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Strategies for Reversing the Trend Towards

Subfertility in Dairy Cattle
ARTICLE in THE VETERINARY JOURNAL AUGUST 2000
Impact Factor: 2.17 DOI: 10.1053/tvjl.1999.0450 Source: PubMed

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The Veterinary Journal 2000, 160, 5360

doi: 10.1053/tvjl.1999.0450, available online at http://www.idealibrary.com on

Review
Strategies for Reversing the Trend Towards Subfertility in
Dairy Cattle
M. ROYAL, G. E. MANN and A. P .F. FLINT
Cattle Fertility Research Group, School of Biological Sciences, University of Nottingham, Sutton Bonington Campus, Loughborough,
Leicestershine LE12 5RD, UK

SUMMARY
The calving rate of the modern dairy cow is declining at approximately 1% per annum. First service conception rates are now below 40%, and the average cow requires more than two inseminations to get her in
calf. We review here genetic, endocrine and nutritional strategies for reducing subfertility, and we emphasize the role of milk progesterone data in the analysis of the condition. Milk progesterone data underpin
three approaches to treating subfertility; firstly, they allow the identification of specific reproductive abnormalities which can be treated pharmacologically. Secondly, they show that at least one endocrinopathy is
heritable, thereby providing the basis for a selective breeding strategy. Thirdly, they provide a means for
monitoring ovarian (and indirectly uterine and conceptus) function during early pregnancy, which are
central to the successful establishment of pregnancy, and are amenable to dietary modification. These
approaches to reversing the falling fertility of dairy cows are characterized by different timescales.
KEYWORDS: Dairy; cow; fertility; reproduction; subfertility.

INTRODUCTION
The three principal problems facing the UK dairy
farmer are subfertility, mastitis and lameness. Of
these, subfertility has the highest economic cost
and is the most difficult to treat. We define subfertility broadly as any condition leading to failure to
establish a pregnancy following completion of uterine involution at 4050 days post partum. Failure to
establish a pregnancy at the expected time may
reflect a number of abnormalities, including failure to ovulate, failure to show oestrus, inappropriate patterns of ovarian cyclicity and loss of
pregnancy. These conditions may reflect dysfunction at the hypothalamic, pituitary, ovarian or uterine level, or in conceptus development. Therefore,
subfertility appears more complex than at first

Correspondence: A.P.F. Flint. Fax: +44 115 951 6302; E-mail:

anthony.flint@nottingham.ac.uk
1090-0233/00/010053 + 08 $35.00/0

2000 Harcourt Publishers Ltd

appears and there are many potential remedies.

The complexity of the condition makes it difficult
to analyse, diagnose and treat. However, treatments
must urgently be found, as the problem is worsening rapidly.
Pregnancy rates to first service in the UK have
reached an all time low of 40% (Darwash et al.,
1999; Royal et al., 2000). This has a major impact
on dairy economics, as the costs of temporary
infertility include lost income from milk sales, feed
and quota costs, and semen costs where animals are
repeatedly artificially inseminated. These amount
to about 3 per cow per day (Esslemont & Peeler,
1993), and 60% of cows can be assumed to be infertile for 30 days per year. To this must be added the
costs of culling and replacing subfertile cows. One
in 10 cows is culled due to difficulty getting them
into calf, at a total cost (milk losses plus replacement costs) of 600 each. Given there are approximately 2.5 million cows in the UK national herd,
2000 Harcourt Publishers Ltd

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THE VETERINARY JOURNAL, 160, 1

producing 14 billion L of milk per annum, these

costs add approximately 2p per L to the price of
milk. At a milk price to the farmer of approximately 18p per L, this represents 10% of the value
of the product at the farm gate. In addition to its
economic value, subfertility also has important animal welfare implications in terms of treatments to
the cow and premature culling of persistently infertile animals. Furthermore, time spent dealing with
subfertility detracts from time available for other
welfare concerns. It is interesting to note that the
impact of subfertility on dairy economics has only
recently been estimated (Esslemont & Peeler, 1993;
Stott et al., 1999). In a recent questionnaire, dairy
farmers cited subfertility as their greatest concern,
more important than mastitis or lameness (Moffitt,
1995), and between 1990 and 1992 poor fertility
accounted for a higher proportion of cows culled
than mastitis, spongiform encephalopathy or lameness (36.5% disposals compared with 10.1, 7.4 and
5.6%, respectively; Esslemont & Kossaibaiti, 1997).
The situation is complicated by interactions
between these conditions, as lameness may lead to
subfertility (Collick et al., 1989).
ASSESSMENT OF SUBFERTILITY
There are a number of ways to monitor fertility in
dairy cattle. These include recording interval to
first service, days open, calving interval, conception
rate to first service and number of services per conception. However, measures of this kind, which may
be described as traditional fertility parameters, are
traits which are biased by management decisions
and by inadvertent data censoring (for instance,
culling animals before they are inseminated).
An alternative range of fertility parameters provided by measurement of milk progesterone levels,
might be called physiological fertility parameters.
These indicate when an animal ovulates, and reflect
the formation and lifespan of the corpus luteum, as
well as whether oestrous cyclicity is typical, and a
pregnancy is maintained. Although physiological
function as monitored by endocrine measures such
as circulating progesterone concentrations may be
affected by an animals body condition (condition
score), by stress or by environmental factors, they
are not likely to be affected by management decisions and data censoring.
Analysis of reproductive parameters based on
milk progesterone levels identifies a number of
abnormalities of ovarian function associated with
reduced fertility. These include a long period of

post partum anoestrus (a delay between calving and

ovulation), a lengthy anoestrous period between
cycles (which might reflect delayed ovulation, presence of a cystic follicle or absence of ovarian function), and a persistent corpus luteum in the first or
subsequent cycles post partum. Each of these atypical patterns has been statistically defined by
Lamming and Darwash (1998). The total number
of cows experiencing one or more of these atypical
characteristics has risen significantly (P < 0.0005)
over the past 20 years by 12 to 44% (Royal et al.,
2000). However, what is particularly striking is that
the incidence of persistent corpus luteum in the
first cycle post partum has risen from 7.3% in 1683
British Friesian cows (2503 lactations) from 20
herds sampled between 1975 and 1982 (Darwash et
al., 1999), to 18.2% in 714 Holstein-Friesians on
seven farms between 199598 (Royal et al., 2000).
The interval to first service lengthened from 73.9
to 77.6 days and the calving rate to first service
declined from 55.6 to 39.7%. A value of approximately 40% for current rate of calving to first service is supported by independent studies of Darwash
et al. (1999) and P. Ball (personal communication).
Data covering the intervening period (Esselmont,
1992) suggest a calving to first service rate of 50%
in 1989. The calving rate to first service therefore
dropped between the two data collection periods
in 197582 and 199598 at approximately 1% per
annum. This compares with a decline in rates of
conception of first service in the USA of approximately 0.45% per annum between 1975 and 1997
(Butler & Smith, 1989; Beam & Butler, 1999).
GENETICS
Analysis of milk progesterone data reveals that
certain endocrine abnormalities are heritable
(Darwash et al., 1997a). In the past, fertility traits
have not generally been considered heritable.
However, the traits examined were traditional fertility parameters, which do not solely reflect the
physiological characteristics of the cow, due to the
influence of managerial decisions. Recent re-examination of such traditional parameters confirms
their low heritability (typically <0.05; Pryce et al.,
1997), although they do display relatively high
coefficients of genetic variation, which would indicate room for genetic improvement. Notwithstanding, genetic improvement based on
traditional fertility parameters would be slow
since in addition to low heritability the use of
these traits is limited by biological (sex and age

CATTLE FERTILITY

limitations) and structural (accuracy/reliability of

data recording) constraints (Darwash et al., 1999).
One particular measure of ovarian function, the
commencement of luteal activity post partum (CLA)
was shown to be moderately heritable (range for
various transformations, h2 = 0.130.28) and repeatable (r2 = 0.28) in the 197582 data base on British
Friesian cows described above (Darwash et al.,
1997a). CLA was also shown to be heritable in
Dutch HolsteinFriesian cattle in their first lactation (Veerkamp et al., 2000; h2 = 0.16). In the British
Friesians, the mean interval to CLA was 28.7 days, in
the Dutch Holstein-Friesians, 29.5 days (Veerkamp
et al., 2000). Shorter intervals were positively correlated with increased fertility in British Friesians
(between 197582), as for each day by which CLA
increased there was a delay of 0.24 and 0.41 days (P
< 0.001), respectively, in the intervals to first service
and conception (Darwash et al., 1997b). Early reestablishment of post partum ovarian activity has
therefore been established as an important pre-requisite for high fertility, which is consistent with
reports by Thatcher and Wilcox (1973), Whitmore
et al. (1974), Silvia et al. (1992) and Stevenson et al.
(1983). However, this remains a controversial area,
since these reports are in disagreement with Ball
and McEwan (1998) and Smith and Wallis (1998),
and therefore require confirmation within the present UK dairy herd of HolsteinFriesians. Other
endocrinopathies associated with subfertility
included length of the inter-luteal interval and persistent corpora lutea (PCL; reflecting delayed luteolysis), but it is not yet known to what extent these
conditions affect fertility.
Although the use of physiological measures of
fertility, such as CLA, in breeding programmes are
ultimately the preferred parameters to help halt
and reverse the decline in fertility (due to reasons
discussed previously), these parameters are in need
of further investigation. For example, estimates of
heritability and phenotypic/genetic correlation
with other economically important traits need
refining for the modern dairy cow population
before successful incorporation into selection
indices can be achieved. This work is currently in
progress at Nottingham University in collaboration
with the Roslin Institute. However, in the meantime, the low heritability of traditional measures of
fertility does not prevent their worthwhile inclusion in breeding programmes (Woolliams, 1997).
In fact, the genetic correlation between one traditional parameter, calving interval, and condition

55

score is currently under investigation at the

Scottish Agricultural Colleges (Edinburgh) and the
University of Edinburgh (Pryce et al., 2000).
Condition score has been recorded by HUKI
(Holstein UK and Ireland; formally HFS) since
1996 as part of its linear type classification scheme.
As calving dates are reliably recorded, most cows
that survive to their second lactation have calving
interval data. Condition score (CS) is believed to
be related to energy balance (Veerkamp, 1998) and
has a heritability much greater than most traditional fertility measures of approximately 0.20.3
(Jones et al., 1999). On the assumption that energy
balance and fertility are related, CS is a possible
selection criterion for fertility. The results of this
research suggest there may be an opportunity to
select indirectly for fertility using a combination of
calving interval and condition score information,
which could be implemented at low cost and in the
near future.
Although the physiological fertility parameters
described above would be an improvement upon
traditional parameters, a further improvement in
response to selection for female fertility could be
gained through the use of indirect selection criteria for these parameters. This might for instance
involve a heritable trait in the male which is measurable in early life and genetically correlated to
female fertility. The LH secretory response to a
GnRH challenge is one possible parameter, which
is heritable in tropical beef bulls, as judged by
testosterone responses (MacKinnon et al., 1991).
The genetic reversal of the trend in subfertility is
a long-term goal; it is unlikely we shall see substantial genetic progress in fertility for at least a decade.
In the meantime, milk progesterone analysis allows
other approaches to the problem. The most immediately productive of these is probably endocrine
therapy.
ENDOCRINE THERAPIES
Having successfully initiated a return to cyclicity
post partum, the cow must meet two other
endocrine criteria. Firstly, she must express oestrus
behaviour sufficiently strongly to be detected, not
by a bull, but by the stockman. Stockmen are not so
sensitive to the onset of heat as bulls (and are probably considerably busier) so a particularly strong
manifestation of heat may be necessary. Secondly,
she must ovulate appropriately and form a corpus
luteum able to secrete sufficient progesterone. The

56

THE VETERINARY JOURNAL, 160, 1

conceptus must produce the maternal recognition

of pregnancy signal (trophoblast interferon; IFN-)
by day 16 post-insemination, in order to prevent
uterine prostaglandin secretion and luteolysis
(Mann et al., 1999), and it must have undergone
elongation in order to produce the signal in sufficient quantities. When conceptus growth is compromised by poor uterine secretory function,
luteolysis (progesterone withdrawal) occurs and
pregnancy fails. Conceptus growth is progesteronedependent, a low level of progesterone effectively
resulting in starvation of the conceptus. Mann et al.
(1999) showed that a late post-ovulatory progesterone rise to low luteal phase levels results in poor
embryo development and little or no interferon
secretion on day 16: the concentration of interferon- in uterine flushes on day 16 is related to circulating concentration of progesterone between
days 4 and 5 (r2 = 0.4; P < 0.05; Mann et al., 1998).
Therefore, in this context, sufficient progesterone
means a concentration high enough to trigger a
normal pattern of uterine secretory function by
day 5 after insemination. Cows with circulating concentrations below this threshold, due to a late postovulatory rise in progesterone secretion, have
higher rates of failed conception or embryonic loss
than those above it. The threshold concentration
in the systemic circulation is reflected in a threshold concentration in milk (which varies between
assay techniques approximately in the range
27 ng/mL), and cows suffering from luteal inadequacy can be identified by milk progesterone assay
on day 5.
There are three therapies that can be applied to
correct these endocrinopathies. A low day-5 milk
progesterone level can be treated by progesterone
administration via an intra-vaginal progesterone
releasing device (CIDR). A meta-analysis of numerous studies using progesterone supplementation
showed an overall 5.2% improvement in pregnancy
rate following treatment compared with a 10.3%
improvement in studies where progesterone was
administered before day 6. A further improvement
in pregnancy rate of 19.3% was seen when progesterone was administered to cows exhibiting low
fertility (Mann & Lamming, 1999). Early progesterone supplementation increases the rate of conceptus development; Garrett et al. (1988) showed
that progesterone administered between days 2
and 5 increases conceptus length 10-fold on day 14.
In view of the dependence of conceptus growth on
progesterone-stimulated uterine secretion of

embryotrophic factors, it is not surprising that

progesterone supplementation should be so effective. Although a 10% improvement from early supplementation may seem a small gain, it is worth
noting that this would return conception rates to
those considered normal 10 years ago; in other
words, it would buy 10 years in the race against
subfertility. Manipulation of circulating progesterone concentrations during early pregnancy has
resulted in increased rates of conceptus survival in
sows in which lowering the plane of nutrition
reduces
hepatic
progesterone
catabolism
(Ashworth & Pickard, 1998).
The only theoretical disadvantage of administering progesterone is that it may reduce pituitary
gonadotrophin secretion, upon which the corpus
luteum depends; although this is not significant
while progesterone is being administered, it may
become so when the treatment is withdrawn if the
corpus luteum has been reduced in size or secretory capability during the period of administration.
In practice, this only occurs with relatively high
rates of progesterone administration and not with
the CIDRs currently in use for this purpose.
A second therapeutic approach to progesterone
supplementation is to block luteal regression by
reducing the luteolytic drive associated with follicular oestrogen secretion. This is the principle underlying use of Buserelin (Receptal, Hoechst Roussel
Vet Limited), a GnRH analogue which induces ovulation or atresia of follicles thus lowering circulating oestrogen concentrations (Mann & Lamming,
1995; Beck et al., 1996). This approach is based on
the fact that uterine secretion of prostaglandin F2,
which causes luteolysis, is stimulated by circulating
oxytocin, and the uterine expression of the oxytocin receptor is triggered by oestrogen. Reduced
follicular secretion of oestrogen leads to reduced
uterine secretion of prostaglandin F2 (Mann &
Picton, 1995). Treatment at first service post partum
with GnRH given on the day of insemination
caused a 6% improvement in pregnancy rate, and
later administration (days 1113 after service) produced a greater increase (1012%; Mee et al., 1990;
Peters et al., 1992).
The third therapeutic strategy should be based
on treating animals that ovulate but fail to show
oestrus. It has been estimated recently using a combination of milk progesterone analysis and Kamar
heatmount detectors (Cox Surgical) that 72% of
first ovulations post partum are silent (i.e. not
accompanied by oestrus), this percentage falling to

CATTLE FERTILITY

an average of 25% at subsequent ovulations (M.

Royal, unpublished data). This illustrates that, in
practice, failure to observe oestrus can be detected
by milk progesterone analysis, which will show ovarian activity in the absence of insemination.
Underlying endocrine causes of undetected
oestrus may include absence of ovarian function,
delayed ovulation (DOV) resulting in various types
of ovarian cyst or PCL. Alternatively, there may be
normal ovarian activity but a poor behavioural
response to the endocrine triggers responsible for
the onset of heat. These conditions are commonly
treated by administering progesterone via a CIDR
and/or prostaglandin F2 analogue. In the case of
PCL or a luteal cyst this treatment induces luteolysis and ovulation. Where there is normal ovarian
activity but failure to show heat, progesterone treatment and withdrawal may enhance the behavioural
manifestation of oestrus. Progesterone administration also has the advantage that it induces follicular
development and ovulation in the case of prolonged post partum anoestrus.
In view of the availability of treatments for some
of the causes of failure to show oestrus, it would be
appropriate to develop optimized therapeutic
protocols. These should be based on strategic
progesterone monitoring. Application of Kamar
heatmount detectors to all cows not observed in
heat by day 35 or 40 will also give valuable information as to an animals endocrine and behavioural
status. Although it is not possible to predict precisely the benefits of these treatments, it may be
that an optimized combination of treatments
would raise oestrus detection considerably.
NUTRITION
In addition to endocrine and genetic approaches
to cattle subfertility, the nutritional approach also
offers considerable potential. High-yielding dairy
cows are selected for an ability to partition energy
into milk production at the expense of body fat
early in lactation, and a cows condition score during lactation is heritable (Coffey, 1998; Jones et al.,
1999). As a result, dairy cows enter a state of negative energy balance at a time when they are
expected to return to ovarian cyclicity. The normal
development of the ovarian follicle involves production both locally and systemically of growth factors, in particular insulin-like growth factor I
(IGF-I; Guidice, 1992; Jones & Clemmons, 1995).
Hepatic IGF-I production (which determines

57

systemic levels of IGF-I) is dependent on growth

hormone. But systemic IGF-I levels fall under these
conditions although GH levels are raised (McGuire
et al., 1995) because undernutrition or a state of
negative energy balance lowers insulin secretion,
which reduces hepatic growth hormone receptor
levels (Pell & Bates, 1990). The local availability of
IGFs also declines, through effects on levels of IGF
binding proteins (IGFBPs). In the bovine follicle
approaching ovulation, the expression of the gene
for one of these IGFBPs (IGFBP-2) decreases,
allowing IGF to exert its effect of increasing granulosa cell growth (Armstrong et al., 1998). The
expression of IGFBP-2 is reduced by insulin, so a
high circulating level of insulin effectively increases
availability of IGF within the follicle, and a low
insulin level decreases it. As a result of these systemic and local relationships between metabolic
hormones and growth factors, milk yield compromises follicular growth (Webb et al., 1999). This is
borne out by the positive relationship between the
change in condition score during early lactation
and CLA (Canfield & Butler, 1990; Beam & Butler,
1997). Regression analysis shows that CLA is
unfavourably correlated to milk, fat and protein
yields, whereas energy balance, liveweight during
lactation and weight gain all have a favourable
genetic correlation with CLA (range = 0.40 to
0.80; Veerkamp et al., 2000).
Changes in systemic IGF-I levels may alter the
function of the uterine IGF system in addition to its
effects on the ovary. The endometrium produces
IGF-II during early pregnancy in ruminants, but
IGF-I in the uterus is probably derived from the circulation (Wathes et al., 1998). The endometrial
glands express IGF receptors during pregnancy, so
IGF-I in the uterus may affect uterine glandular
secretion of embryotrophic factors. IGFBPs are
also expressed in the uterus, and expression of at
least one of these proteins (IGFBP-3) is reduced in
the bovine uterus in the presence of a conceptus.
The operation of the IGF system in both the ovary
and the uterus therefore points to the benefits of
maintaining high IGF-I levels. If a nutritional
regime could be devised that would maintain high
insulin levels at this critical time of lactation,
thereby maintaining hepatic expression of the
growth hormone receptor and IGF-I secretion, the
adverse effects of negative energy balance on fertility may be overcome.
A second connection between diet and fertility
involves circulating concentrations of ammonia

58

THE VETERINARY JOURNAL, 160, 1

and/or urea. The rapidly digestable diets fed for

maximum yield contain high levels of rumen
degradable protein. This results in conversion of
dietary protein into ammonia and urea. In some
cases ammonia is produced at rates exceeding
hepatic clearance to urea. As a result, these diets
can lead to raised circulating levels of ammonia
and urea, which have deleterious effects on the follicle, the embryo and the uterus (Butler et al., 1996;
McEvoy et al., 1997; Sinclair et al., 1999). One
approach to improving fertility is therefore to provide a diet that raises insulin and lowers ammonia/urea during a critical period of follicular
development. However, since follicular development from the preantral stage to ovulation takes 4
months, it is not certain at present when such a diet
could be fed without compromising milk yield.
Further research is urgently needed in this area.
Experiments with high-producing (top 5% UK
herd) and low-producing (average for UK herd)
dairy cows show that in both groups, animals are
lactating to about 75% of their metabolic capacity
(Knight, 1999). That is, if they are given experimental galactopoietic stimuli, such as frequent
milking, recombinant bovine somatomammotropin (BST) and thyroxine, cows from both high
and low genetic merit lines will increase their yield
by 25%. However, when stimulated in this way, ovarian cyclicity (and in fact all follicular development)
ceases (Webb et al., 1999), and milk production
soon drops. It seems therefore that within the 25%
unused metabolic capacity, there is a trade off
between reproduction and yield. One aim of nutritional research should be to find how to maintain
yield without compromising fertility; alternatively,
we must accept that a high peak yield is not always
compatible with a short calving interval.
STRATEGIES
The three approaches to treating dairy cow subfertility, through endocrinology, nutrition and
genetics, are likely to be effective within different
timescales. The strategy for reversing the trend in
dairy cow subfertility must clearly include all the
cost-effective options open to us. In the short-term,
endocrine therapies should be introduced immediately. Although their widespread use may be criticized on welfare grounds in the long-term (since
they involve administering hormones), they will
provide immediate gains. There will always be a
proportion of problem cows that require

endocrine treatments, although in continuing to

breed from these animals the potential inheritance
of reproductive abnormalities should be borne in
mind.
Whilst endocrine therapy can be used now, the
nutritional and genetic approaches are likely to
take some time to develop. The genetic approaches
are the most long-term strategy, largely because of
the lack of and cost of collecting substantial quantities of objective and reliable data to analyse. There
is however one approach to genetic improvement
of fertility which may be introduced relatively soon.
Using milk progesterone data, Veerkamp et al.
(1998) and Darwash et al. (1999) suggested that it
may be possible to exploit the heritability of CLA
and the relationship between CLA and fertility, to
develop a selection index that could be incorporated into progeny testing for bulls. These methods
could exploit the fact that large numbers of daughters are tested per sire for milk quality and yield. If
the milk samples collected were also analysed for
progesterone at monthly intervals or during a critical interval post partum (e.g. 1725 days post partum), a bull could be assigned a breeding value for
female fertility, this being estimated from the average fertility of his daughters, since fertile daughters
ovulate earlier post partum than those that are less
fertile (Darwash et al., 1997b). The advantage of
this approach is that it could be implemented at
the time of progeny testing, using milk samples
obtained for fat and protein analysis; the information would then be available at the same time as
milk data upon which the value of the sire is currently based. Success of this strategy would require
breeding companies to develop a market for semen
of bulls with high male but more specifically high
female breeding values for fertility. However, this is
largely dependent upon the farmer perceiving an
advantage in paying a premium for female fertility.
Once bulls with high breeding values for female
fertility are identified and their semen marketed,
genetic improvements in female fertility could
potentially begin, although the response to selection depends upon the weighting applied to fertility within the selection index, the selection
intensity imposed on both the sire and the dam,
the heritability of CLA (which is moderate) and the
long generation interval of the cow.
We feel that fertility traits, preferably those free
of management bias, should be incorporated into
progeny testing with a minimum of delay. Bearing
in mind that the relationship between milk yield

CATTLE FERTILITY

and fertility may be genetically antagonistic, this

should be with appropriate economic weighting. A
programme to raise awareness of fertility problems,
within the industry, should be commenced immediately, in order to develop a market for semen
from sires with fertile daughters. In the mediumterm, nutritional approaches should be exploited.
These will involve further research into the compositions of diets providing fertility benefits, and
when they should be fed. Lastly, a long-term
genetic approach should be developed, based on
use of molecular markers in marker-assisted selection protocols and introgression of genes from
more fertile breeds. This will allow exploitation of
the heritable fertility traits revealed by milk progesterone analysis (see above). Opportunities for
development of markers are now (or will shortly
be) offered by the current advances in DNA arraying technologies and the bovine genome mapping
programme. Furthermore, basic research on the
physiological processes involved in fertility and subfertility will lead to the identification of candidate
genes which may also be used as markers. This may
be some way in the future, since it will be 510 years
before markers become available, but as with all
these options, it is vital for the future health of the
UK dairy industry that we find the resources
required to support their development now.

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