et al., 2004
chromosome-
wise significance
level)
25 CM ILSTS102- *(chromosome) Holmberg &
RM404 NS Andersson-Eklund,
(genome) 2004a
27 CM Klungland et al., 2001
(putative)

72
Chromo Trait Closest P(%) Location Reference
marker confidence
(position interval
cM) (cM).2/
14 CM(cofacto BMS1747- <0.01(P-value at 25 Schulman et al., 2004
r analysis) BMS740 chromosome-
wise significance
level)
21 CM(cofacto RM151- 0.01(P-value at 23 Schulman et al., 2004
r analysis) INRA103 chromosome-
wise significance
level)
6 Quality of 0.000/0.008 89 Hiendleder et al., 2003
udder
3 SCC BMC5227 0.0317(chromoso 171 Schrooten et al., 2000
me-wise P-value)
7 SCC BMS2258- 0.025(chromoso 107 Kuhn et al., 2003
(putative) OarAE129 me-wise)
9 SCC CSSM56 **(chromosome) Holmberg &
Andersson-Eklund,
NS(genome) 2004a
11 SCC BMS7169 ****(chromosom Holmberg &
e) Andersson-Eklund,
2004a
**(genome)
18 SCC BM7109- 0.0103(chromoso 70 Schrooten et al., 2000
ILSTS002 me-wise P-value)
18 SCC TGLA227 0.058(genome- 117 Kuhn et al., 2003
wise)
23 SCC BM1443 *(chromosome) Holmberg &
NS Andersson-Eklund,
(genome) 2004a
27 SCC BM3507- 0.004(chromoso 8 Kuhn et al., 2003
(putative) TGLA179 me-wise)
10 SCC TGLA378- 0.027(chromoso 49 Kuhn et al., 2003
(putative) TGLA102 me-wise)
1 SCS <0.01(P-value at 59 Schulman et al., 2004
(cofactor chromosome-
analysis) wise significance
level)
3 SCS 0.01 (P-value at 105 Schulman et al., 2004
chromosome-
wise significance
level)
5 SCS BL37- 9.1(F statistic) 54 Ashwell et al., 2004
BM1819 (chromosome-
wise) (P<0.01)
7 SCS BM6117- 8.8(F statistic) 61 Ashwell et al., 2004
BMS2258 (chromosome-
wise) (P<0.01)
7 SCS BM6117- 3.2 (F statistic) 67 Ashwell et al., 2004
BMS2258 (chromosome-
wise)
(P<0.01)
11 SCS 0.03 (P-value at 63 Schulman et al., 2004
chromosome-
wise significance

73
Chromo Trait Closest P(%) Location Reference
marker confidence
(position interval
cM) (cM).2/
level)

14 SCS 0.01 (P-value at 63 Schulman et al., 2004

chromosome-
wise significance
level)
SCS
35_45 /
(genome- BMS2684
15 0.05 22_48 Boichard et al., 2003
wise
significant)
15 SCS BMS2684- 9.8(F statistic) 34 Ashwell et al., 2004
HBB (chromosome-
wise) (P<0.01)
18 SCS 0.02 (P-value at 113 Schulman et al., 2004
chromosome-
wise significance
level)
20 SCS RM310- 11.8(F statistic) 29 Ashwell et al., 2004
TGLA126 (chromosome-
wise) (P<0.01)
21 SCS <0.01(P-value at 51 Schulman et al., 2004
(cofactor chromosome-
analysis) wise significance
level)
22 SCS BMS875- 3.32 (F statistic) 80 Ashwell et al., 2004
BM4102 (chromosome-
wise)
(P<0.01)
23 SCS BB705- 12.7(F statistic) 50 Ashwell et al., 2004
BM1818 (chromosome-
wise) (P<0.01)
23 SCS BB705- 3.02 (F statistic) 41 Ashwell et al., 2004
BM1818 (chromosome-
wise)
(P<0.01)
23 SCS 0.01(P-value at 7 Schulman et al., 2004
(cofactor chromosome-
analysis) wise significance
level)
24 SCS <0.01(P-value at 28 Schulman et al., 2004
(cofactor chromosome-
analysis) wise significance
level)
26 SCS Centromere- 11.2(F statistic) 0 Ashwell et al., 2004
BM1314 (chromosome-
wise)
(P<0.01)
26 SCS Centromere- 11.0(F statistic) 0 Ashwell et al., 2004
BM1314 (chromosome-
wise)
(P<0.01)

74
 Chromo Trait Closest P(%) Location Reference
marker confidence
(position interval
cM) (cM).2/
26 SCS Centromere- 2.72 (F statistic) 0 Ashwell et al., 2004
BM1314 (chromosome-
wise)
(P<0.01)
27 SCS 0.02 (P-value at 3 Schulman et al., 2004
chromosome-
wise significance
level)
29 SCS BMC1206- 8.0 (F statistic) 50 Ashwell et al., 2004
BMS1948 (chromosome-
wise)
(P<0.01)
29 SCS 0.01 (P-value at 14 Schulman et al., 2004
chromosome-
wise significance
level)
9 SCS BMS1967 1.3 Boichard et al., 2003
(chromosom
e-wise
significant)
21 SCS TGLA122 1.5 Boichard et al., 2003
(chromosom
e-wise
significant)
23 SCS RM33 1.7 Boichard et al., 2003
(chromosom
e-wise
significant)
10 SCS DIK20 0.9 66_90 / Boichard et al., 2003
(genome- 6_92
wise
significant)
5 Udder depth 0.046 106 Hiendleder et al., 2003
(chromosome- (location)
/experiment-wise
thresholds)
6 Udder depth 0.000/0.008 89 Hiendleder et al., 2003

9.3 Maternal behaviour scores at tagging

Description of maternal behaviour score (MBS)a MBS Score interpretationb

Ewe flees at the approach of the shepherd, shows no 1
interest in the lambs and does not return.
Ewe retreats further than 10m but comes back to her 2 Poor mothers – cull annually
lambs as the shepherd leaves them.
Ewe retreats to such a distance that tag identification is 3 Good mothers
difficult (5-10m)
Ewe retreats but stays within 5m 4
Ewe stays close to the shepherd during the handling of her 5 Excellent mothers
lambs.
a
As described by O’Connor et al. (1985)
b
Everett-Hincks et al., 2005

75
9.4 Trait checklist

Trait Av. Definition Ram, Age of Ease of Expensive Key references

Heritability ewe or measurement recording to record
lamb
Breeding for resistance to disease
Nematodes
Faecal egg count 0.2 – 0.4 Number of eggs per gram of faeces after a Lamb FEC 1: 6-8 weeks Time Yes Morris et al., 1997 and
period of known challenge of infective after weaning consuming 2000a; Woolaston and
larvae FEC 2: 6-8 weeks Piper, 1996;
after FEC1 Woolaston and
Windon, 2001
Immunoglobulin 0.57 Activity of IgA against a somatic extract Lamb 7-9 months Involves Yes but Strain et al., 2002
A (IgA) levels of 4th stage larvae from T. circumcincta blood less than Davies et al., 2005
sampling & FEC
lab analysis
Eosinophil 0.35 Concentration of cells in blood Lamb - Involves Yes Stear et al., 2002
counts blood Davies et al., 2005
sampling &
lab analysis
Fructosamine 0.39 Concentration in plasma Lamb - Involves Yes Stear et al., 2001
concentrations blood Davies et al., 2005
sampling &
lab analysis
Footrot
Footrot lesion 0.28 – 0.53 4 to 8-point scale based on degree of Any Any Subjective No Skerman et al, 1988
score infection severity Raadsma et al., 1994
Footrot Gene- genetic variation within the ovine MHC - - - Yes Hickford, 2000
Marker Test loci in the class II region, specifically at
(New Zealand) the DQA2 gene
Mastitis
Somatic Cell 0.04 – 0.24 Counts of epithelial and inflammatory Ewe 1st lactation Time Yes Conington et al., 2005
Counts cells (per ml) from a milk sample (usually onwards consuming
taken on ‘test’ days, i.e. at regular and difficult
intervals during lactation). in meat
breeds
Breeding for enhanced flock fertility

76
Trait Av. Definition Ram, Age of Ease of Expensive Key references
Heritability ewe or measurement recording to record
lamb
Ovulation rate 0.15 Number of corpora lutea on each ovary Ewe From first Laparoscopy Yes Safari & Fogarty, 2003
oestrous
Fertility 0.08 Number of ewes lambing per ewe joined Ewe At lambing Easy No Safari & Fogarty, 2003
Litter size 0.13 Number of lambs born per ewe lambing Ewe At lambing Easy No Safari & Fogarty, 2003
Scrotal 0.21 Greatest diameter of the scrotal sac Ram Optimum 90-180 Easy No Safari & Fogarty, 2003
circumference lamb days
Serving capacity 0.25 Rate at which rams attain successful Ram From 14 months Time Yes, Stellflug et al., 2006
matings when housed with estrous consuming probably Snowder et al., 2002
females
Breeding for improved maternal ability and lamb survival
Maternal traits
Gestation length 0.20 – 0.29 Days from joining to birth of lamb(s) Ewe First parity Easy No Osinowo et al., 1994
onwards Vatankhah et al., 2000
Lambing 0.05 – 0.17 Interval between the first definite sign of Ewe During parturition Easy but No Cloete et al., 2002
ease/length of impending parturition in the ewe to the time
parturition birth of a specific lamb. consuming
Pelvic capacity - Mean pelvic dimensions Ewe - Involves CT Yes Bilbe et al., 2005
scanning
Ewe lamb- 0.06 Ratio of lambs weaned to lambs born Ewe When lambs are Easy No Cloete et al., 2003
rearing ability weaned
Maternal 0.13 5-point scale based on the distance a ewe Ewe When lambs are Easy No O’Connor
Behaviour Score (Lambe et retreats from her lambs when the tagged et al., 1985
al,2001) shepherd is tagging them (appendix 1) Lambe et al., 2001

0.09
(Everett-
Hincks et
al., 2005)
Maternal - 6-point scale based on active cooperation Ewe Straight after Easy but No Cloete et al., 2003
cooperation score of the ewe post-partum (standing still, parturition time
adopting a slightly hunched posture to consuming
enable access to the udder and nudging
the lamb in a position to facilitate
suckling.

77
Trait Av. Definition Ram, Age of Ease of Expensive Key references
Heritability ewe or measurement recording to record
lamb
Period ewe 0.20 (just 1 Ewes regarded as having left their birth Ewe Straight after Easy but No Cloete et al., 2003
remains on or study) sites permanently after having moved parturition time
near birth site away more than 15m for more than 2h consuming
Milk score 0.08 – 0.26 6-point score based on udder size and Ewe Straight after Easy but No Snowder et al.,
udder milkiness (often divided into 3 lambing, 1st subjective 2001a,b
categories: ‘low’, ‘medium’ and ‘high’) parity onwards Sawalha et al., 2005
Lamb traits
Lamb survival 0.06-0.13 Survival time Lamb Birth to weaning Easy No Sawalha et al., 2006
(and beyond if
desired)
Time taken to 0.08 – 0.12 Interval from birth to apparently suckling Lamb Birth Easy but No Cloete et al (2002)
stand and suck requires
careful
observation
Cold resistance 0.3 – 0.7 Time taken for the lambs’ rectal Lamb Few days after Easy No Samson and Slee, 1981
temperature to fall to 35 0C when birth Slee and Stott, 1986
immersed in a cooling water bath Slee et al., 1991
Breeding for other traits
Longevity 0.06-0.08 Days in the flock or days to final joining Ewe At culling Easy No Brash et al., 1994c
Conington et al., 2001
Bone quality - Areas and density of spongy and compact Ewe Any age Involves CT Yes Conington et al., 2004
bone in the tibia scanning
Body - Prediction of depletion and repletion of Ewe - Involves CT Yes Conington pers.
composition internal and carcase fat and muscle scanning comm.
Wool shedding - - Ewe - Easy No Conington, 1990
and Vipond, 2006
ram
Temperament 0.23 Behavioural tests measuring approach and Any Any Time Yes Murphy et al., 1994
avoidance behaviour and degree of consuming Martin et al., 2004
anxiety in isolation and requires
special
equipment

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ISBN 1-84555-000-5

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