Animal Reproduction Science 96 (2006) 282296

The effect of nutritional management of the

dairy cow on reproductive efficiency
James F. Roche
UCD, School of Agriculture, Food Science and Veterinary Medicine, College of Life Sciences,
University College Dublin, Beleld, Dublin 4, Ireland
Available online 3 August 2006

Abstract
The cause of low fertility in dairy cows is multifactorial. Poor nutrition during the dry and early
postpartum periods results in reduced glucose, insulin, insulin-like growth factor (IGF-I) and low LH
pulse frequency with concomitant increases in -hydroxy butyrate, non-esterified fatty acids (NEFA) and
triacylglycerol. Cows must mobilize large lipid, but also some protein reserves, with a consequent increased
incidence of such metabolic disorders as hypocalcaemia, acidosis, ketosis, fatty liver and displaced
abomasums. The occurrence of milk fever and ketosis affects uterine contractions, delays calving and
increases the risk of retained foetal membranes (RFM) and endometritis. The nutritional risk factors that
cause RFM are hypocalcaemia, high body condition score (BCS) at calving and deficiencies in Vitamin E
and selenium. The risk factors for endometritis are hypocalcaemia, RFM, high triacylglycerol and NEFA.
Thus, metabolic disorders predispose cows to gynaecological disorders, thereby reducing reproductive
efficiency. Cows that are overconditioned at calving or those that lose excess body weight are more likely to
have a prolonged interval to first oestrus, thereby prolonging days open. Nutritionally induced postpartum
anoestrus is characterized by turnover of dominant follicles incapable of producing sufficient oestradiol to
induce ovulation due to reduced LH pulse frequency.
High nutrition can also increase metabolic clearance rate of steroid hormones such as progesterone or
oestradiol. Lower concentrations of oestradiol on the day of oestrus are highly correlated with the occurrence of suboestrus, thereby making the detection of oestrus in high yielding cows even more difficult.
Nutrition also affects conception rate (CR) to AI. Cows that develop hypocalcaemia, ketosis, acidosis or
displaced abomasums have lower CRs and take longer to become pregnant. Excessive loss of BCS and
excess protein content of the ration can reduce CR while supplemental fats that attenuate the production of
F2 can improve CR. The increased metabolic clearance rate of progesterone (P4), which decreases blood
concentrations during early embryo cleavage up to the blastocyst stage is associated with decreased CRs.

This paper is part of the special issue entitled Nutrition and Fertility in Dairy Cattle, Guest Edited by A. Evans and
F.J. Mulligan.
E-mail address: james.roche@ucd.ie.

0378-4320/$ see front matter 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.anireprosci.2006.08.007

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In conclusion, poor nutritional management of the dairy cow, particularly before and after calving, is a key
driver of infertility.
2006 Elsevier B.V. All rights reserved.
Keywords: Fertility; Dairy cow; Metabolic disorders; Negative energy; Body condition; Retained placenta; Anoestrus;
Conception rate; Endometritis; Progesterone; LH; Transition period; Postpartum; Reproductive management

1. Introduction
The cause of low fertility in the modern Holstein Friesian cow is multifactorial involving genetic
improvement, inadequate nutrition, poor reproductive management, an increased incidence of
disease and overall poor cow welfare (Lucy, 2001). The partitioning of the relative impact of
these various factors on infertility is not well understood. It is also clear that there are interactions
between these factors and different contributions from individual farm factors depending on
the specific farm management strategy. It is becoming increasingly clear that good reproductive
management is dependent on proper attention to the optimum nutrition of the cow, whose nutrient
requirements vary depending on physiological state and the specific nutrient demands to prevent
metabolic disorders in the peri-parturient period (Boland et al., 2001; Overton and Waldron, 2004).
Thus, increasingly emphasis must be placed on the nutritional management of the cow before
and during the dry period, the length of the dry period and the actual body condition score (BCS
on a scale of 15) targets and rate of BCS loss to be met. The underlying theme of this paper
is that poor nutritional management of the cow, particularly before calving, is the key driver of
low reproductive efficiency in high yielding dairy cows, when fed to allow her meet her genetic
potential for milk production.
High reproductive efficiency is necessary for efficient milk production and it, therefore, has
an important influence on herd profitability (Pryce et al., 2004). Low reproductive efficiency
decreases herd profitability by: (i) prolonging the calving interval, which results in less milk
produced per cow and fewer calves born per year; (ii) increasing culling due to infertility and
therefore, increased replacement costs; (iii) increased labour, semen costs and veterinary bills;
(iv) an extended low production or dry period can result in over conditioned cows calving in too
high a BCS (>3.5) which results in a subsequent prolonged period of negative energy balance
(NEB) and low reproductive efficiency.
The two key components of high reproductive efficiency are:
High submission rate of cows for insemination.
High conception rate to each service.
Submission rate is defined as the percent eligible cows (e.g. calved >42 days so that the majority
should be cyclic) presented for artificial insemination (AI) in a 24-day period (representing an
oestrous cycle). The two key factors that affect it are:
Percent cows that are anoestrus.
Heat detection efficiency in the herd.
The target is to achieve >80% of cows inseminated in a 24-day period of the breeding season.
Therefore, to obtain high submission rate to AI, it is necessary to obtain proper uterine involution
by day 50 or so postpartum and get >90% of cows resuming cyclicity with normal oestrous cycles

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Table 1
The postpartum (pp) reproductive targets to be met to obtain high reproductive efficiency and the associated key risk
factors affecting these targets
Reproductive process

Target to be achieved

Risk factors affecting targets

Normal uterine involution

Day 50 pp

Dystocia
RFM
Uterine infection

Resumption of ovulation

90% by day 42

Loss of > 0.5 BSC unit

Low feed intake
Uterine health

High oestrous detection

85% per cycle

Infrequent checks
Sub-oestrus
High yield

High conception rate to AI

50% per breeding

Excess BCS loss

Prior uterine problems
Low P4 days 47 of pregnancy

before breeding. The postpartum reproductive targets to aim for are presented in Table 1. Thus high
reproductive efficiency is dependent on obtaining normal uterine involution, early resumption of
ovulation, high efficiency of oestrous detection and high conception rates per service.
2. Uterine involution
Following calving, there is a rapid decrease in size and weight of the uterus due to vasoconstriction and reduction in size of the cotyledons within 9 days (Gier and Marion, 1968). There is
dissolution and detachment of superficial layers of the cotyledons with increased cellular debris
and fluid production. Most of the necrotic layer has been removed by 1014-day (Gier and Marion,
1968). Peristaltic contractions of the uterus continue for the first few days, presumably under the
influence of the prostaglandin (PG) F2, which remains elevated, but declines over the next 20
days after calving (Lindell and Kindahl, 1982). Changes in the cervix resulted in decrease in size
and cervical diameter from about 15 cm after calving, to 911, 78 and 56 cm by days 10, 20
and 60 after calving (Gier and Marion, 1968). Physical involution, i.e. return to its normal nonpregnant size is completed by 3040 days post-calving but the endometrium may not be capable
of normal embryonic development and maternal recognition of pregnancy until day 60 or so after
calving. The key factors that affect progression of involution are retained placenta, dystocia and
the occurrence of metabolic disorders such as milk fever (Gier and Marion, 1968; Marion et al.,
1968; Morrow et al., 1966). Therefore, delayed uterine involution will reduce reproductive efficiency. There are no effective ways to speed it up although some authors use multiple injections
of PGF2 in the early postpartum period. Results of its efficiency are not clear cut and this is
partially related to the high but declining concentrations of PGF2 in the early postpartum period
(Lindell and Kindahl, 1982) and the short half life of PGF.
Postpartum uterine health plays a central role in determining the reproductive efficiency of dairy
herds. Postpartum local infections of the uterus delay uterine involution, cause inflammation of
the endometrium, may delay the interval to first ovulation postpartum, reduce conception rate
(CR) to first insemination and increase the risk of culling for infertility (Sheldon, 2004). Uterine
problems can also have a negative effect on dry matter intake of cows, cause poor welfare and
increased stress, probably resulting in high cortisol secretion (Wischral et al., 2001).

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2.1. Retained foetal membranes

Retained foetal membranes occur due to any interference with the normal detachment of
cotyledons from the caruncle resulting in its failure to be expelled within 12 h of calving. The
enzymes involved in tissue remodelling such as collagenases and proteases play an important role
in concert with myometrial contractions in the early postpartum period. Decreased neutrophil
function, manifest as reduced placental attraction, is implicated as a primary cause (Kimura et al.,
2002). Interleukin-8, a potent chemoattractant and the immune status were reduced in cows that
develop RFM. Increased metabolic stress could be implicated in the earlier but reduced synthesis
of cortisol, the reduced oestradiol and prostaglandin PGE and F metabolites before parturition
in cows with RFMs (Wischral et al., 2001). RFM delay uterine involution, predispose cows to
endometritis or metritis and decrease fertility (Grohn and Rajala-Schultz, 2000; Maizon et al.,
2004). Nutrition plays an important role in causing RFM. The factors involved are: high BCS
at calving, dietary deficiencies of Vitamins A, D and E and deficiencies in selenium, iodine and
perhaps zinc (Laven and Peters, 1996; Gupta et al., 2004; Han and Kim, 2005). Hypolcalcaemia is
also a key risk factor for RFM (Houe et al., 2001). Thus, maintenance of normal uterine physiology
by good nutritional management during the dry and transition periods is important to reduce the
incidence of RFM, which is an important risk factor for endometritis. Hence, its occurrence has
a negative effect on reproduction.
2.2. Endometritis
Endometritis is a superficial inflammation of the endometrium without systemic or overt signs
of illness that often occurs before first ovulation (Sheldon, 2004). The presence of pathogens trigger an innate immune response including neutrophil migration, cytokine release, macrophage and
serum compliment increases to overcome the local infection. The uterus of most cows is exposed to
bacterial contamination after calving; the proportion of uteri contaminated with bacteria declined
from 78% by days 1630 to 50% by days 3145 and 9% by days 4660 postpartum (Sheldon,
2004). The problem arises where pathogenic bacteria multiply due both to the suppressed immune
system around parturition and the resultant decrease in migration and the functional capacity of
neutrophils to eliminate uterine bacteria. The incidence of endometritis is affected by the method
used to diagnose it because of perceived insensitivity and lack of specificity of rectal examination
of the uterus and observation of a genital discharge (Gilbert et al., 2005; Sheldon et al., 2006).
Current evidence suggests the most efficacious methods for its diagnosis are the presence of a
muco-purulent discharge following vaginoscopy, the presence of an enlarged cervix and/or the
presence of inflammatory cells in uterine aspirates. The prevalence of endometritis varies with
method of diagnosis used, production system employed, yield and parity of cows being examined. The incidence of clinical/subclinical endometritis in Canada, Ireland and the US, has been
reported as 17% of 1865 cows, 24% of 6500 cows and 54% of 141 cows, respectively (Mee and
Buckley, 2003; LeBlanc et al., 2002; Gilbert et al., 2005). It causes damage to the endometrium,
histological lesions, and may delay uterine involution.
The consequences of it on reproductive efficiency are partly dependent on when it occurs, its
severity in the herd and the prevalence of the key risk factors on a herd basis. However, its incidence
is such that it can have serious negative effects on herd reproductive efficiency. It can decrease
submission rate by: prolonging the interval to first ovulation, reducing the growth rate of the first
postpartum dominant follicle and decreasing the secretion of oestradiol from dominant follicles
in the early postpartum period (LeBlanc et al., 2002; Sheldon and Dobson, 2004). This, therefore,

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increases the chances that the dominant follicles that develop will produce insufficient oestradiol
to induce GnRH release, the LH/FSH preovulatory surge and ovulation, with a consequent delay
to first ovulation. The production of endotoxins can also suppress LH pulse frequency, which will
reduce oestradiol secretion from dominant follicles thereby increasing the chances of dominant
follicle artesia, rather than ovulation. Endometritis can also have detrimental effects on first service
conception rates if it occurs between 20 and 33 days postpartum (LeBlanc et al., 2002) and it
increases the number of inseminations per pregnancy (Gilbert et al., 2005). Thus, it will prolong
the calving to first service interval, the calving to conception interval, days open and increase the
risk of cows being culled for infertility.
There are many risk factors for the establishment of endometritis previously reviewed (Sheldon
and Dobson, 2004; Sheldon, 2004). Those that relate to nutritional involvement are RFM, high
triacylglycerol concentrations associated with liver disease (Zerbe et al., 2000), overconditioned
cows at calving, high non-esterified fatty acid levels as an indicator of NEB (Jorritsma et al.,
2003) and hypolcalcemia (Whiteford and Sheldon, 2005). The incidence of metabolic disorders
is inter-related and the occurrence of one can predispose cows to related disorders (Laven and
Peters, 1996). Milk fever and ketosis affect uterine muscle contractions, delay the process of
calf delivery and also the ability of the uterus to expel the placenta within 12 h of calving. Thus,
endometritis is prevalent in the postpartum period, its incidence is affected by nutrition and it is
a serious cause of reduced reproductive efficiency in the modern Holstein Friesian dairy cow.
3. Resumption of ovulation and oestrus
Following calving, the negative feedback effects of the high steroid concentrations in late
pregnancy on gonadotrophin releasing hormone (GnRH) are removed. This is followed by an
increase in FSH concentrations for 35 days between days 7 and 14 postpartum (Fig. 1). The
increase in FSH results in the emergence of the first postpartum follicle wave (Savio et al., 1990a).

Fig. 1. Follicular dynamics, ovulation and concentrations of FSH, progesterone, oestradiol (E2) and LH pulse (in panels)
in cows that resume ovulation early postpartum (cyclic cow) or in those that fail to ovulate (anoestrous cow) within 3040
days after calving.

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A cohort of oestrogen active antral follicles is detectable by ultrasound scanning on the ovaries.
The follicles produce both oestradiol and inhibin, which in turn suppress FSH secretion. Thus,
the growing follicles are now faced with a decline in FSH. However, one follicle (generally the
largest of the cohort) develops increased amounts of LH receptors and insulin-like growth factor
(IGF-I) binding protein proteases, which allow the maintenance of high levels of bioactive IGF-I
by degrading the IGF binding proteins. Thus, this follicle is selected to continue growing due to
these local paracrine changes within the follicle in spite of declining systemic concentrations of
FSH, which prevents other FSH-dependent cohort members developing. This dominant follicle
(DF) is now mainly LH-responsive and thus, its fate is determined by the LH pulse frequency it
is exposed to at this time. The fate of the first postpartum DF based on recent research (Savio
et al., 1990b; Beam and Butler, 1997; Sartori et al., 2004; Sakaguchi et al., 2004) is one of the
following:
it ovulates in 3080% of cows;
it undergoes atresia in 1560% of cows;
it becomes cystic in 15% of cows.
The occurrence of the first transient 34 days increase in FSH concentrations is not affected
by plane of nutrition or presence of a calf (Stagg et al., 1998). The incidence of anoestrus in dairy
cows based on twice weekly assay of progesterone in milk has increased from 7% in 1986 (Fagan
and Roche, 1986) to 1122% in UK and Netherlands (Lamming and Darwash, 1998; Opsomer et
al., 2000).
There are significant metabolic changes associated with parturition and early lactation. The
transient but crucial period of NEB in the early postpartum period induces significant metabolic
changes, which have a negative impact on reproductive efficiency (Jorritsma et al., 2003). The
ability of each cow to adapt physiologically to these metabolic changes and limit the amount of
body tissue mobilization has an important influence on inherent cow fertility (Jorritsma et al.,
2003). Following calving, concentrations of growth hormone (GH) are increased which induces
lipolysis and suppresses peripheral tissue insulin responsiveness. The increased GH induces a
catabolic state and results in loss of BCS and weight of cows (Lucy, 2003). There are also reduced
GH receptors in the liver, which results in declining IFG-I concentrations despite the increased GH
concentrations in the early postpartum period (Vandehaar et al., 1995; Lucy, 2001). This decline in
IGF-I is an important indicator of nutritional status on the hypothalamic-pituitary-ovarian axis in
the early postpartum period. It has been hypothesized that measurement of serum concentrations
of IGF-I could be a useful predictor of nutritional status and hence reproductive efficiency in dairy
cows (Zula et al., 2002a).
3.1. First ovulation
The key factors determining whether or not the first DF ovulates are: size of the DF, the LH
pulse frequency it is exposed to and the systemic concentrations of IGF-I. Follicles less than
1 cm in diameter rarely ovulate. The LH pulse frequency required for ovulation is about 1 pulse
per hour. This pulse frequency ensures there is a significant increase in oestradiol concentrations
necessary to induce positive feedback and ovulation to occur. If the first postpartum DF ovulates,
oestrous behaviour is not generally expressed and the ovulation is silent (Kyle et al., 1992). This
is because ruminants require prior exposure to progesterone in order for them to express oestrous
behaviour in response to the pre-ovulatory increase in oestradiol (Kyle et al., 1992) that causes

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first postpartum ovulation. Thus, it is necessary to pre-treat anoestrous cows with progesterone in
order to increase the chances of behavioural oestrus being shown at first ovulation. In addition, the
length of the first luteal phase is also often short, due to the premature release of PGF2 arising
from the increased oestradiol produced from the formation of the post-ovulatory DF on days 58
of the cycle. Thus, the CL regresses prematurely at days 810 of the cycle and the DF from the
first post-ovulatory follicle wave ovulates around days 911 after the first ovulation. The second
ovulation is generally accompanied by oestrous behaviour and a normal length luteal phase. Prior
exposure to progesterone also plays a key role in determining whether or not a short cycle occurs.
Treatment of anoestrous cows with progesterone and oestradiol induces oestrous and shortens the
postpartum interval to conception (Rhodes et al., 2003).
Cows can have two, three or four follicle waves during the oestrous cycles that recur in the
early postpartum period (Savio et al., 1990a; Sartori et al., 2004). Holstein cows tend to have
two follicle waves per 1823 days cycle (Sartori et al., 2004). Certain factors that regulate LH
pulse frequency such as progesterone concentration may affect the duration of dominance of the
sequential DFs that develop during the luteal phase of the oestrous cycle. Holstein cows have lower
concentrations of progesterone during the cycle than cyclic heifers (Sartori et al., 2004; Wolfenson
et al., 2004). Thus, the lower concentrations of progesterone may allow a subtle increase in LH
pulse frequency and allow the DF to continue to grow rather than undergoing atresia, thereby
preventing the development of a third follicle wave. Cows with longer than normal luteal phases
can have a fourth follicle wave (Savio et al., 1990a). Follicle waves also continue during early
pregnancy (Savio et al., 1990a). The number of follicle waves will affect the duration of dominance
of the DF and, in general, the shorter the duration of dominance of the pre-ovulatory follicle, the
higher conception rate to AI will be, where all other factors are consistent (Austin et al., 1999).
Thus, nutrition by altering metabolic clearance rates of progesterone can affect the number of
follicle waves per cycle, and hence, indirectly, conception rates.
3.2. Failure of rst postpartum DF to ovulate
The factors that influence whether the first postpartum DF ovulates are BCS, yield, parity
season and disease (Beam and Butler, 1997; Stagg et al., 1998). Cows that calve down in poor
BCS (<2.5) are more likely to have a prolonged anoestrous period (Fig. 1) due, presumably, to low
LH pulse frequency and subsequent reduced concentrations of oestradiol, which are ineffective
to induce an LH surge and ovulation. Cows in poor BCS after calving have decreased diameter of
the DF, reduced insulin and IGF-I concentrations and low LH pulse frequency. Current evidence
suggests cows should calve down in a BCS of 2.753 and not lose more than 0.5 of a unit of BCS
between calving and first service (Overton and Waldron, 2004) rather than the previous higher
recommendations of 33.5 (Buckley et al., 2003). Cows that lose 1 unit BCS have a longer
postpartum interval to first ovulation. Thus, monitoring of BCS score from calving to first service
is an important aspect of good reproductive management.
Current Holstein cows produce very high yields of milk. This high yield in the early postpartum
period forces the cow to mobilize mainly fat but also protein to meet the energetic demands of
milk production. Cows can lose 5075 kg body weight. It is necessary to prevent a steep decline
in NEB and to shorten its duration postpartum by increasing dry matter intake (DMI) in the early
postpartum period. Cows that are mobilizing tissue at a high rate have increased blood concentration of non-esterified fatty acids, -hydroxy butyrate, triacylglycerol but reduced concentrations
of insulin, glucose and IGF-I (Grummer et al., 2004). This metabolic status increases the risk
of hypocalcaemia, acidosis, ketosis, fatty liver and displaced abomasums (Grohn and Rajala-

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Schultz, 2000; Overton and Waldron, 2004; Maizon et al., 2004). The occurrence of subclinical
hypocalcaemia increases the risk of reproductive problems associated with parturition viz such
as dystocia and retained placenta while the occurrence of metabolic disorders after calving can
affect uterine functions viz endometritis. Cows thus affected are more prone to mastitis, lameness,
anoestrus and reduced conception rate to AI (Fourichon et al., 1999; Grohn and Rajala-Schultz,
2000; Lucy, 2001; Lopez-Gatius et al., 2005; Maizon et al., 2004). This, in turn, means these cows
are more likely to be culled for infertility. Therefore, high yielding cows must be nutritionally
managed to ensure only a mild, but not severe, NEB deficit post calving occurs, have adequate
Ca mobilization in the peri parturient period and are given the appropriate vitamin and mineral
allowances, particularly selenium, iodine, vitamins A, D and E to maintain their health, welfare
and reproductive status. Current strategies to mimimize BCS loss involve keeping cows in BCS
2.753.00 at calving, shortening the dry period and maintenance of normal rumen function.
3.3. Formation of a cystic follicle
These occur when the first DF that develops in the early postpartum period fails to ovulate
and continues to grow to diameters >25 mm over a 1040 days period in the absences of a corpus
luteum (CL) in the dairy cow (Savio et al., 1990a; Gumen et al., 2002; Hatler et al., 2003). The
continuation of the growth of the cyst seems to be due to lack of positive feedback induced by
oestradiol and thus failure of occurrence of an LH/FSH pre-ovulatory surge. Thus, progesterone
concentrations are low but oestradiol concentrations are elevated much above normal proestrous
concentrations (Savio et al., 1990b; Hatler et al., 2003). This elevation in oestradiol affects their
behaviour, and cows with follicular cysts tend to be members of the sexually active group, and
thus can facilitate oestrous detection. The secretion of high concentrations of oestradiol from the
cyst and also presumably inhibin, result in the suppression of FSH recurrent increases; hence no
new follicle wave emerges during this period. It then appears that the cyst becomes oestrogen
inactive while still morphologically present and still detectable by ultrasound examination. This is
followed by new follicle wave emergence and subsequent ovulation often when the original large
physiologically active cyst is still present. This difference in functionality is important in deciding
whether or not to treat a cow with a cystic follicle. Multiple cysts can develop simultaneously
and these are often accompanied by very high concentrations of oestradiol being produced during
their early physiologically active phase. Many cysts that occur in the early postpartum period self
correct but some persist and further cysts can develop which may be hard to eliminate. Cows with
ovarian cysts take 611 days longer to first service and 2030 more days to conception (Fourichon
et al., 1999). The metabolic risk factors involved in the pathogenesis of cysts are overconditioned
cows, a reduction in insulin and IGF-I and increased non-esterified fatty acids (Vanholder et al.,
2005; Zula et al., 2002b). Cows with an abnormal puerperium were 1.9 times more likely to
develop cysts while an increase of 1 unit of BCS increased the risk 8.4 times; thereby indirectly
implicating nutrition as a secondary causative factor (Lopez-Gatius et al., 2002).
3.4. Manifestation of irregular cycle lengths
Some cows have irregular cycles, either shorter or longer than 3 weeks in the absence of the
embryo. These irregular cycles generally relate more to premature or prolonged CL regression.
The risk factors associated with irregular cycles are dystocia, RFM, abnormal vaginal discharge
and metritis (Opsomer et al., 2000). Cows with atypical ovarian progesterone patterns before first
service had longer intervals to conception, a higher number of services per conception and lower

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first service pregnancy rates (Lamming and Darwash, 1998). The incidence of prolonged cycles
in non-pregnant cow has increased from 3% (Fagan and Roche, 1986) to 1122% (Lamming and
Darwash, 1998; Opsomer et al., 2000; Shrestha et al., 2004).
4. Detection of oestrus
The efficiency of detection of oestrus in cows at farm level is variable but often low (Stevenson,
2001). This is a major limitation to achieving high submission rates to A1. The problem is due
to the short duration of oestrus and exacerbated by the occurrences of low intensity oestrous
behaviour. Continuous monitoring of dairy cows using pressure activated radiotelemetric devices
showed that the duration of oestrus was 7.14 h (range of 5.510.6 h) with cows being mounted 8.5
times per oestrus (range of 6.412.8 times) (Dransfield et al., 1998). Similar results were reported
by Xu et al. (1998) for dairy cows at pasture. The intensity of expression of oestrus is affected
by housing, floor surface, yield, lameness and number of herd mates in oestrus simultaneously
(Diskin and Sreenan, 2000). High ambient temperatures causing heat stress result in physical
lethargy and reduced oestrous detection efficiency in cows (Peralta et al., 2005). Cows with high
milk production (40 kg/day) have shorter duration of oestrus (6.2 0.5 h versus 10.9 0.7 h),
less total mounts per cow (6.3 0.4 s versus 8.8 0.6 s) and overall shorter duration of total time
standing to be mounted (21.7 1.9 s versus 28.2 1.9 s) than lower producing cows measured at
the same postpartum interval (Lopez et al., 2004). Moreover, the high producing cows have more
low intensity/short duration oestrus periods (53%) compared with low producing cows (32%).
In Japan, it has been reported that 33% of cows apparently in oestrus, based on expression of
secondary oestrous behavioural signs, were not detected to stand to be mounted when checked
every 4 h (Yoshida and Nakao, 2005). Thus, as production levels of cows continue to increase,
due to improved genetics and nutritional management, the duration and intensity of expression
of oestrus have been reduced, thereby exacerbating the problems of oestrous detection, often in
association with reduced labour input and sometimes associated with reduced technical knowledge
of the stockmen hired.
The physiological causes of these detrimental effects of high milk production on duration and
expression of oestrus are in part related to a reduction in the concentrations of oestradiol on the day
of oestrus (Lyimo et al., 2000). The increased milk production was associated with a corresponding
increase in DMI, which increased liver blood flow and metabolic clearance rates of the key steroid
hormones, progesterone and oestradiol (Sangsritavong et al., 2002). It is hypothesized that this
increased metabolic clearance rate of the key reproductive steroid hormones is a major cause of
the reduced oestradiol concentrations on the day of oestrus and thus it decreases the probability
of detecting oestrous as DMI intake increases in high producing cows. Furthermore, heifers have
higher concentrations of oestradiol than dairy cows (Sartori et al., 2004; Wolfenson et al., 2004) on
the day of oestrus. The changes in peripheral steroid hormones during the oestrous cycles of high
producing cows induced by high metabolic clearance rates of the key gonadotrophin regulating
steroid hormones could alter the follicular wave patterns, duration of dominance and the time of
ovulation relative to the onset of oestrus (Austin et al., 1999).
The optimum time to inseminate cows has been reconfirmed to be within 412 h of observation
of oestrus (Dransfield et al., 1998) since the cow generally ovulates 2428 h after the onset of
oestrus. Ovulation is induced by positive feedback of oestradiol on GnRH secretion, which results
in the pre-ovulation LH/FSH surges and ovulation. Thus, the attenuation of the oestradiol increase
on the day of oestrus could affect the timing of the LH/FSH surge, and hence the time of ovulation.
This could potentially affect the optimum time to inseminate cows with reduced concentrations

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of oestradiol on the day of oestrus. Acute reduction in the plane of nutrition of heifers attenuated
both the proestrous increase in oestradiol and the magnitude of the LH surges (Mackey et al.,
1999). There was a significant relationship between peak proestrous concentrations of oestradiol
and pregnancy rates with reduced pregnancy rates occurring in heifers with attenuated oestradiol
increases. This again highlights that nutrition can affect, indirectly, oestradiol concentrations on
the day of oestrus, which, in turn, can affect the duration and intensity of oestrous behaviour and
perhaps also fertility by affecting the time of ovulation in the cow.
5. Conception rate to AI
The optimum time to inseminate the cow is 1218 h before ovulation using viable fresh or
frozen (thawed) semen. The fertile life span of bull spermatozoa is 2430 h (Dalton et al., 2001)
but the viable life span of the ovulated oocyte is only 612 h (Gordon, 1996). Breeding heifers
based on the use of secondary signs of oestrus such as triggered heat mount detector, non-standing
behavioural signs based on mounting and chin resting, or rectal palpation, resulted in significantly
reduced conception rates (Donovan et al., 2003). The nutritional factors that affect conception
rates include.
5.1. Disease status
Cows that develop hypocalcemia, ketosis, acidosis or displaced abomasum can have lower CR
to first service, require more inseminations per pregnancy, or take longer postpartum to become
pregnant (LeBlanc et al., 2002; Kim and Kang, 2003; Gilbert et al., 2005). These metabolic
diseases are key risk factors for the development of gynaecological problems during or after parturition and in the early postpartum period during the critical period of uterine involution. The
key risk factors for delayed conception identified by Grohn and Rajala-Schultz (2000) were RFM,
metritis and cysts while Maizon et al. (2004) found that days from first breeding to conception
increased in cows with dystocia, stillbirth, RFM, metritis or ovulation dysfunction. Conception
rates were reduced by 14, 15 and 21%, respectively, in cows with RFM, metritis or cysts compared
with cows not diagnosed with these problems (Grohn and Rajala-Schultz, 2000). Other studies
show that delayed conception can be related to sub-clinical disease, stressful environmental, social
conditions and nutritional stress (Lopez-Gatius et al., 2005). Thus, it is clear that inadequate nutritional management of the cow in the dry period and after calving has a significant negative impact
on subsequent conception rate, services per conception and interval from calving to conception.
These effects of nutrition are manifest through different mechanisms including BCS changes,
increase incidence of metabolic disease, the resultant abnormal metabolic profiles precipitated,
the increased incidence of gynaecological problems with their attendant detrimental effects on
the uterus and the abnormal profiles of cytokines and endotoxins produced which can act locally
or systemically on the developing embryo.
5.2. Rate of BCS loss
Dry matter intake and energy balance during the dry period and post calving transitional
period have a crucial influence on herd health, productivity and reproductive efficiency. They
affect endocrine parameters, metabolic hormones and key concentrations of metabolic factors.
There is a normal decline in DMI prepartum in cows, as the foetus grows, which is exacerbated
in cows that subsequently develop metabolic or gynaecological problems at or after parturition

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(Grummer et al., 2004). Thus, excessive drop in DMI prepartum could serve as an early warning
signal of the propensity of cows to develop problems at or after parturition. Overconditioned cows
have increased incidence of peri- and post-parturient problems (Zula et al., 2002b; Lacetera et
al., 2005). However, it is becoming increasingly clear that massive increases in BCS followed
by abrupt loss in the early postpartum period are not well tolerated by the cow. Overconditioned
cows lose more BCS, have reduced DMI postpartum (Zula et al., 2002b; Jorritsma et al., 2003)
and can be more difficult to get in calf. Accordingly, it may be more important to precondition
the physiology and metabolism of the cow to moderate changes in NEB and minimal BCS loss
(<0.5 unit) in order to attenuate the massive changes in metabolites and metabolic hormones
that occur in the early postpartum period. Thus, minimizing BCS changes pre and post calving,
maintenance of optimal rumen function and prevention strategies to prevent metabolic disorders
are key management targets, in association with improved cow comfort and reduced stress, to
improve health, productivity and reproductive efficiency of high yielding dairy cows. Furthermore,
feeding high roughage diets at the start of the dry period to minimize BCS gain and maintaining
change in BCS score of 0.5 of a unit are increasingly important goals to achieve for high conception
rates and herd pregnancy rates (Overton and Waldron, 2004).
5.3. Dietary effects
It is important to supply cows with the specific amino acid requirements for milk synthesis,
reproduction and general body maintenance. These are derived from metabolizable proteins of
either microbial or dietary origin. The protein content of the diet affects milk production and
composition but many studies show that increasing the crude protein content of the diet reduces
fertility (reviewed by Laven and Drew, 1999) by reducing conception rates to service, particularly
in older cows, thereby prolonging days open. Some studies have shown that the degradability
of the protein affects fertility, but this is conflicting (Laven and Drew, 1999; Westwood et al.,
2000). Where excess protein is fed to cows in NEB, the energetic demands of excreting this as
urea may exacerbate the effects of the NEB on reproduction, thereby decreasing fertility. This
is supported by the fact that high crude protein diets with associated elevated urea levels, did
not affect embryonic survival in heifers (Kenny et al., 2002). The source of rumen undegradable
protein may be important. Where fish meal is used, improvements in fertility have been obtained
in some studies but it is not clear if this effect is due to its protein content or fatty acid profile
provided.
There is increasing evidence that the fatty acid composition provided by feeding supplemental
by-pass fats can alter the fatty acid profile in the blood of cows and increase linoleic acid (Thatcher
et al., 2006). This results in increased arachadonic acid levels, which in turn result in increased PGF
synthesis. PGF plays an important role in uterine involution, in upregulation of the immune system
and neutrophil function in the early postpartum period. Thatcher et al. (2006) have hypothesized
that a dietary strategy of feeding certain by-pass fats to increase PGF production in the uterus and
enhance immune competency is a novel approach to reduce uterine infections and assist normal
uterine involution in the transition period with consequent beneficial effects on fertility. The type
of fat fed during the breeding season should provide increased levels of n 3 fatty acids in order
to reduce PGF synthesis at the expense of increased PGE or other non-luteolytic PGs. This
reduced or delayed synthesis of PGF could be important in giving a longer period for maternal
recognition of pregnancy (MRP) to take place from smaller embryos producing less interferon tau
often in association with reduced progesterone concentrations before MRP (Mann et al., 2006).
The feeding of fish oil reduces uterine PGF synthesis and may benefit fertility in cows.

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293

6. Oocyte and embryo developmental capacity

A major reason for reduced conception rates in dairy cows is the increased embryonic mortality
prior to MRP (Sartori et al., 2004). This effect can be manifest as early as day 5 after AI. It is
not yet clear whether this increased loss is due to ovulation of an incompetent oocyte capable of
being fertilized but not able to develop or due to reduced embryonic development as a result of
oviduct-uterine environmental aberrations. It is clear that progesterone concentrations are reduced
by high DMI intake (Sangsritavong et al., 2002) and that embryonic loss is increased in cows
with reduced progesterone concentrations days 47 post-insemination due in part to the presence
of reduced interferon tau concentrations (Mann et al., 2006; Stronge et al., 2005). A reduction
in oocyte competency in high yielding Holstein Friesian cows has been reported (Snijders et al.,
2000; Leroy et al., 2005). Thus, it appears that both oocyte competency and embryonic survival are
compromised. Nutrition is also implicated in oocyte competency because during NEB, the levels
of non-esterified fatty acids are elevated which can compromise oocyte developmental capacity
(Leroy et al., 2005) in the early postpartum period.
7. Conclusions
Nutrition plays a pivotal role in determining fertility of postpartum cows. It affects submission
rate of cows to AI during the breeding season by increasing the incidence of anoestrus due to its
suppressive effect on LH pulse frequency, and reduced oestradiol production from the dominant
follicle which then undergoes atresia rather than ovulation. The incidence of anoestrus in dairy
cows has increased. The occurrence of metabolic disorders in their own right often delay the
interval to first detected oestrus as well as predisposing cows to gynaecological disorders such
as dystocia, RFM, endometritis and metritis. These disorders also reduce submission rate to AI.
Their incidence is also increasing. There appears to be an interaction between the occurrence of
metabolic diseases such as hypocalcemia, ketosis and fatty liver and an increased incidence of
dystocia, RFM and endometritis. Thus, prevention of metabolic diseases is one important component of attaining high reproductive efficiency in dairy cows and points to the interaction of
good nutritional management on fertility. Over conditioned cows (BCS >3.5) have reduced DMI
post calving, lose excess BCS post calving, have lower fertility and increased risk of developing
cystic follicles. Management of BCS is a critical component of nutritional management of cows
for high fertility. Major increases in or loss of BCS are undesirable. It is clear that optimum
reproductive efficiency is obtained when BCS loss is at or below 0.5 unit during the transition
period. To achieve this important target, cows should be maintained at a BCS of 2.53.0 in association with maintenance of proper rumen function through adequate dietary fibre, shortening the
dry period (68 weeks maximum), reduction in the incidence of subclinical metabolic disorders,
particularly hypocalcemia and minimizing mobilization of body reserves in the early postpartum
period.
Nutrition and dietary factors also affect conception rate and the number of services per conception. Cows that have had metabolic disorders and/or gynaecological problems are more likely
to have lower conception rates and to be culled due to infertility. The rate of BCS loss is also
a critical component with cows losing most BCS and in low BCS at time of AI having reduced
fertility. The energy status of the diet is, therefore, critical. However excess protein, particularly
in cows in NEB can reduce conception rates to AI. In addition the fatty acid profile of the diet,
the type of fat fed and the use of fish meal can affect uterine health and conception rates. Cows
with high metabolic loads will metabolize progesterone faster resulting in reduced progesterone

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concentrations during early embryonic development, thereby reducing conception rates due to
high embryonic loss in such cows.
In order to meet the challenge of increasing fertility of the modern dairy cow, genetic strain
selected, optimizing nutritional management, reduction in the incidence of metabolic and gynaecological disorders, and improved reproductive management in herds of increasing size with
reduced labour are all critical components of fertility. Thus, multi-disciplinary approaches using
the latest technology are fundamental to overcoming this major problem, which is a key component
of determining the level of productivity, profitability and cow comfort in the herd.

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