Body Condition and Suckling As Factor Influencing The Duration of Postpartum Anestrum in Cattle.

Animal Reproduction Science 85 (2005) 126
Body condition and suckling as factors influencing

the duration of postpartum anestrus in cattle:
a review
F. Montiel , C. Ahuja
Department of Reproduction, Faculty of Veterinary Medicine, University of Veracruz, Circunvalacin S/N
Esquina Yaez, Cdigo Postal 91710, Veracruz, Mexico
Received 21 March 2003; received in revised form 10 October 2003; accepted 14 November 2003
Abstract
Prolonged postpartum anestrus is a main factor limiting reproductive efficiency in cattle, partic-
ularly in Bos indicus and Bos taurus/Bos indicus cows from tropical regions, because it prevents
achievement of a 12 month calving interval. During anestrus, ovulation does not occur despite
ovarian follicular development, because growing follicles do not mature. Although many factors
affect postpartum anestrus, nutrition and suckling are the major factors influencing the resumption
of postpartum ovarian cycles, as they affect hypothalamic, pituitary and ovarian activity and thus
inhibit follicular development. Under-nutrition contributes to prolonged postpartum anestrus, par-
ticularly among cows dependent upon forages to meet their feed requirements and it apparently
interacts with genetic, environmental or management factors to influence the duration of anestrus.
The nutritional status or balance of an animal is evaluated through body condition score (BCS), as
it reflects the body energy reserves available for metabolism, growth, lactation and activity. There
is a converse relationship between energy balance and time to resumption of postpartum ovarian
activity; inadequate nutrient intake results in loss of weight and BCS and finally cessation of estrous
cycles. Suckling interferes with hypothalamic release of GnRH, provoking a marked suppression
in pulsatile LH release, resulting in extended postpartum anestrus. The effects of suckling on reg-
ulation of tonic LH release are determined by the ability of the cow to identify a calf as her own or
as unrelated. Vision and olfaction play critical roles in the development of the maternal-offspring
bond, allowing the cow to identify her own calf, and abolition of both senses attenuates the negative
effects of suckling on LH secretion. Thus, the maternal-offspring bond is essential for prolonged
postpartum suckling-induced anovulation, and the suppressive influence of suckling is independent
of neurosensory pathways within the teat or udder.
2003 Elsevier B.V. All rights reserved.
Keywords: Anestrus; Body condition score; Cattle; Suckling; Postpartum
Corresponding author. Tel.: +52-29-9342075; fax: +52-29-9344053.
E-mail address: fmontielp@hotmail.com (F. Montiel).
0378-4320/$ see front matter 2003 Elsevier B.V. All rights reserved.
doi:10.1016/j.anireprosci.2003.11.001
2 F. Montiel, C. Ahuja / Animal Reproduction Science 85 (2005) 126
1. Introduction
Anestrus is the state of ovarian acyclicity, reflected by complete sexual inactivity without
manifestation of estrus (Fallas et al., 1987; Wright and Malmo, 1992), and anovulation
accompanied by serum progesterone concentrations less than 0.5 ng/ml (Arregun et al.,
1997). In cattle, anestrus and infertility in the postpartum period were recognized as a
problem for the first time more than 60 years ago (Short et al., 1990). Postpartum anestrus
is the period after parturition during which cows do not show behavioral signs of estrus.
The anestrus condition is associated with the presence of static ovaries, and even though
there is follicular development, none of the ovarian follicles that start growing becomes
mature enough to ovulate. As a result of this lack of follicular maturation, ovulation does
not occur while anestrus is present (Moro et al., 1994). In suckled beef cows the main cause
of prolonged postpartum anestrus is not the failure of the dominant follicles to develop, but
rather their failure to ovulate (Murphy et al., 1990; Stagg et al., 1995).
In the postpartum cow there is a normal anestrus period. In grazing Bos taurus/Bos
indicus cows, anestrus is considered as abnormal when it extends beyond an average of
90 days (Fallas et al., 1987). Prolonged postpartum periods of anestrus (>150 days) are
characteristic of Bos indicus and Bos taurus/Bos indicus cattle of tropical regions; in this
type of cattle prolonged postpartum anestrus is one of the main infertility problems due
to its great incidence, and results in economic losses due to failure to achieve a desirable
inter-calving interval of 12 months (Escobar et al., 1982a; Anta et al., 1989a; Vaccaro,
1990). Postpartum anestrus is more marked in beef than in dairy cattle (Galina and Arthur,
1989b; Galina et al., 1989).
The main factors affecting the duration of postpartum anestrus in cattle are the nutri-
tional status (measured by BCS) and suckling (Short and Adams, 1988; Randel, 1990).
Some other factors such as breed, age, number of calvings, milk yield, calving season,
presence or absence of the bull, delay uterine involution, dystocias and general health status
influences duration of postpartum anestrus (Morrow et al., 1969; Macmillan and Clayton,
1980; Oyedipe et al., 1982; Peters, 1984; Galina and Arthur, 1989b; Galina et al., 1989).
Nonetheless, Wettemann (1994) states that any factor considered as a probable cause of
postpartum anestrus, other than nutrition and calf presence, only modulates the effects
provoked by these two major factors.
Although the roles of gonadotropins and other endocrine regulators of follicular function
during the early postpartum anestrus period are not clear, ultrasonography now makes it
possible to compare changes in the ovarian follicular growth with endocrine and metabolic
variables in order to delineate the endocrine cause of prolonged anestrus in cows (Murphy
et al., 1990; Stagg et al., 1995).
2. Body condition
The existence of a relationship between reproduction and nutritional status in cattle

has been established (Moore and Campos da Rocha, 1983; Galina and Arthur, 1989b;
Arregun et al., 1997), with under-nutrition being reported to be the main hindering factor
in bovine production systems in tropical regions. Under-nutrition, or the inadequate intake
F. Montiel, C. Ahuja / Animal Reproduction Science 85 (2005) 126 3
of nutrients relative to metabolic demands, is a major factor contributing to prolonged

postpartum anestrus, particularly among cows dependent upon natural forages for most or
all of their feed requirements (Jolly et al., 1995). Under-nutrition may also interact with
genetic, environmental or management factors to influence duration of postpartum anestrus,
and although the precise nature of these factors and their interactions is complex and often
unclear, many of them appear to act via common hormonal mechanisms (Jolly et al., 1995).
2.1. Body condition score evaluation
The nutritional status or balance of an animal, evaluated through BCS, reflects the body
reserves available for basic metabolism, growth, lactation and activity (Wright et al., 1987).
Body fat is indicative of the energy stored. To maintain health, reproductive function and
productive capacity, females must have adequate amounts of body reserves, particularly
dairy cows (Edmonson et al., 1989). The rate of synthesis or degradation of muscular
protein can vary in response to changes in nutritional and endocrine status experienced by
cows after parturition, and loss of muscle protein during lactation has been reported in some
(Chilliard and Robelin, 1983), but not in all (Butler-Hogg et al., 1985) studies with dairy
cows. Although the maximum amount of fat a cow can lose without affecting its basic vital
functions is not known, there are serious limitations when 25% of the body protein is lost
in lactating dairy cows (Botts et al., 1979). Therefore, the amount of muscle protein loss
can be a source of information to estimate the metabolic status in these cows (Zurek et al.,
1995).
Evaluation of BCS is an effective way to measure, by sight and touch, the amount of
metabolic energy stored as subcutaneous fat and muscle on an animal (Wildman et al.,
1982; Wagner et al., 1988; Edmonson et al., 1989; Houghton et al., 1990). The technique
for BCS evaluation, initially adapted for beef cattle (Lowman et al., 1976), is also used in
dairy cattle, and involves palpation of the back bones and lumbar processes, feeling their
sharpness and covering with muscle and fat, scoring the animals on a scale of 1 (emaciated)
to 5 points (obese), with quarters or half points in the middle; however, to make BCS more
practical and for situations where it is not possible to palpate the animals, evaluation can
be done visually (Wildman et al., 1982; Edmonson et al., 1989). Using a 19 scale, Wagner
et al. (1988) determined the BCS accounts for about 80% of the variation in carcass fat
in beef cows. Despite differences between temperate cattle, for which this technique was
devised, and tropical cattle, the same measurement scale is used with similar results in both
types (Montiel, 2001). Several authors recommend evaluation of BCS because it reflects
the nutritional status (Short et al., 1990; Arregun et al., 1997), in beef (Rae et al., 1996)
and dairy cows (Wildman et al., 1982; Edmonson et al., 1989). In dairy cows, this technique
has been widely recommended as a method for evaluating their nutritional management
(Gillund et al., 2001).
When cows have low body energy reserves they may have a greater probability of suffering
from diseases, metabolic disorders, reproductive failure and reduction in milk yield, and
heifers are older at puberty (Edmonson et al., 1989). Dairy cows with excessive fat reserves
or over-condition at calving may have a greater risk of lower milk yield and of increased
health and reproductive disorders, such as dystocia and fatty liver (fat cow syndrome). Dairy
heifers with excessive BCS have difficulties in getting pregnant as well as disorders in
development of the mammary gland, resulting in lower productivity (Morrow, 1976; Fronk
et al., 1980; Boisclair et al., 1986; Gearhart et al., 1990). Nonetheless, fat or over-conditioned
cows are not common among high producing dairy cows (Domecq et al., 1997a), nor in Zebu
or dual-purpose cows of tropical regions, fed only by grazing (Montiel, 2001). A number
of authors have pointed out the effects of BCS at calving and its influence on lactation,
health and fertility in high producing dairy cows (Gearhart et al., 1990; Ruegg et al., 1992;
Waltner et al., 1993; Ruegg and Milton, 1995; Heuer et al., 1999). In several studies, the
proportion of overconditioned cows at calving was less than 10% (Ruegg and Milton, 1995;
Markusfeld et al., 1997; Heuer et al., 1999).
2.2. Energy balance
In dairy cows, attempts have been made to relate the interval between calving and first
ovulation with metabolic status (Zurek et al., 1995), and a converse relationship between
energy balance and time elapsed until resumption of postpartum ovarian activity has been
reported (Butler and Smith, 1989; Senatore et al., 1996; De Vries et al., 1999). Likewise,
the lack of nutrient intake results in loss of weight and BCS and cessation of estrous
cycles in beef cows (Richards et al., 1989; Bishop and Wettemann, 1993). In dairy cows,
some authors (Butler et al., 1981; Allrich et al., 1987) have noticed a significant positive
relationship between mean energy balance in the first weeks after calving and interval to
first ovulation, whereas others (Villa-Godoy et al., 1988; Spicer et al., 1990) have found no
relationship between mean negative energy balance and duration of postpartum anestrus.
Butler et al. (1981), Canfield et al. (1990) and Canfield and Butler (1991), found that first
ovulation occurred approximately 10, 14, and 14 days after the maximum negative energy
balance, respectively. When ovulation occurred, energy balance was still negative, but in
all cases was in a decreasing status (Zurek et al., 1995).
Nutritional status in the pre- and postpartum periods influences subsequent reproductive
performance. Thus, inadequate protein and energy intake during pregnancy or early lactation
results in low BCS at calving and a longer inter-calving period in beef cows (Laflamme and
Connor, 1992). During the early postpartum period, under-nutrition increases percentage of
anestrous cows (Robinson, 1990). In beef cattle, this reduces the possibility for having a high
percentage of cows that have initiated estrous cycles during the breeding season (Dziuk and
Bellows, 1983; Richards et al., 1986). Cows fed only by grazing have greater probabilities
for being anestrous (Roche et al., 1992; McDougall et al., 1995b), and under-nutrition can
induce anestrus, particularly in Bos indicus breeds (Jolly et al., 1995). During the early
postpartum period in beef cattle there is a suppression of cylces of ovarian function, that
worsens if the female has a poor BCS. Energy deficit during growth and in the pre- and
postpartum periods inhibits estrus onset and reduces fertility in females (Williams, 1989),
because during a negative energy balance, growth and follicular maturation are inadequate,
resulting in poor estrus manifestation and in deficient results in programs for synchronization
of estrus (Stevenson et al., 1987).
After calving, dairy cows experience a slow increase in dry matter intake, a rapid increase
in milk production and increased mobilization of fat tissue. Therefore, during early lactation
BCS is evaluated to estimate the amount and mobilization of fat tissue, and to find out
whether there is a negative energy balance, which has been found to occur in 80% of cows
(Garnsworthy and Topps, 1982; Butler and Smith, 1989; Edmonson et al., 1989; Gearhart
et al., 1990; Nebel and McGilliard, 1993). The negative energy balance prior to peak milk
production occurs because most dairy cows are not able to meet the energy requirements
for growth, maintenance and milk production (Waltner et al., 1993).
During early lactation, reproductive performance of dairy cows, particularly conception
rate, may be negatively associated with the magnitude of the negative energy balance (Butler
and Smith, 1989; Nebel and McGilliard, 1993). Positive relationships between interval from
calving to first ovulation and magnitude of negative energy balance, and between interval
from calving to minimum energy balance (nadir) have been found, and it has been shown that
dairy cows start lactation in negative energy balance (values below zero), which reaches its
minimum level at the first or second week, but thereafter it steadily increases until reaching
a maximum level, then starts to slowly decrease (Butler et al., 1981; Butler and Smith,
1989; Canfield and Butler, 1990; Zurek et al., 1995; Beam and Butler, 1998; De Vries et al.,
1999). Senatore et al. (1996) found a negative effect of energy deficit on ovulation during
early lactation in first lactation dairy cows.
Therefore, during early lactation nutritional level is quite important. Cows use reserves
for milk production, which means a significant loss of body weight and a progressive
decrease of BCS. This leads to a delay in resumption of postpartum ovarian activity (Gallo
et al., 1996), because the cows priority is milk production and not the establishment and
maintenance of estrous cycles (Garca-Winder, 1990).
After calving, individual cows respond to the negative energy balance through differ-
ent combinations of feed intake, fat tissue mobilization and a decrease in milk yield
(Garnsworthy and Topps, 1982; Butler and Smith, 1989; Nebel and McGilliard, 1993).
Cows with low BCS have a more rapid increase in dry matter intake and achieve a positive
energy balance before cows with greater BCS. Cows with greater BCS at calving lose more
BCS than cows with lesser BCS (Garnsworthy and Topps, 1982; Ducker et al., 1985; Butler
and Smith, 1989). Garnsworthy and Topps (1982) suggest that the relationship between
BCS at calving and milk yield varies, and cows with greater BCS at calving generally lose
more BCS during lactation, which might negatively influence milk yield, whereas low pro-
ducing dairy cows lose less BCS than high producing dairy cows (Gallo et al., 1996). Frood
and Croxton (1978) showed that cows with low BCS at calving (1.5 in 15 scale) did not
lose BCS during early lactation, and cows with moderate BCS (3.0) lost more BCS until the
second month after parturition. In dairy heifers with greater BCS at calving, the extended
negative energy balance may contribute to reduced fertility (Villa-Godoy et al., 1990).
2.3. Pre- and postpartum nutrition
In the beef cow, prepartum nutrition reflected by BCS at calving, is a greater determinant
on the length of postpartum anestrus than postpartum nutrition is (Dunn and Kaltenbach,
1980; Dziuk and Bellows, 1983; Richards et al., 1986; Selk et al., 1988). There is a negative
correlation between BCS at calving and duration of postpartum anestrus (Richards et al.,
1986; Wright et al., 1987, 1992). Angus cows fed on a greater nutritional plane 55 days
before and 40 days after calving averaged 35 kg of weight more and had a shorter interval
from calving to first estrus than cows fed a medium nutritional plane (Montgomery et al.,
1985). BCS at the time of parturition is likely the most important factor affecting subsequent
net calf-crop in mature beef cows. A BCS of 5 (19 scale) at calving is important to ensure
acceptable postpartum reproduction in multiparous cows. Cows with BCS >6 or <5 may be
managed to lose or gain condition during the last trimester of pregnancy, to attain BCS of 5
at calving. In either case, subsequent reproductive performance should be equal to cows that
have maintained BCS. Therefore, reproductive performance of cows calving in moderate
BCS is not influenced by large changes in body energy reserves during the last trimester of
pregnancy (Morrison et al., 1999).
Severe under-nutrition of beef cows in the last trimester of pregnancy and postpartum may
result in the absence of ovarian follicles 5 mm in diameter or of larger (>8 mm) follicles
than can produce appreciable amounts of estradiol (Perry et al., 1991; Jolly, 1992). Perry
et al. (1991) found that growth of follicles 8 mm was virtually abolished in beef cows fed
a low energy diet before and after calving. Low energy intake after calving also resulted
in a reduced rate of appearance of smaller follicles (5.07.9 mm), which persisted in the
ovaries for prolonged periods, perhaps reflecting the absence of functionally dominant large
follicles (Perry et al., 1991). Likewise, in Bos indicus cows, no underfed cows had estrogenic
follicles >8 mm, and half of them had no follicles >5 mm present in either ovary at 610
weeks postpartum. However, in small (24 mm) and medium (57 mm) sized follicles that
were present, estradiol synthesis by individual follicles in culture was not different between
cows fed maintenance or restricted diets (Jolly, 1992). These findings are consistent with
others suggesting that chronic under-nutrition has little effect on the steroidogenic function
of small- or medium-sized follicles in beef cows (Prado et al., 1990; Rutter and Manns,
1991; Ryan et al., 1994).
The effects of postpartum nutrition on the length of postpartum anestrus are apparently
inconsistent in beef cows, with no effect (Wettemann et al., 1986; Wright et al., 1992) or
significant effects (Rutter and Randel, 1984; Richards et al., 1986; Wright et al., 1987).
This inconsistency may reflect interactions among pre- and postpartum nutrition, negative
energy balance, BCS, milk yield and suckling, as well as other environmental factors that
influence duration of postpartum anestrus.
In tropical regions, cows calving with acceptable BCS, and that maintain it after calving,
have greater reproductive rates than cows calving in poor BCS (Galina and Arthur, 1989a),
due to greater hypophyseal function and as a consequence a greater reproductive potential,
that translates into an early return to estrus after calving (Rutter and Randel, 1984). In
dairy cows, an excessive (4.0 points, scale 15) or insufficient (<2.0 points) BCS at
calving reduces subsequent fertility rates. In addition, loss or gain of one BCS after calving
significantly reduced fertility in cows with acceptable BCS before parturition (Gaines,
1989), and affected the general health status of the animal (Ferguson, 1991). Otto et al. (1991)
reported that a one BCS change in dairy cattle equals a 2560 kg change in live weight.
Likewise, Wright et al. (1987) indicate that each BCS is equivalent to 53 kg of body weight
in beef cows, and that postpartum anestrus extended 43 days for each BCS lost at calving.
Although BCS at calving has particular importance for the early re-establishment of
ovarian activity in cattle (Haresign, 1984; Holness, 1984), in tropical regions there is marked
seasonal variation in both the quantity and the quality of available forage throughout the
year, resulting in prolonged periods of sub-maintenance nutrition (McIvor, 1981). This is
particularly critical during the peripartum period, when superior quality forage is required
for at least the last trimester of pregnancy and for 812 weeks postpartum. This minimum
period of 6 months means that in regions where feed intake depends on the availability
of forage, which is influenced by the rainy season (as in most of tropical regions), it is
difficult to provide the cattle with the necessary energy and protein for this critical stage
in their reproductive life (Galina and Arthur, 1989b). In Bos indicus crossbred beef cattle,
nutritional stress is particularly severe in late pregnant and lactating cows during the dry
season, resulting in a major loss of live weight and BCS (Jolly et al., 1996). Under these
conditions, prolonged postpartum anestrous periods are common, and are one of the main
limiting factors for Zebu cattle (Entwistle, 1983). Early weaning of Zebu calves has reduced
weight loss in cows pospartum (Moore and Campos da Rocha, 1983).
2.4. Secretion of hypothalamic and hypophyseal hormones
The complexity of the relationship between nutritional status and reproductive function
has been reviewed (Short and Adams, 1988; Randel, 1990). According to Short and Adams
(1988), blood glucose concentrations may link nutritional status with reproductive function
at hypothalamus in beef cattle. When cows are in negative energy balance (blood glucose
levels <30 mg/dl) fertility is reduced (McClure, 1970). Dairy cows with reduced concentra-
tions of glucose in blood have reduced progesterone in plasma (Villa-Godoy et al., 1988).
Body fat reserves regulate secretion of hypothalamic and hypophyseal hormones that control
ovarian function (Roberson et al., 1992). An important mechanism by which energy deficit
impairs reproductive activity is by suppressing release of GnRH, and as a consequence, the
frequency of LH pulses necessary to achieve follicular growth (Schillo, 1992). As energy
balance, mean plasma LH concentrations and number of episodic LH peaks increase, the
first ovulation occurs in dairy cows (Canfield et al., 1990; Canfield and Butler, 1991).
The possible mechanism by which under-nutrition may inhibit resumption of postpartum
ovarian activity in cows is by the inhibition of hypothalamic GnRH secretion. However, little
is known about the specific ways in which information about nutritional state is translated
into neuroendocrine signals that affect GnRH secretion (Jolly et al., 1995). Some mech-
anisms include metabolic hormones involved in regulation of intermediary metabolism
(particularly insulin), as well as the availability of specific amino acids (as tyrosine) and
oxidizable metabolic fuels (Schillo, 1992). In recent years, it has been suggested that cen-
tral effects of under-nutrition on hypothalamic GnRH secretion may be partly mediated
by the neuropeptide Y, which also appears to play an important role in the regulation of
feed intake and energy balance in ruminants (Miner, 1992; McShane et al., 1993). Jolly
et al. (1995) state that it is still unknown whether nutritional effects on follicular growth
and function in cows are due simply to changes in gonadotropin secretion, or involve local
effects within the ovary, possibly mediated by metabolic hormones or growth factors. Thus,
chronic under-nutrition in suckled postpartum beef cows may induce a state analogous to
that of hypophysectomy, totally suppressing the maturation of follicles which are dependent
upon gonadotropin support (Perry et al., 1991; Jolly, 1992).
Growth of bovine follicles beyond 45 mm, and their ability to produce considerable
amounts of estradiol, are dependent upon adequate FSH and LH support (Driancourt, 1991;
Fortune, 1994). FSH controls the growth of follicles greater than 5 mm, while LH is asso-
ciated with the maintenance of dominant follicles and their ability to produce substantial
amounts of estradiol necessary for induction of estrus, preovulatory LH surge and ovulation
(Fortune, 1994). Inadequate LH, and normal FSH secretion, during nutritionally induced
anestrus is associated with less estradiol production by dominant follicle (Bossis et al.,
1999). Hourly pulses of LH are required for maturation of follicles (Jolly et al., 1995). LH
is also responsible for the amount of estradiol a large follicle can produce, as its synthesis
by the granulosa cells depends upon the supply of aromatizable substrate (predominantly
androstenedione) from the adjacent theca interna cells which are governed by LH (McNatty
et al., 1984). Recovery of pulsatile LH secretion requires the re-establishment of the estradiol
positive feedback on hypothalamus and anterior hypophysis, which is suppressed during the
early postpartum period, but which is also typically recovered within 23 weeks after calv-
ing when nutrition is not limiting (Alam and Dobson, 1987; Nolan et al., 1988). Resumption
of growth of large follicles after calving results from increased FSH secretion within 34
days after calving in adequately fed cows (Lamming et al., 1981), but the ability of follicles
to grow and become dominant, and the duration of their functional dominance, probably
depends upon the extent of pulsatile LH secretion, which in turn reflects the amount of tonic
GnRH secretion from the hypothalamus (Jolly et al., 1995).
Short et al. (1990) and Schillo (1992) have reviewed effects of under-nutrition on go-
nadotropin secretion in postpartum cows. These effects are thought to be mediated by inhibi-
tion of the estradiol positive feedback on the preovulatory gonadotropin-surge mechanism,
or by enhanced estradiol negative feedback on the hypothalamus, as well as by direct (ovary
independent) inhibition of GnRH release (Schillo, 1992). In beef heifers, Imakawa et al.
(1987) suggested that inadequate plasma LH concentrations reflect a steroid-independent
ovarian reduction in LH secretion by the hypothalamichypophyseal axis. Deficient nu-
trition in postpartum cows modifies the normal neuroendocrine changes, particularly the
frequency of episodic LH release (Robinson, 1990). Some metabolic components have been
observed to act on the hypothalamichypophysealovarian axis when BCS of the female
decreases (Randel, 1990; Short et al., 1990). Richards et al. (1991) reported that circulating
LH and IGF-1 concentrations decreased during feed restriction in non-pregnant and beef
cows that were not be suckled by calves, and suggested that these responses might be phys-
iologically linked, because there is evidence that IGF-1 is involved in modulation of brain
and hypophyseal functions (Tannenbaum et al., 1983).
In lactating cows, recovery of the GnRH surge mechanism and of the high frequency
pulsatile LH secretion postpartum are inhibited by undernutrition (Echternkamp et al.,
1982; Schallenberger and Prokopp, 1985; Alam and Dobson, 1987; Nolan et al., 1988;
Wright et al., 1990, 1992; Perry et al., 1991). However, thin anestrous beef cows had
greater concentration of GnRH in the infundibular stalk median eminence than fat cows
(Rasby et al., 1992). Failure of the mechanism(s) controlling final ovarian follicle maturation
and ovulation in moderate under-nutrition may involve inhibition of the greater frequency
pulsatile LH release necessary for stimulating greater estradiol secretion and/or central
inhibition of the GnRH surge response to estradiol (Jolly et al., 1995). The transition from
decreasing to increasing energy balance in early lactating dairy cows was associated with
an increase in pulsatile LH frequency (Canfield and Butler, 1990), which suggests that
pulsatile LH secretion may be inhibited until a negative energy balance nadir is reached,
after which pulsatile LH frequency increases. Reductions in the size and persistence of
dominant follicles associated with undernutrition may reflect inhibitory effects on frequency
of pulsatile LH secretion (Jolly et al., 1995).
Prolonged acyclic intervals resulting from under-nutrition are associated with suppression
of both tonic and surge release of GnRH from the hypothalamus, and hence LH and FSH
secretion from the anterior hypophysis. A poor nutritional status is associated with minimal
concentrations of LH and estradiol in plasma, delay in follicular development, and delay in
occurrence or decrease in the magnitude of the ovulatory surge of LH induced by estradiol
(Jolly et al., 1995). In beef cows, Richards et al. (1989) indicated that anestrus induced by
a lesser nutritional plane was characterized by decreased pulsatile LH secretion, whereas
Wright et al. (1992) noted that postpartum maintenance nutrition had greater effect than
BCS at calving on pulsatile LH frequency. In dual purpose cattle, if cows calve with poor
BCS, the hormonal signals to stimulate the ovary and to resume estrous cycles are not
released, and consequently the interval from calving to first ovulation is extended (Anta
et al., 1989b; Moro et al., 1994). When postpartum Bos indicus cows consume small or
minimal amounts of quality forages (Jolly et al., 1991), concentrations of FSH and LH are
more profoundly suppressed (deep anestrus), than when diets are adequate.
FSH appears to be essential for stimulation of growth of large ovarian follicles, and the
absence of follicles >5 mm in diameter or of large estrogenic follicles, occurs in severely
underfed cows (Perry et al., 1991; Jolly, 1992). This may reflect inadequate FSH stimulation
or inhibition of FSH-dependent processes within the ovary. Similarly, marked suppression
of hypothalamic and hypophyseal function may be responsible for the prolonged (36
months or longer) postpartum anestrus in lactating Bos indicus cows during adverse seasonal
conditions in the dry tropics, which markedly constrains reproductive rates in this region
(Jolly, 1992).
In underfed heifers, estradiol negative feedback on gonadotropin secretion may be related
to increased concentrations of unbound (free) steroid in plasma, due to reduced metabolic
clearance or plasma protein binding (Jolly et al., 1995). Lesser energy intakes have been
associated with reduced binding capacity of sex steroid-binding protein(s) in plasma of
intact, but not ovariectomized heifers (Lermite and Terqui, 1991). Circulating concentrations
of steroid hormones that affect feedback regulation of gonadotropin secretion might be
determined by both ovarian outputs and factors affecting protein binding or metabolic
clearance rates (Jolly et al., 1995).
2.5. Postpartum follicular dynamics
At parturition, follicles >5 mm in diameter are generally absent in beef cows (Braden
et al., 1986; Ryan et al., 1994), but regular patterns of growth and regression of large folli-
cles similar to those during the estrous cycle and early pregnancy are evident from as early
as 1 week after calving in most dairy and beef cows (Roche et al., 1992; Fortune, 1993).
In most adequately fed lactating dairy cows, there is ovulation from the first dominant fol-
licle that develops 23 weeks postpartum (Savio et al., 1990). In contrast, in adequately
fed suckled beef cows in both temperate (Murphy et al., 1990) and tropical (Jolly et al.,
1995) environments, recurrent growth and regression of several (16) dominant follicles
may occur before ovulation. The degree of under-nutrition can alter these typical patterns
of follicle growth and ovulation (Jolly et al., 1995). In beef cows, before the first ovulation
postpartum a greater number of small palpable follicles and a consistently smaller size of
the largest palpable follicle occur in cows on sub-maintenance diets than in cows receiving
greater than maintenance requirements (Wiltbank et al., 1964). McDougall et al. (1995a)
demonstrated that moderate amounts of under-nutrition may delay the resumption of post-
partum estrous cycles in pasture-fed dairy cows calving with lesser BCS. This effect is
associated with final maturation of ovarian follicles (Savio et al., 1990; Roche et al., 1992).
When dairy cows received diets that contained lesser amounts of nutrients than the es-
tablished requirements the first dominant follicle did not ovulate, the interval from calving
to first ovulation was extended (43 5 days) and the pattern of postpartum follicular de-
velopment was similar to that described for adequately fed suckled beef cows (Jolly et al.,
1996) (growth and regression of 19 dominant follicles, Savio et al., 1990; Roche et al.,
1992). McDougall et al. (1995a) found that spontaneous ovulation did not occur until about
6 weeks postpartum also in moderately underfed dairy cows. Moreover, Jolly et al. (1995)
have suggested that restriction of prepartum or postpartum feed intake may also affect the
maximum size and persistence of anovulatory follicles that develop after calving. Murphy
et al. (1990) found that long-term energy deprivation of beef heifers was associated with
regular estrous cycles and reductions in both dominant follicle diameter and persistence.
2.6. Pregnancy rate
Besides influencing the length of postpartum anestrus, BCS also influences pregnancy
rate. Martnez and Castillo (1995) in Bos indicus and Bos taurus/Bos indicus grazing cows,
using a BCS evaluation scale of 19 (Wagner et al., 1988), obtained pregnancy rates of 0, 3
and 11% for cows with BCS of 1, 2 and 3, respectively, and of 85% for cows with BCS of 7,
8 and 9. Weaver (1992) indicates that the desirable BCS at calving is 3 (scale 15), varying
from 2.5 to 3.5, whereas Richards et al. (1986) stated that, in multiparous beef cows, a BCS
of 5 (scale 19) at calving seems to be the critical level affecting subsequent reproductive
performance, and that this critical BCS of 5 seems to be consistent among a number of beef
breeds and crosses (Tinker et al., 1989). Basurto et al. (1998) obtained greater first service
conception rates in dual-purpose cows with BCS >2.5 than with BCS <2.5 (scale 15). In
Gyr cows, Moore and Campos da Rocha (1983) reported that the interval from calving to
conception was reduced from 160 to 116 days in cows fed a high energy diet.
Several studies with dairy cows and heifers have shown that females with excessively high
BCS at calving may require a greater number of services to get pregnant (Ducker et al., 1985;
Villa-Godoy et al., 1990). To this respect, Ferguson (1991) reported that change in BCS from
calving to first breeding is a good predictor of conception from first AI service in dairy cows,
as cows that lost 1.0 BCS had conception rates that were less than cows with no BCS loss or
gain. Similarly, Domecq et al. (1997b) indicate that BCS loss between calving and AI service
can negatively influence conception rates. Cows that lost 0.51.0 BCS had a conception rate
to first AI service of 53%, whereas when cows lost more than 1.0 BCS between parturition
and time of AI the rate was 17% (Butler and Smith, 1989; Ferguson and Otto, 1989).
Ruegg and Milton (1995) indicated that, although they were not able to demonstrate a
statistically significant effect of BCS on reproductive performance in dairy cows, there was
some indication that cows that experience excessive BCS loss (>1.0 point) prior to breeding
may require more AI services. Conversely, others indicate that BCS of dairy cows at calving
or at the time of AI appears not to affect conception rates or AI services per conception
(Bourchier, 1987; Jones and Garnsworthy, 1988; Butler and Smith, 1989; Ruegg et al., 1992).
Although Ruegg and Milton (1995) did not find relationships among energy balance, BCS
and reproductive performance in dairy cows, Domecq et al. (1997b) state that associations
among energy balance, BCS and conception to first AI services are probably greatest in
early lactation. Therefore, the use of BCS to monitor changes in the negative energy balance
during the non-lactating period and during the first 30 days of lactation may be important
to enhance conception to first AI service.
3. Suckling
Prolonged postpartum anestrus is a major limitation to achieve a optimal reproductive

efficiency in beef cattle, and suckling is one of the main factors that influences the length
of the postpartum anestrus period (Stagg et al., 1998). If suckling is continuously practiced
during lactation, it blocks ovulation with the consequent long period of postpartum anestrus,
contributing to a lesser reproductive efficiency (Wettemann, 1994). In tropical cattle, a long
postpartum anestrus is one of the main factors affecting reproduction, and to shorten it is
crucial to achieve the desirable goal of one calving every 12 months (Ramrez-Iglesia et al.,
1992).
Suckling and milk yield can affect activity of the hypothalamus, hypophysis and ovaries,
inhibiting follicular development and extending the anestrous period. The amount of milk
produced during the postpartum period can affect the interval from calving to ovulation.
Suckling reduces the hypothalamic release of GnRH, which results in insufficient pulsatile
LH release (Williams, 1990). Inhibition of pulsatile LH secretion during lactation and the
corresponding increase in pulsatile LH secretion 26 days after removal of the suckling
stimulus is well established (Walters et al., 1982; Smith et al., 1983; Williams et al., 1987).
3.1. LH secretion
The primary endocrine deficit resulting in the establishment of postpartum anovulation

in suckled beef cows is the failure to resume a pattern of LH secretion that will support
the development and final maturation of an estrogen-active, preovulatory follicle. This
deficit depends upon central mechanisms associated with control of GnRH-secreting neu-
rons (Williams et al., 1983). At parturition, LH concentrations in the anterior hypophysis
are depleted due to the inhibitory effect of placental-derived estradiol on the synthesis of LH
and subunits (Nett, 1987). This negative feedback sensitivity is potentiated by suckling
and may be a major factor regulating resumption of normal LH secretion (Acosta et al.,
1983). However, estradiol antagonists have not been consistently successful in shortening
the postpartum anovulatory interval (Schramm et al., 1991). The inhibitory effect of estra-
diol ends at calving, and by 23 weeks postpartum stores of LH are similar to amounts
in cows with estrous cycles (Williams et al., 1982; Nett et al., 1988). A new pattern of
LH secretion would be expected coincident with repletion of LH stores, and the onset of
follicular maturation and ovulatory competence. However, this does not occur consistently,
and anovulatory periods in some females persist for 100 days or more after calving (Casida,
1971). In dairy cows, the biopotency of LH secreted increases from calving until about 12
days after, and may play a role in the sequence of events leading to first ovulation (Weesner
et al., 1987). Resumption of estrous behavior after calving is partly determined by the ability
of the hypophysis to secrete adequate amounts of LH (Jaeger et al., 1995).
In the early postpartum beef cow, at 26 days after removal of the suckling stimulus,
there is an increase in pulsatile LH secretion which has been inhibited during the postpar-
tum anovulatory status (Walters et al., 1982; Smith et al., 1983). Thus, if calves are weaned
at birth, resumption of pulsatile LH secretion is observed within 714 days postpartum,
followed by resumption of ovarian cyclicity (Williams et al., 1983). If weaning is delayed
until at least 2 weeks after calving, a different response is observed than that seen immedi-
ately after calving (Williams and Griffith, 1995). As suckling delays release of hormones
needed for resumption of estrous cycles after parturition, cows that lose their calf at birth
commonly show estrus earlier than cows with suckling calves (Wettemann, 1994).
Unlike changes observed following calf removal at birth, acute weaning of chronically
suckled anestrous beef cows is characterized by a rapid increase in pulsatile LH secretion
within 4896 h (Smith et al., 1983; Shively and Williams, 1989). This is initially accom-
panied with a rapid decline (secretion > synthesis) followed by an increase (synthesis >
secretion) in content of GnRH within the hypothalamus (Malven et al., 1986). The respon-
siveness of the anterior hypophysis to GnRH is increased with the secretion of LH into
the peripheral circulation (Walters et al., 1982). After resumption of ovarian cycles, release
of content and secretion of both hypothalamic GnRH and hypophyseal LH revert to those
for cows weaned at birth. The acute response to weaning can be markedly attenuated by
the premature return of calves, and it may require up to 144 h for all cows to respond to
temporary weaning (Shively and Williams, 1989; Cutshaw et al., 1991).
Inhibition of episodic LH release provoked by suckling may be influenced by the secre-
tion of an hypothalamic opioid peptid -endorphin, in response to the suckling stimulus
(Malven et al., 1986; Boland et al., 1990). Concentrations of endogenous opioid peptides in
neural tissue are affected by suckling, and short-term administration of opioid antagonists
increase LH secretion in suckled anestrous cows (Williams and Griffith, 1995). During late
pregnancy, large amounts of placental-derived estradiol inhibit LH synthesis by direct action
on pituitary gonadotrops (Nett, 1987). If both estradiol negative feedback and opioid inhi-
bition at the hypothalamus are increased, it is possible that gestational increases in estradiol
induce changes in the microenvironment of GnRH-secreting neurons that, although perhaps
not irreversible, require a recovery period before normal function is restored. Suckling may
impede the recovery rate of the hypothalamuspituitary axis from a state of greater opi-
oid inhibition that is established during late gestation. Secretion of opioids may decrease
because physiological effects of suckling are eventually reduced and most cows initiate
normal ovarian cycles (Williams and Griffith, 1995).
3.2. Restricted and unrestricted suckling
Although Randel (1981) indicates that suckling by a single calf less than twice daily
substantially and consistently reduces the postpartum anovulatory interval in first calf Bos
indicus/Bos taurus heifers, other authors (Day et al., 1987; Williams, 1990) state that fre-
quency and duration of ad libitum suckling by a single calf apparently are not related to
duration of anestrus, except under extreme circumstances. The effect of suckling in Bos in-
dicus cows (Montoni and Riggs, 1978; Escobar et al., 1982b; Moore and Campos da Rocha,
1983; Bastidas et al., 1984) appears to be related to the suckling frequency rather than to
the nutrients taken from the cow, and when the suckling stimulus is reduced, improved
reproductive performance occurs without affecting calf growth. Reducing the traditional
weaning age from 7 to 4 months, can advance the next calving date by 39 days in Bos
indicus cows (Schottler and Williams, 1975).
When suckling occurs twice or three times a day, duration of ovarian acyclicity increases,
plasma LH concentrations decrease, sensitivity of the hypothalamichypophyseal axis to the
inhibitory effects of estradiol on LH increases (Walters et al., 1982; Acosta et al., 1983; Shiv-
ely and Williams, 1989; Short et al., 1990; Williams, 1990) and GnRH secretion is inhibited
in the absence of estradiol (Zalesky et al., 1990), compared with suckling once a day. Com-
plete weaning, partial weaning (less suckling episodes) or temporary weaning (for 4872 h),
together with treatments to synchronize estrus, can provoke an increase in plasma LH con-
centrations and hasten the onset of ovulation and ovarian cycles (Wright and Malmo, 1992).
In Bos taurus cattle, suckling can prolong postpartum anestrus for more than 90 days,
and is a main cause of lesser reproductive efficiency (Wright et al., 1987). This also occurs
in dual-purpose cows (Bos taurus/Bos indicus) (Acosta et al., 1983). In beef cattle, temporal
weaning for 48 h usually increases conception rate to fixed time AI with different estrous
synchronization protocols (Smith et al., 1979; Kiser et al., 1980; Yelich et al., 1995). Like-
wise, early weaning, or suckling only once a day, shortens the interval from calving to first
estrus in primiparous cows less than 85 days postpartum and calving in good BCS, which
does not happen in cows that are greater than 85 days postpartum (Bell et al., 1998).
In Bos indicus cows, restriction of suckling to once (Shively and Williams, 1989; Williams,
1990) or twice a day (Tegegne et al., 1992) reduces duration of anestrus compared with un-
restricted suckling, as was also indicated by Del Vecchio et al. (1988) and Ruas et al. (1991).
The effect of suckling frequency on duration of postpartum anestrus remains unclear in dual
purpose cows that maintain BCS during the early postpartum period (Montiel, 2001).
Suckling delays postpartum ovarian activity in Bos indicus cows (Moore, 1984), and
early weaning reduces the calving interval (Azevedo et al., 1981). Weaning calves during
the first week after calving dramatically reduces the time from calving to the onset of
ovarian activity (Arora et al., 1971; Moore and Campos da Rocha, 1983; Wells et al., 1985).
However, the effect of weaning calves beyond the first week after calving results in varied
responses (Maree et al., 1974; Salcedo et al., 1977; Pimentel et al., 1979; Alberio et al.,
1984; Lishman and Harwin, 1985).
Restricted suckling of Bos indicus cows increases pregnancy rate (Bastidas et al., 1984).
In Bos indicus and Bos taurus/Bos indicus cows, restricted suckling has a more beneficial ef-
fect on primiparous heifers than in multiparous cows (Arora et al., 1971; Montoni and Riggs,
1978; Randel, 1981). In first calf Bos indicus crossbred beef heifers, the effect of weaning
on resumption of ovarian activity and on duration of postpartum anestrus was greater than
the effect of feed supplementation during the postpartum period (McSweeney et al., 1993,
1993a). In Bos indicus cows, early weaning (23 months of age) reduced loss in weight and
BCS during the dry season, and increased pregnancy rates in some herds, but responses var-
ied widely (Schlink et al., 1988, 1994; Sullivan et al., 1992). In Bos indicus and Bos taurus
cows, the delayed response to weaning has been related to the low body weight or BCS of
the cow (Mukasa-Mugerwa et al., 1991; Bishop et al., 1994). It has also been proposed that
inhibitory effects of suckling and under-nutrition interact, because the increase in suckling
intensity has been associated with lesser milk production in underfed cows (Loudon, 1987).
The nature of this interaction may depend on the relative partitioning of nutrients between
milk synthesis and maternal growth and storage, a process which also varies according
to genotype (Hunter and Magner, 1988). Partitioning of nutrients during lactation may be
regulated by insulin, growth hormone, and the IGF-1, all of which are sensitive to changes
in the nutritional status (Brockman and Laarveld, 1986; Gluckman et al., 1987).
In tropical regions with a predominance of Bos indicus cattle, it is usual to share the milk of
the cow with the calf. In these cows, milk let-down only occurs in the presence of their calves,
and consequently, calves often suckle for 89 months before weaned (Mukasa-Mugerwa
et al., 1991). In Zebu cows maintained continuously with calves, the onset of first estrus
postpartum was 169 days, whereas cows that suckled only 2 h in the morning and 2 h in the
afternoon it was 140 days, and shortened to 110 days when suckling was only for 2 h in
the morning (Brito, 1974). Likewise, Randel and Welker (1976) found that the first estrus
postpartum was 69 days when dam and calf were separated the whole day, allowing the
calf to suckle only for 0.5 h, in comparison with 116 days when the calf remained with its
dam all the time. In Gyr cows suckled only during one month, return to estrus averaged
40 days postpartum, which was 23 days before cows suckling for longer periods (Moore
and Campos da Rocha, 1983). Bluntzer et al. (1989) found and improvement in conception
and pregnancy rates at 90 days in primiparous Bos indicus crossbred cows with once a day
suckling compared with cows with unrestricted suckling. In Bos indicus cows, Bastidas et al.
(1981) also found that suckling twice a day increased pregnancy rate by 33% compared
with continuous suckling.
3.3. Suckling own or unrelated calves
Although several authors (Short et al., 1990; Williams, 1990) have reviewed the physi-
ological basis of anovulation and infertility in postpartum cattle focusing on suckling and
its neuroendocrine events, new concepts have emerged regarding the role of the suckling
calf in the attenuation of the hypothalamichypophysealovarian axis (Silveira et al., 1993;
Viker et al., 1993; Williams et al., 1993). Maternal identification and selectivity for the
cows own calf influences the response due to suckling (Williams and Griffith, 1995).
Under natural conditions, most cows only allow their own calves to suckle; therefore,
it is possible that only a cows own calf can attenuate gonadotropin release (Williams
and Griffith, 1995). Nonetheless, when a cow adopts an additional calf, the postpartum
anestrous period is extended markedly (Wettemann et al., 1978). Hence, Williams and
Griffith (1995) tested the hypothesis that exteroceptive cues responsible for the suppression
of LH secretion are specifically attributable to the dams own calf and, therefore, are a
product of the maternal-offspring bond (Silveira et al., 1993). Williams and Griffith (1995)
found that removal of natural calves from cows resulted in the expected increases in serum
LH concentrations and pulse frequency within 48 h, and suckling by unrelated calves every
6 h for 4 days did not prevent these increases; however, suckling by cows own calf at 6 h
intervals maintained the suppressed pattern of LH release typical of suckled, anovulatory
females. Moreover, mean postpartum intervals to onset of luteal activity in weaned and
unrelated-suckled groups were nearly identical, both of which were markedly reduced
compared with cows suckling their own calves.
According to Williams and Griffith (1995), the neural input at the teat by the suck-
ling calf is irrelevant to the inhibition of LH secretion, for mechanical, electrical and
thermal hyper-stimulation of sensory neurons in the teat failed to simulate the effects of
suckling; therefore, neither the free-running pattern of LH secretion in ovariectomized,
non-lactating females (Williams et al., 1984), nor the postweaning LH increase in lactating
cows (Cutshaw et al., 1991) was impeded. Conversely, neither mechanical masking (McVey
and Williams, 1991) nor complete mammary de-nervation (Williams et al., 1993) prevented
suckling-induced inhibition of LH, and intervals from parturition to first ovulation were not
affected. In this respect, mastectomized cows whose calves were removed at calving exhib-
ited slightly shorter postpartum anovulatory intervals than did weaned-intact cows (Short
et al., 1972), whereas mastectomized cows maintained with their natural calves with unlim-
ited interaction exhibited anovulatory periods similar to suckled-intact cows (Viker et al.,
1989) and to milked cows maintained with muzzled calves with unlimited oral access to the
inguinal region (Macmillan, 1983). Calves maintained with mastectomized cows exhibited
chronic pseudosuckling, which was defined as positioning by the calf in the reverse par-
allel or perpendicular position, bunting, oral manipulation of skin of the leg or flank, and
characteristic head to tail contact between cow and calf (Williams and Griffith, 1995). If
cowcalf interaction is restricted mechanically (Williams et al., 1987) or spatially (Viker
et al., 1993; Stevenson et al., 1994), eliminating direct oral contact with the inguinal region,
the anovulatory state is not maintained. Hence, the direct oral contact with the inguinal re-
gion and, therefore, the mere perception of being suckled, may be sufficient for prolonging
the period of anovulation (Williams and Griffith, 1995).
3.4. Role of vision and olfaction
The role of vision and olfaction in suckling-mediated inhibition of LH secretion in cattle

has been investigated (Griffith and Williams, 1994). Olfaction plays a critical role in the
development of maternal selectivity, with a minimum time required for sensitization of
5 min, with up to 5 h of effect in the absence of the calf (Hudson and Mullord, 1977).
However, cows rendered anosmic after the maternal bond has been established, recognized
their calves visually, allowed them to suckle and continued to exhibit an inhibited pattern of
LH secretion in a suckled environment (Griffith and Williams, 1994). Neither blindness nor
anosmia in cows nursing their own calves caused disinhibition of gonadotropin secretion
(Williams and Griffith, 1995). At 3 weeks after calving cows were separated from their calves
and allowed to be suckled by either their own calf or an unrelated calf every 6 h for 6 days.
Blind, olfactory-intact cows recognized their calves by smell, and anosmic, sighted cows
recognized their calves by sight. Cows that were both blind and anosmic, and were forced to
suckle unrelated calves, exhibited increases in pulse frequency of LH for 6 days. When both
senses were abolished, cows could not identify the calf as their own or as unrelated, behaving
as cows with weaned calves and having the typical increase in LH secretion. In summary,
these results indicate that the net effects of suckling or pseudo-suckling on the central
regulation of tonic LH release are determined by the ability of the cow to identify the calf as
its own or as unrelated when a cow is suckled. Both weaning (total absence of calf-related
stimuli) and suckling by an unrelated calf results in a rapid escape from the inhibitory
influence previously maintained by the cows own calf. Thus, the maternal-offspring bond
is essential for the suckling-induced anovulation, and cows can use both olfactory and visual
cues to identify own and unrelated offspring (Williams and Griffith, 1995).
A sequence of estrogen priming, genital stimulation (oxytocinergic effect) and olfac-
tion may be the primary communication link that drives the full establishment of maternal
behavioral responses in the sheep, and although no data are available, it is assumed that
similar events characterize the development of the maternal behavior in cows (Williams
and Griffith, 1995). In cattle, maximum maternal responsiveness is critical at parturition,
when there is an increased opioid effect (Williams and Griffith, 1995), which is maintained
during early lactation in suckled beef cows, suppressing LH secretion (Malven et al., 1986;
Whisnant et al., 1986). Although no relationship between intracerebral oxytocin release and
suckling-induced inhibition of gonadotropin secretion has been found in cattle, there is an
inverse relationship between factors facilitating maternal behavior and factors modulating
gonadotropin secretion (Silveira et al., 1993). Suckling stimulates peripheral oxytocin re-
lease and increases opioid tone in cattle; however, cows forced to suckle unrelated calves
showed reduced frequency of peripheral oxytocin release and rapid resumption of pulsatile
LH secretion, relative to cows suckling their own calves. Hence, suckling by unrelated calf
might not maintain a hypothalamic opioid tone that can inhibit neuronal stimulation of
GnRH secretory neurons (Williams and Griffith, 1995).
Moreover, the generic precursor of the -endorphins, proopiomelanocortin, and the
GnRH fibres are closely intermingled at the portal capillaries and in the preoptic area of the
brain, and may increase mRNA encoding oxytocin in this area during lactation (Brooks,
1992). Williams and Griffith (1995) hypothesize a close linkage between physiological
variables that modulate maternal behavior, opioidergic tone, and the LH pulse generator
during suckling-mediated anovulation in cows.
It is now clear that the suppressive influence of suckling is independent of the neu-
rosensory pathways within the teat or the udder (Williams et al., 1984; Cutshaw et al.,
1991; McVey and Williams, 1991; Viker et al., 1993; Williams et al., 1993), and that the
maternal-offspring bond is the essential component of the prolonged postpartum anestrus
induced by suckling in beef cows (Silveira et al., 1993; Lamb et al., 1997). It has been
shown that olfaction and vision are equally effective in allowing the calf identification by
its dam (Griffith and Williams, 1996), and that abolition of both senses attenuate the neg-
ative effects of suckling on LH secretion (Stagg et al., 1998). Although temporal weaning
provokes an increase in pulsatile LH frequency (Shively and Williams, 1987), several au-
thors question its usefulness to induce ovarian cyclicity in postpartum cows, lest at the same
time a progestogen-based treatments are applied (Troxel et al., 1980; Walters et al., 1982;
Smith et al., 1983; Williams et al., 1987).
4. Summary
In summary, reproduction is a major component of the productive efficiency for a cowcalf

system, and one of the main causes affecting reproductive performance is the calving
interval, which is directly influenced by postpartum anestrus. Duration of postpartum
anestrus is mainly affected by the nutritional status and suckling (Short and Adams, 1988;
Randel, 1990). Adequate nutrition is essential for resumption of postpartum ovarian activity
in cattle (Robinson, 1990). When nutrient intake is inadequate and body energy reserves are
depleted, interval from calving to first estrus is extended; thus, to establish the relationship
between nutrition and reproductive efficiency, it is necessary to estimate the nutritional sta-
tus of cows during the pre- and postpartum periods through evaluation of BCS (Wettemann,
1994; Butler and Elrod, 1995). When nutrition is improved, particularly during the prepar-
tum period, postpartum anestrus and calving intervals are reduced. In beef cows, when BCS
at calving is very poor its effect on calving interval is greatest, when BCS is adequate or
excellent at calving the interval to conception is reduced and nutrient intake and suckling
have lesser effects on performance (Selk et al., 1988).
Grazing Bos indicus cattle experience long periods of postpartum anestrus, reaching up
to 440 (Oyedipe et al., 1982) or 810 days (Pullan, 1979). This results in very long intervals
from calving to first breeding postpartum, and consequently, excessively long intercalving
intervals (Galina and Arthur, 1989b). In grazing cattle from the Mexican tropics, calving
intervals for Zebu cows have been reported at 382 3.7 days (Hinojosa et al., 1980) and
452 days (Eversbusch, 1978), and 444 days for Bos taurus/Bos indicus cows (Eversbusch,
1978). In this region, lesser BCS at calving and during the early postpartum period, together
with suckling twice a day, increased duration of postpartum anestrus in Bos indicus cows, by
depressing development of the largest ovarian follicles between days 11 and 69 postpartum
(Arregun et al., 1997). In Bos taurus suckling beef cows, the interval from calving to first
estrus is 4048 days, while in dairy cattle it is 3076 days (Wright and Malmo, 1992).
However, in dairy cattle, postpartum anestrus may extend more than 3040 days in cows
with a very negative energy balance (Staples et al., 1990).
Jolly et al. (1996) suggested that even Bos taurus/Bos indicus cows with very poor
BCS may resume ovarian cyclicity if the stress of lactation and/or suckling is removed and
adequate postweaning nutrition is provided. This is consistent with other findings which have
suggested that the predominant factors inhibiting postpartum reproductive performance in
Bos indicus genotypes are those associated with lactation and/or suckling, rather than with
liveweight per se (Frisch et al., 1987; Mackinnon et al., 1989; McSweeney et al., 1993a).
The relative importance of postpartum nutrition and suckling on duration on postpartum
anestrus is not established. However, BCS is the major regulator, since weaning calves
from thin cows does not induce initiation of ovulation immediately, and fatter cows can
have calves suckling them and still have a shorter postpartum interval to ovulation than
thinner cows managed in the same manner.
Acknowledgements
The authors gratefully acknowledge Dr. Robert P. Wettemann for the revision of this
manuscript.
References
Acosta, B., Tarnavsky, T.E., Platt, T.E., Hamernik, D.L., Brown, J.L., Schoenmann, H.M., Reeves, J.J., 1983.
Nursing enhances the negative effect of estrogen on LH release in the cow. J. Anim. Sci. 57, 15301536.
Alam, M.G.S., Dobson, H., 1987. Pituitary responses to a challenge test of GnRH and oestradiol benzoate in
postpartum and regularly cyclic dairy cows. Anim. Reprod. Sci. 14, 19.
Alberio, R., Butler, H., Palma, G., Schiersmann, G., Mihura, H., 1984. Efecto de un destete temporario sobre la
reactivacin posparto de vacas de cra primparas. Rev. Argentina de Prod. Anim. 4, 933939.
Allrich, R.D., Berghorn, K.A., Noller, C.H., 1987. Influence of energy balance on ovarian activity and estrous
behavior in postpartum dairy cows. J. Dairy Sci. 7 (Suppl. 1), 184.
Anta, E., Rivera, J.A., Galina, C., Porras, A., Zarco, L., 1989a. Anlisis de la informacin publicada sobre la
eficiencia reproductiva de los bovinos. II. Parmetros reproductivos. Vet. Mx. 20, 1118.
Anta, E., Rivera, J.A., Galina, C., Porras, A., Zarco, L., 1989b. Anlisis de la informacin publicada sobre la
eficiencia reproductiva de los bovinos. III. Factores que la afectan. Vet. Mx. 20, 1925.
Arora, K.L., Tomar, S.S., Singhal, S.P., 1971. Factors affecting the reproductive efficiency of Hariana cattle: the
duration of post-partum breeding interval. Indian Vet. J. 48, 10151019.
Arregun, J.A.A., Santos, R.E., Villa-Godoy, A., Romn-Ponce, H., 1997. Dinmica folicular ovrica en vacas Ceb
con diferente condicin corporal y frecuencia de amamantamiento durante el perodo anovulatorio posparto.
Divisin de Educacin Continua, Unam, F.M.V.Z. (Eds.), VII Curso Internacional de Reproduccin Bovina.
Mx., D.F., pp. 210240.
Azevedo, N.A., Silva, H.M., Moreira, H.A., Norte, A., 1981. Efeito do perodo de aleitamento sobre o desempenho
reprodutivo de vacas azebuadas. Arq. Esc. Vet. Univ. Fed. Minas Gerais 33, 327334.
Bastidas, P., Troconiz, J., Verde, O., Silva, O., 1981. Effect of restricted suckling on pregnancy rates and calf
performance in Brahman cows. Theriogenology 16, 1325.
Bastidas, P., Troconiz, J., Verde, O., Silva, O., 1984. Effect of restricted suckling on ovarian activity and uterine
involution in Brahman cows. Theriogenology 21, 525532.
Basurto, C.H., Alonso, D.M., Mora, M.C., 1998. Efecto del tiempo pos-parto y condicin corporal sobre la
respuesta al estro y fertilidad en vacas doble propsito tratadas con progesterona y Ammveb, P.M.S.G. (Ed.),
XXII Congreso Nacional de Buiatra, pp. 361362.
Beam, S.W., Butler, W.R., 1998. Energy balance, metabolic hormones, and early postpartum follicular development
in dairy cows fed prilled lipid. J. Dairy Sci. 81, 121131.
Bell, D.J., Spitzer, J.C., Burns, G.L., 1998. Comparative effects of early weaning or once-daily suckling on
occurrence of postpartum estrus in primiparous beef cows. Theriogenology 50, 707715.
Bishop, D.K., Wettemann, R.P., 1993. Pulsatile infusion of gonadrotropin-releasing hormone initiates luteal activity
in nutritionally anestrous beef cows. J. Anim. Sci. 71, 27142720.
Bishop, D.K., Wettemann, R.P., Spicer, L.J., 1994. Body energy reserves influence the onset of luteal activity after
early weaning of beef cows. J. Anim. Sci. 72, 27032708.
Bluntzer, J.S., Forrest, D.W., Harms, P.G., Beverly, J.R., Long, C.R., 1989. Effect of suckling manipulation on
pospartum reproduction in primiparous Brahman-cross cows. Theriogenology 32, 893899.
Boisclair, Y., Grieve, D.G., Stone, J.B., Allen, O.B., MacLeod, G.K., 1986. Effect of prepartum energy, body
condition, and sodium bicarbonate on production of cows in early lactation. J. Dairy Sci. 69, 26362647.
Boland, M.P., Murphy, M.G., Roche, F.S., 1990. The use of ultrasound to monitor ovarian function in farm animals.
Ag. Biotech. News Inf. 2, 841844.
Bossis, I., Wettemann, R.P., Welty, S.D., Vizcarra, J.A., Spicer, L.J., Diskin, M.G., 1999. Nutritionally induced
anovulation in beef heifers: ovarian and endocrine function preceding cessation of ovulation. J. Anim. Sci. 77,
15361546.
Botts, R.L., Hemken, R.W., Bull, L.S., 1979. Protein reserves in the lactating dairy cow. J. Dairy Sci. 62, 433440.
Bourchier, C.P., 1987. The relationship between milk yield, body condition and reproductive performance in
high-yielding dairy cows. Anim. Prod. 44, 460.
Braden, T.D., Manns, J.G., Cermak, D.L., Nett, T.M., Niswender, G.D., 1986. Follicular development following
parturition and during the estrous cycle in beef cows. Theriogenology 25, 833843.
Brito, R., 1974. Estudio de los efectos de la reduccin del tiempo de permanencia del ternero junto a la vaca Ceb,
sobre su actividad sexual y el desarrollo de sus cras. Ciencia Veterinaria 5, 2330.
Brockman, R.P., Laarveld, B., 1986. Hormonal regulation of metabolism in ruminants a review. Livest. Prod. Sci.
14, 313334.
Brooks, P.J., 1992. The regulation of oxytocin mRNA levels in the medial preoptic area: relationship to maternal
behavior in the rat. In: Pedersen, C.A., Caldwell, J.D., Jirkowski, G.F., Insel, T.R. (Eds.), Oxytocin in Maternal,
Sexual and Social Behaviors. The New York Academy of Sciences, New York, pp. 271285.
Butler, W.R., Elrod, C.C., 1995. Reproduction in high-yielding dairy cows as related to energy balance and protein
intake. AIBIR, Divisin de Educacin Continua, Unam, F.M.V.Z. (Eds.), Sexto Congreso Internacional de
Reproduccin Bovina, Mxico, D.F., pp. 2027.
Butler, W.R., Smith, R.D., 1989. Interrelationships between energy balance and postpartum reproductive function
in dairy cows. J. Dairy Sci. 72, 767783.
Butler, W.R., Everett, R.W., Coppock, C.E., 1981. The relationships between energy balance, milk production and
ovulation in postpartum Holstein cows. J. Anim. Sci. 53, 742748.
Butler-Hogg, B.W., Wood, J.D., Bines, J.A., 1985. Fat partitioning in BritishFriesian cows: the influence of
physiological state on dissected body composition. J. Agric. Sci. (Camb.) 104, 519.
Canfield, R.W., Butler, W.R., 1990. Energy balance and pulsatile luteinizing hormone secretion in early postpartum
dairy cows. Dom. Anim. Endocrinol. 7, 323330.
Canfield, R.W., Butler, W.R., 1991. Energy balance, first ovulation and the effects of naloxone on LH secretion in
early postpartum dairy cows. J. Anim. Sci. 69, 740746.
Canfield, R.W., Sniffen, C.J., Butler, W.R., 1990. Effects of excess degradable protein on postpartum reproduction
and energy balance in dairy cattle. J. Dairy Sci. 73, 23422349.
Casida, L.E., 1971. The postpartum interval and its relation to fertility in the cow, sow and ewe. J. Anim. Sci.
32 (Suppl. 1), 6672.
Chilliard, R., Robelin, J., 1983. Mobilization of body proteins by early lactating dairy cows measured by slaughter
and D20 dilution techniques. In: Proceedings of the 4th International Symposium on Protein Metabolism and
Nutrition. Clermont-Ferrand, France, p. 195.
Cutshaw, J.L., Hunter, J.F., Williams, G.L., 1991. Effects of transcutaneous thermal and electrical stimulation of
the teat on pituitary luteinizing hormone prolactin and oxytocin secretion in ovariectomized estradiol treated
beef cows following acute weaning. Theriogenology 37, 915934.
Day, M.L., Imakawa, K., Clutter, A.C., Wolfe, P.L., Zalesky, D.D., Nielsen, M.K., Kinder, J.E., 1987. Suckling
behavior of calves with dams varying in milk production. J. Anim. Sci. 65, 12071212.
Del Vecchio, R.P., Randel, R.D., Neuendorff, D.A., Peterson, L.A., 1988. Effect of suckling manipulation on
pospartum reproduction in primiparous Brahman-cross cows. Theriogenology 32, 893.
De Vries, M.J., Van Der Beek, S., Kaal-Lansbergen, L.M.T.E., Ouweltjes, W., Wilmink, J.B.M., 1999. Modeling
of energy balance in early lactation and the effect of energy deficits in early lactation on first detected estrus
postpartum in dairy cows. J. Dairy Sci. 82, 19271934.
Domecq, J.J., Skidmore, A.L., Lloyd, J.W., Kaneene, J.B., 1997a. Relationship between body condition scores
and milk yield in a large dairy herd of high yielding Holstein cows. J. Dairy Sci. 80, 101112.
Domecq, J.J., Skidmore, A.L., Lloyd, J.W., Kaneene, J.B., 1997b. Relationship between body condition scores
and conception at first artificial insemination in a large dairy herd of high yielding Holstein cows. J. Dairy Sci.
80, 113120.
Driancourt, M.A., 1991. Follicular dynamics in sheep and cattle. Theriogenology 35, 5579.
Ducker, M.J., Haggett, R.A., Fisher, W.J., Morant, S.V., Bloomfield, G.A., 1985. The effect of level of feeding
in late pregnancy and around the time of insemination on the reproductive performance of first lactation dairy
heifers. Anim. Prod. 41, 1.
Dunn, T.G., Kaltenbach, C.C., 1980. Nutrition and the postpartum interval of the ewe, sow and cow. J. Anim. Sci.
51 (Suppl. 2), 2937.
Dziuk, P.J., Bellows, R.A., 1983. Management of reproduction in beef cattle, sheep and pigs. J. Anim. Sci.
57 (Suppl. 2), 355362.
Echternkamp, S.E., Ferrell, C.L., Rone, J.D., 1982. Influence of pre-and post-partum nutrition on LH secretion in
suckled postpartum beef heifers. Theriogenology 18, 283295.
Edmonson, A.J., Lean, I.J., Weaver, L.D., Farver, T., Webster, G., 1989. A body condition scoring chart for Holstein
dairy cows. J. Dairy Sci. 72, 6878.
Entwistle, K.W., 1983. Factors influencing reproduction in beef cattle in Australia. Aust. Meat Res. Committ. Rev.
43, 30.
Escobar, F.J., Galina, H.C., Fernndez-Baca, S., Jara, S.L., 1982a. Estudio de la actividad reproductiva postparto en
vacas Ceb, criollas y F1 (Ceb/Holstein). AMMVEB (Ed.), VIII Congreso Nacional de Buiatra, pp. 213248.
Escobar, F.J., Fernndez-Baca, S., Galina, C.S., Saltiel, A., Berruecos, J.M., 1982b. Estudio del intervalo entre
partos en bovinos productores de carne en una explotacin del altiplano y otra de la zona tropical hmeda. Vet.
Mx. 13, 5360.
Eversbusch, J.Ch., 1978. ndices reproductivos en un hato de ganado Charolais comparado con Ceb (raza
Brahman) y sus cruzas en el trpico mexicano. Tesis Licenciatura. FMVZ, UNAM, Mxico.
Fallas, M.R., Zarco, Q.L., Galina, C.S., Basurto, H., 1987. Efecto del amamantamiento sobre la actividad ovrica
posparto en vacas F1 (Holstein Indobrasil) en dos tipos de pasto. INIFAP (Ed.), Reunin de Investigacin
Pecuaria en Mxico, pp. 348349.
Ferguson, J.D., 1991. Nutrition and reproduction in dairy cows. Vet. Clin. N. Am. Food Anim. Pract. 7, 483507.
Ferguson, J.D., Otto, K.A., 1989. Managing Body Condition in Dairy Cows. Cornell Univ. (Ed.), Proc. Cornell
Nutr. Conf. Feed Manuf., pp. 75.
Fortune, J.E., 1993. Follicular dynamics during the bovine estrous cycle: a limiting factor in improvement of
fertility? Anim. Reprod. Sci. 33, 111125.
Fortune, J.E., 1994. Ovarian follicular growth and development in mammals. Biol. Reprod. 50, 225232.
Frisch, J.E., Munro, R.K., ONeill, C.J., 1987. Some factors related to calf crops of Brahman, Brahman crossbred
and Hereford Shorthorn cows in a stressful tropical environment. Anim. Reprod. Sci. 15, 126.
Fronk, T.J., Schultz, L.H., Hardie, A.R., 1980. Effect of dry period overconditioning on subsequent metabolic
disorders and performance of dairy cows. J. Dairy Sci. 63, 1080.
Frood, M.J., Croxton, D., 1978. The use of condition scoring in dairy cows and its relationship with milk yield
and live weight. Anim. Prod. 27, 285.
Gaines, J., 1989. The relationship between nutrition and fertility in dairy herds. Food Anim. Pract. Vet. Med. 10,
9971002.
Galina, C.S., Arthur, G.H., 1989a. Review of cattle reproduction in the tropics. 2. Parturition and calving intervals.
Anim. Breed. Abstr. 57, 679686.
Galina, C.S., Arthur, G.H., 1989b. Review of cattle reproduction in the tropics. 3. Puerperium. Anim. Breed. Abstr.
57, 889910.
Galina, C.S., Murcia, C., Beatty, A., Navarro-Fierro, R., Porras, A., 1989. Reproductive performance of Zebu
cattle in Mexico. FAO/IAEA (Eds.), Proceedings, Final Research Coordination Meeting of the FAO/IAEA
Coordinated Research Programme entitled Regional network for improving the reproductive management of
milk and meat-producing livestock in Latin America with the aid of radio-immunoassay techniques. Bogot,
Colombia.
Gallo, L., Carnier, P., Cassandro, M., Mantovani, R., Bailoni, L., Contiero, B., Bittante, G., 1996. Change in body
condition score of Holstein cows as affected by parity and mature equivalent milk yield. J. Dairy Sci. 79,
10091015.
Garca-Winder, M.J., 1990. Efecto de la nutricin sobre la reproduccin. INIFAP (Ed.), Reunin Anual de
Investigacin Pecuaria, Tabasco, Mxico, pp. 573580.
Garnsworthy, P.C., Topps, J.H., 1982. The effect of body condition of dairy cows at calving on their food intake
and performance when given complete diets. Anim. Prod. 35, 113.
Gearhart, M.A., Curtis, C.R., Erb, H.N., Smith, R.D., Sniffen, C.J., Chase, L.E., Cooper, M.D., 1990. Relationship
of changes in condition score to cow health in Holsteins. J. Dairy Sci. 73, 31323140.
Gillund, P., Reksen, O., Grhn, Y.T., Karlberg, K., 2001. Body condition related to ketosis and reproductive
performance in Norwegian dairy cows. J. Dairy Sci. 84, 13901396.
Gluckman, P.D., Breier, B.H., Davis, S.R., 1987. Physiology of the somatotropic axis with particular reference to
the ruminant. J. Dairy Sci. 70, 442446.
Griffith, M.K., Williams, G.L., 1994. Role of visual and olfactory cues during suckling in control of maternal
behavior, lactation and suckling-mediated inhibition of LH secretion in beef cows. J. Anim. Sci. 72 (Suppl. 1),
366.
Griffith, M.K., Williams, G.L., 1996. Roles of maternal vision and olfaction in suckling-mediated inhibition of
luteinizing hormone secretion, expression of maternal sensitivity, and lactational performance of beef cows.
Biol. Reprod. 54, 761768.
Haresign, W., 1984. Underfeeding and reproduction: physiological mechanisms. INRA (Ed.), Reproduction des
ruminants en zone tropicale. Reunion Internationale, Ponit-a-Pitre, Guadeloupe Paris, France, pp. 339345.
Heuer, C., Schukken, Y.H., Dobbelaar, P., 1999. Postpartum body condition score and results from the first test day
milk as predictors of disease, fertility, yield, and culling in commercial dairy herds. J. Dairy Sci. 82, 295304.
Hinojosa, A., Franco, A., Bolio, I., 1980. Genetic and environmental factors affecting calving interval in a
commercial herd in a semi-humid tropical environment. Trop. Anim. Prod. 5, 165171.
Holness, D.H., 1984. The effects of pre- and post-partum levels of nutrition on fertility in cattle. INRA (Ed.),
Reproduction des ruminants en zone tropicale. Reunion Internationale, Ponit-a-Pitre, Guadeloupe Paris, France,
pp. 379388.
Houghton, P.L., Lemenager, R.P., Moss, G.E., Hendrix, K.S., 1990. Prediction of postpartum beef cow body
composition using weight to height ratio and visual body condition score. J. Anim. Sci. 68, 14291437.
Hudson, S.J., Mullord, M.M., 1977. Investigations of maternal bonding in dairy cattle. Appl. Anim. Ethol. 3,
271276.
Hunter, R.A., Magner, T., 1988. The effect of supplements of formaldehyde-treated casein on the partitioning of
nutrients between cow and calf in lactating Bos indicus Bos taurus heifers fed a roughage diet. Austr. J.
Agric. Res. 39, 11511162.
Imakawa, K., Day, M.L., Zalesky, D.D., Clutter, A., Kittok, R.J., Kinder, J.E., 1987. Effects of 17beta-estradiol
and diets varying in energy on secretion of luteinizing hormone in beef heifers. J. Anim. Sci. 64, 805815.
Jaeger, J.R., Olson, K.C., Corah, L.R., Beal, W.E., 1995. Prostaglandin F2 and naloxone therapy in the anoestrus
postpartum beef cows. Theriogenology 43, 657666.
Jolly, P.D., 1992. Physiological and Nutritional Aspects of Postpartum Acyclicity in Bos indicus Cows. Ph.D.
Thesis, James Cook University of North Queensland, Townsville.
Jolly, P.D., McDougall, S., Fitzpatrick, L.A., Macmillan, K.L., Entwistle, K.W., 1995. Physiological effects of
undernutrition on postpartum anoestrus in cows. J. Reprod. Fert. 49 (Suppl.), 477492.
Jolly, P.D., McSweeney, C.S., DOcchio, M.J., Entwistle, K.W., 1991. Nutritional, suckling and ovarian feedback
effects on plasma LH and FSH during post-partum anoestrus in Brahman-cross cows. Proc. Austr. Soc. Reprod.
Biol. 23, 152.
Jolly, P.D., McSweeney, C.S., Schlink, A.C., Houston, E.M., Entwistle, K.W., 1996. Reducing post-partum
anoestrous interval in first-calf Bos indicus crossbred beef heifers. III. Effect of nutrition on responses to
weaning and associated variation in metabolic hormone levels. Austr. J. Agric. Res. 47, 927942.
Jones, G.P., Garnsworthy, P.C., 1988. The effects of body condition at calving and dietary protein content on dry
matter intake and performance in lactating dairy cows given diets of low energy content. Anim. Prod. 47, 321.
Kiser, T.E., Dunlap, S.E., Benyshek, L.L., Mares, S.E., 1980. The effect of calf removal on estrous response and
pregnancy rate of beef cows after Syncro-Mate B treatment. Theriogenology 13, 381389.
Laflamme, L.F., Connor, M.L., 1992. Effect of postpartum nutrition and cow body condition at parturition on
subsequent performance of beef cattle. Can. J. Anim. Sci. 72, 843851.
Lamb, G.C., Lynch, J.M., Greiger, D.M., Minton, J.E., Stevenson, J.S., 1997. Ad libitum suckling by unrelated calf
in the presence or absence of a cows own calf prolongs postpartum anovulation. J. Anim. Sci. 75, 27622769.
Lamming, G.E., Wathes, D.C., Peters, A.R., 1981. Endocrine patterns in the post-partum cow. J. Reprod. Fert.
30 (Suppl.), 155170.
Lermite, V., Terqui, M., 1991. Plasma sex steroid-binding protein in mature heifers: effects of reproductive status,
nutritional level, and porcine growth hormone, and estradiol-17 treatments. Biol. Reprod. 44, 864870.
Lishman, A.W., Harwin, G.O., 1985. Failure to induce ovulation by short-term calf removal in lactating beef cows
on dry-lot. South African J. Anim. Sci. 15, 2122.
Loudon, A., 1987. Nutritional effects on puberty and lactational infertility in mammals: some interspecies
considerations. Proc. Nutr. Soc. 46, 203216.
Lowman, B.G., Scott, N.A., Somerville, S.H., 1976. Condition scoring of cattle. East of Scotland College of
Agriculture (Ed.), Bull. No. 6, Animal Production, Advisory Development Department, UK.
Mackinnon, M.J., Hetzel, D.J.S., Taylor, J.F., 1989. Genetic and environmental effects on the fertility of beef cattle
in a tropical environment. Austr. J. Agric. Res. 40, 10851097.
Macmillan, K.L., 1983. Postpartum interval to oestrus in monozygous twin cows and possible effects of maternal
bonding. N. Z. J. Agric. Res. 26, 451454.
Macmillan, K.L., Clayton, D.G., 1980. Factors influencing the interval to post-partum oestrus, conception date
and empty rate in an intensively managed dairy herd. Proc. N. Z. Soc. Anim. Prod. 40, 236239.
Malven, P.V., Parfet, J.R., Gregg, D.W., Allrich, R.D., Moss, G.E., 1986. Relationships among concentrations of
opioid neuropeptides and luteinizing hormone in neural tissues of beef cows following early weaning. J. Anim.
Sci. 62, 723733.
Maree, C., Jurriaanse, A., Venter, H.A.W., 1974. The occurrence of post partum anoestrus in Bonsmara cows on
supplemented sourveld grazing. South African J. Anim. Sci. 4, 181183.
Markusfeld, O., Galon, N., Ezra, E., 1997. Body condition score, health, yield and fertility in dairy cows. Vet.
Rec. 141, 6772.
Martnez, J.C., Castillo, S.P., 1995. Condicin corporal y gestacin en vacas cebunas bajo condiciones de pastoreo
en el trpico seco. Avances en Investigacin Pecuaria 4, 6368.
McClure, T.J., 1970. An experimental study of the causes of a nutritional and lactational stress infertility of
pasture-fed cows, associated with loss of body weight at about the time of mating. Res. Vet. Sci. 11, 247254.
McDougall, S., Burke, C.R., Macmillan, K.L., Williamsom, N.B., 1995a. Patterns of follicular development during
periods of anovulation in pasture-fed dairy cows after calving. Res. Vet. Sci. 58, 212216.
McDougall, S., Burke, C.R., Macmillan, K.L., Williamson, N.B., 1995b. Follicle patterns during extended periods
of postpartum anovulation in pasture-fed dairy cows. Res. Vet. Sci. 58, 212216.
McIvor, J.G., 1981. Seasonal changes in the growth, dry matter distribution and herbage quality of three native
grasses in northern Queensland. Aust. J. Exp. Agric. Anim. Husbandry 21, 600609.
McNatty, K.P., Heath, D.A., Henderson, K.M., Lun, S., Hurst, P.R., Ellis, L.M., Montgomery, G.W., Morrison, L.,
1984. Some aspects of thecal and granulosa cell function during follicular development in the bovine ovary. J.
Reprod. Fertil. 72, 3953.
McShane, T.M., Petersen, S.L., McCrone, S., Keisler, D.H., 1993. Influence of food restriction on neuropeptide-Y,
pro-opiomelanocortin, and luteinizing hormone-releasing hormone gene expression in sheep hypothalami. Biol.
Reprod. 49, 831839.
McSweeney, C.S., Fitzpatrick, L.A., DOcchio, M.J., Reid, D., Entwistle, K.W., 1993. Reducing post-partum
anoestrous interval in first-calf Bos indicus crossbred beef heifers. I. Effects of pre and post-partum
supplementation strategies. Aust. J. Agric. Res. 44, 10631077.
McSweeney, C.S., Kennedy, P.M., DOcchio, M.J., Fitzpatrick, L.A., Reid, D., Entwistle, K.W., 1993a. Reducing
post-partum anoestrous interval in first-calf Bos indicus crossbred beef heifers. II. Responses to weaning and
supplementation. Aust. J. Agric. Res. 44, 10791092.
McVey Jr., W.R., Williams, G.L., 1991. Mechanical masking of neurosensory pathways at the calf teat interface:
endocrine reproductive and lactational features of the suckled anestrous cow. Theriogenology 35, 931994.
Miner, J.L., 1992. Recent advances in the central control of intake in ruminants. J. Anim. Sci. 70, 12831289.
Montgomery, G.W., Scott, I.C., Hudson, N., 1985. An interaction between season of calving and nutrition on the
resumption of ovarian cycles in post-partum beef cattle. J. Reprod. Fert. 73, 4550.
Montiel, F., 2001. Actividad ovrica post-parto en bovinos de doble propsito en el trpico hmedo mexicano.
Tesis Doctoral, FMVZ, UNAM, Mxico.
Montoni, D., Riggs, J.K., 1978. Efecto del amamantamiento limitado sobre el comportamiento productivo y
reproductivo de un rebao Brahman. Agronoma Tropical. 28, 551571.
Moore, C.P., 1984. Early weaning for increased reproduction rates in tropical beef cattle. World Anim. Rev. 49,
3950.
Moore, C.P., Campos da Rocha, C., 1983. Reproductive performance of Gyr cows: the effect of weaning age of
calves and postpartum energy intake. J. Anim. Sci. 57, 807814.
Moro, J., Castaeda, O.G., Ruiz, F., Romn, H., 1994. Aplicacin de un sistema de registro de la produccin en
ganaderas de doble propsito. INIFAP (Ed.), VII Reunin Cientfica del Sector Agropecuario y Forestal del
estado de Veracruz, Veracruz, Mxico.
Morrison, D.G., Spitzer, J.C., Perkins, J.L., 1999. Influence of prepartum body condition score change on
reproduction in multiparous beef cows calving in moderate body condition. J. Anim. Sci. 77, 10481054.
Morrow, D.A., 1976. Fat cow syndrome. J. Dairy Sci. 59, 16251629.
Morrow, D.A., Roberts, S.J., McEntee, K., 1969. Pospartum ovarian activity and involution of the uterus and
cervix in dairy cattle. II. Involution of uterus and cervix. Cornell Vet. 59, 190198.
Mukasa-Mugerwa, E., Tegegne, A., Franceschini, R., 1991. Influence of suckling and continuous cowcalf
association on the resumption of post-partum ovarian function in Bos indicus cows monitored by plasma
progesterone profiles. Reprod. Nutr. Dev. 71, 241247.
Murphy, M.G., Boland, M.P., Roche, J.F., 1990. Pattern of follicular growth and resumption of ovarian activity in
post-partum beef suckler cows. J. Reprod. Fertil. 90, 523533.
Nebel, R.L., McGilliard, M.L., 1993. Interactions of high milk yield and reproductive performance in dairy cows.
J. Dairy Sci. 76, 32573268.
Nett, T.M., 1987. Function of the hypothalamichypophyseal axis during the post-partum period in ewes and cows.
J. Reprod. Fert. 34 (Suppl.), 201213.
Nett, T.M., Cermak, D., Braden, T., Manns, J., Niswender, G., 1988. Pituitary receptors for GnRH and estradiol
and pituitary content of gonadotropins in beef cows. II. Changes during the postpartum period. Dom. Anim.
Endocrinol. 5, 8189.
Nolan, C.J., Bull, R.C., Sasser, R.G., Rudder, C.A., Panlasigui, P.M., Schoeneman, H.M., Reeves, J.J., 1988.
Postpartum reproduction in protein restricted beef cows: effect on the hypothalamicpitutitaryovarian axis. J.
Anim. Sci. 66, 32083217.
Otto, K.L., Ferguson, J.D., Fox, D.G., Sniffen, C.J., 1991. Relationship between body condition score and
composition of ninth to eleventh rib tissue in Holstein dairy cows. J. Dairy Sci. 74, 852861.
Oyedipe, E.O., Buvanendran, V., Eduvie, L.O., 1982. Some factors affecting the reproductive performance of
White Fulani (Bunaji) cattle. Trop. Agric. (Trinidad) 59, 231234.
Perry, R.C., Corah, L.R., Cochran, R.C., Beal, W.E., Stevenson, J.S., Minton, J.E., Simms, D.D., Brethour, J.R.,
1991. Influence of dietary energy on follicular development, serum gonadotrophins, and first postpartum
ovulation in suckled beef cows. J. Anim. Sci. 69, 37623773.
Peters, A.R., 1984. Reproductive activity of the cow in the post-partum period. I. Factors affecting the length of
the post-partum acyclic period. Br. Vet. J. 140, 7684.
Pimentel, C.A., Deschamps, J.C., Oliveira, J.A.F., Cardelino, R., Pimentel, M.A., 1979. Effects of early weaning
on reproductive efficiency in beef cows. Theriogenology 11, 421427.
Prado, R., Rhind, S.M., Wright, I.A., Russel, A.J.F., McMillen, S.R., Smith, A.J., McNeilly, A.S., 1990. Ovarian
follicle populations, steroidogenicity and micromorphology at 5 and 9 weeks postpartum in beef cows in two
levels of body condition. Anim. Prod. 51, 103108.
Pullan, N.B., 1979. Productivity of White Fulani cattle on the Jos Plateau, Nigeria. I. Herd structures and
reproductive performance. Trop. Anim. Hlth. Prod. 11, 231238.
Rae, D.O., Kunkle, W.E., Chenoweth, P.J., Sand, R.S., Tran, T., 1996. Body condition: influences on beef cattle
reproductive performance. AMMVEB (Ed.), XX Congreso Nacional de Buiatra, Acapulco, Gro., Mxico,
pp. 339343.
Ramrez-Iglesia, L., Soto Balloso, E., Gonzalez Stangnaro, C., Soto Castillo, G., Rincon Urdaneta, E., 1992.
Factors affecting postpartum ovarian activity in crossbred primiparous tropical heifers. Theriogenology 38,
449460.
Randel, R.D., 1981. Effect of once-daily suckling on postpartum interval and cowcalf performance of first-calf
Brahman Hereford heifers. J. Anim. Sci. 53, 755757.
Randel, R.D., 1990. Nutrition and postpartum rebreeding in cattle. J. Anim. Sci. 68, 853862.
Randel, R.D., Welker, G.A., 1976. Once daily suckling effect on calfcow performance. J. Anim. Sci. 43, 301312.
Rasby, R.J., Wettemann, R.P., Harms, P.G., Lusby, K.S., Wagner, J.J., 1992. GnRH in the infundibular stalk-median
eminence is related to percentage body fat in carcasses of beef cows. Domest. Anim. Endocrin. 9, 7176.
Richards, M.W., Spitzer, J.C., Warner, M.B., 1986. Effect of varying levels of postpartum nutrition and body
condition at calving on subsecuent reproductive performance in beef cattle. J. Anim. Sci. 62, 300306.
Richards, M.W., Wettemann, R.P., Schoenemann, H.M., 1989. Nutritional anestrus in beef cows: body weight
change, body condition, luteinizing hormone in serum and ovarian activity. J. Anim. Sci. 67, 15201526.
Richards, M.W., Wettemann, R.P., Spicer, L.J., Morgan, G.L., 1991. Nutritional anestrus in beef cows: effects of
body condition and ovariectomy on serum luteinizing hormone and insulin-like growth factor-1. Biol. Reprod.
44, 961966.
Roberson, M.S., Stumpf, T.T., Wolfe, M.W., Cupp, A.S., Kojima, N., Werth, L.A., Kittok, R.J., Kinder, J.E., 1992.
Circulating gonadotrophins during restricted energy intake in relation to body condition in heifers. J. Reprod.
Fert. 96, 461469.
Robinson, J.J., 1990. Nutrition in the reproduction of farm animals. Nutr. Res. Rev. 3, 267268.
Roche, J.F., Crowe, M.A., Boland, M.P., 1992. Postpartum anoestrus in dairy and beef cows. Anim. Reprod. Sci.
28, 371378.
Ruas, J.R.M., Saturnino, H.M., Machado, G.V., Paulino, M.F., Effect of suckling on the incidence of oestrus
durintg the breeding season in Nelore cows. In: Proceedings of the IX Congresso Brasileiro de Reproducao
Animal. Belo Horizonte, Brasil, vol. 2, 1991, pp. 377384.
Ruegg, P.L., Milton, R.L., 1995. Body condition scores of Holstein cows on Prince Edward Island, Canada:
relationships with milk yield, reproductive performance, and disease. J. Dairy Sci. 78, 552564.
Ruegg, P.L., Goodger, W.J., Holmberg, C.A., Weaver, L.D., Huffman, M.E., 1992. Relation among body condition
score, serum urea nitrogen and cholesterol concentrations, and reproductive performance in high producing
Holstein dairy cows in early lactation. Am. J. Vet. Res. 53, 1014.
Rutter, L.M., Manns, J.G., 1991. Insulin-like growth factor-1 in follicular development and function in postpartum
beef cows. J. Anim. Sci. 69, 11401146.
Rutter, L.M., Randel, R.D., 1984. Postpartum nutrient intake and body condition: effect on pituitary function and
onset of estrus in beef cattle. J. Anim. Sci. 58, 265274.
Ryan, D.P., Spoon, R.A., Griffith, M.K., Williams, G.L., 1994. Ovarian follicular recruitment, granulosa cell
steroidogenic potential and growth hormone/insulin-like growth factor-1 relationships in suckled beef cows
consuming high lipid diets: effects of graded differences in body condition maintained during the puerperium.
Dom. Anim. Endocrinol. 11, 161174.
Salcedo, E., Gonzlez-Padilla, E., Rodrguez, O.L., Ramos, F., 1977. Efecto del destete precoz en el
comportamiento reproductivo de vacas empadradas en agostadero. Tc. Pec. Mx. 3, 3640.
Savio, J.D., Boland, M.P., Hynes, N., Roche, J.F., 1990. Resumption of follicular activity in the early postpartum
period of dairy cows. J. Reprod. Fertil. 88, 569579.
Schallenberger, E., Prokopp, S., 1985. Gonadotrophins and ovarian steroids in cattle. IV. Re-establishment of the
stimulatory feedback action of oestradiol-17-beta on LH and FSH. Acta Endocrinol. 109, 4449.
Schillo, K.K., 1992. Effects of dietary energy on control of luteinizing hormone secretion in cattle and sheep. J.
Aim. Sci. 70, 12711282.
Schlink, A.C., Gibson, D.S., Liang, Z.J., Dixon, R., 1988. Calf management strategies and reproductive
performance in a northern Australian cattle herd. Proc. Austr. Soc. Anim. Prod. 17, 326329.
Schlink, A.C., Houston, E.M., Entwistle, K.W., 1994. Impact of long term early weaning on the productivity of
Bos indicus-cross cows. Proc. Austr. Soc. Anim. Prod. 20, 339.
Schottler, J.H., Williams, W.T., 1975. The effect of early weaning of Brahman cross calves on calf growth and
reproductive performance of the dam. Austr. J. Exp. Agric. Anim. Husb. 15, 456459.
Schramm, R.D., Roberge, S., Reeves, J.J., 1991. Enclomiphene does not alter the postpartum interval of suckled
beef cows. J. Anim. Sci. 69, 41124116.
Selk, G.E., Wettemann, R.P., Lusby, K.S., Oltjen, J.W., Mobley, S.L., Rasby, R.J., Garmendia, J.C., 1988.
Relationship among weight change, body condition and reproductive performance of range beef cows. J.
Anim. Sci. 66, 31533159.
Senatore, E.M., Butler, W.R., Oltenacu, P.A., 1996. Relationships between energy balance and post-partum ovarian
activity and fertility in first lactation in dairy cows. J. Anim. Sci. 62, 1723.
Shively, T.E., Williams, G.L., 1987. Premature calf return attenuates neuroendocrine and ovarian response to
temporary weaning in anestrous cows. J. Anim. Sci. 65 (Suppl. 1), 432.
Shively, T.E., Williams, G.L., 1989. Patterns of tonic luteinizing hormone release and ovulation frequency in
suckled anestrous beef cows following varying intervals of temporary weaning. Dom. Anim. Endocrinol. 6,
379387.
Short, R.E., Adams, D.C., 1988. Nutritional and hormonal interrelationships in beef cattle reproduction. Can. J.
Anim. Sci. 68, 29.
Short, R.E., Bellows, R.A., Moody, E.L., Howland, B.E., 1972. Effects of suckling and mastectomy on bovine
postpartum reproduction. J. Anim. Sci. 34, 7074.
Short, R.E., Bellows, R.A., Staigmiller, R.B., Custer, E.E., 1990. Physiological mechanisms controlling anestrus
and infertility in postpartum beef cattle. J. Anim. Sci. 68, 799816.
Silveira, P.A., Spoon, R.A., Ryan, D.P., Williams, G.L., 1993. Evidence of maternal behaviour as a requisite link
in suckling-mediated anovulation in cows. Biol. Reprod. 49, 13381346.
Smith, M.F., Burrell, W.C., Shipp, L.D., Sprott, L.R., Songster, W.N., Wiltbank, J.N., 1979. Hormone treatments
and use of calf removal in postpartum beef cows. J. Anim. Sci. 48, 12851294.
Smith, M.F., Lishman, A., Lewis, G., Harms, P., Ellersick, M., Inskeep, E., Wiltbank, J., Amos, M., 1983. Pituitary
and ovarian responses to gonadotropin releasing hormone, calf removal and progestogen in anestrous beef
cows. J. Anim. Sci. 57, 418424.
Spicer, L..J., Tucker, W.B., Adams, G.D., 1990. Insulin-like growth factor-I in dary cows: relationships among
energy balance, body condition, ovarian activity, and estrous behavior. J. Dairy Sci. 73, 929937.
Stagg, K., Diskin, M.G., Sreenan, J.M., Roche, J.F., 1995. Follicular development in long-term anoestrus suckler
beef cows fed two levels of energy postpartum. Anim. Reprod. Sci. 38, 4961.
Stagg, K., Spicer, L.J., Sreenan, J.M., Roche, J.F., Diskin, M.G., 1998. Effect of calf isolation on follicular wave
dynamics, gonadotropin and metabolic hormone changes, and interval to first ovulation in beef cows fed either
of two energy levels postpartum. Biol. Reprod. 59, 777783.
Staples, C.R., Thatcher, W.W., Clark, J.H., 1990. Relationship between ovarian activity and energy status during
early lactation in high producing dairy cows. J. Dairy Sci. 73, 938947.
Stevenson, J.S., Knoppel, E.L., Minton, J.E., Salfen, B.E., Garverick, H.A., 1994. Estrus, ovulation, luteinizing
hormone, and suckling-induced hormones in mastectomized cows with and without unrestricted presence of
the calf. J. Anim. Sci. 72, 690699.
Stevenson, J.S., Lucy, M.C., Call, E.P., 1987. Failure of timed inseminations and associated luteal function in dairy
cattle after two injections of prostaglandin F2 . Theriogenology 28, 937946.
Sullivan, R.M., ORourke, P.K., Robertson, D.L., Cooke, D., 1992. Effects of once-yearly weaning on some aspects
of herd productivity in an extensive herd in the semi-arid tropics of northern Australia. Austr. J. Exp. Agric.
32, 149156.
Tannenbaum, G.S., Guyda, H.J., Posner, B.I., 1983. Insulin-like growth factors: a role in growth hormone negative
feedback and body weight regulation via brain. Science 220, 7779.
Tegegne, A., Entwistle, K.W., Mukasa-Mugerwa, E., 1992. Effects of supplementary feeding and suckling intensity
on postpartum reproductive performance of Small East African Zebu Cows. Theriogenology 38, 97106.
Tinker, E.D., Buchanan, D.S., Wettemann, R.P., Frahm, R.R., Knori, L.W., 1989. Effect of breed combination and
body condition score on postpartum cyclicity of various two-breed-cross cows. J. Anim. Sci. 67 (Suppl. 1),
432438.
Troxel, T.R., Kesler, D.J., Noble, R.C., Carlin, S.E., 1980. Ovulation and reproductive hormones following steroid
pretreatment, calf removal and GnRH in postpartum suckled beef cows. J. Anim. Sci. 51, 652659.
Vaccaro, R., 1990. Comportamiento de bovinos para doble propsito en el trpico, Seminario Internacional sobre
Lechera Tropical. Villahernosa, Tabasco, Mxico, pp. 1435.
Viker, S.D., Larson, R.L., Kiracofe, G.H., Stewart, R.E., Stevenson, J.S., 1993. Prolonged postpartum anovulation
in mastectomized cows requires tactile stimulation by the calf. J. Anim. Sci. 71, 9991003.
Viker, S.D., McGuire, W.J., Wright, J.M., Beeman, K.B., Kiracofe, G.H., 1989. Cowcalf association delays
postpartum ovulation in mastectomized cows. Theriogenology 32, 467474.
Villa-Godoy, A., Hughes, T.L., Emery, R.S., Chapin, L.T., Fogwell, R.L., 1988. Association between energy
balance and luteal function in lactating dairy cows. J. Dairy Sci. 71, 10631072.
Villa-Godoy, A., Hughes, T.L., Emery, R.S., Stanisiewski, E.P., Fogwell, R.L., 1990. Influence of energy balance
and body condition on estrus and estrous cycles in Holstein heifers. J. Dairy Sci. 73, 27592765.
Wagner, J.J., Lusby, K.S., Oltjen, J.W., Rakestraw, J., Wettemann, R.P., Walters, L.E., 1988. Carcass composition
in mature Hereford cows: estimation and effect on daily metabolizable energy requirement during winter. J.
Anim. Sci. 66, 603615.
Walters, D.L., Kaltenbach, C.C., Dunn, T.G., Short, R.E., 1982. Pituitary and ovarian function in postpartum beef
cows. I. Effect of suckling on serum and follicular fluid hormones and follicular gonadotropin receptors. Biol.
Reprod. 26, 640646.
Waltner, S.S., McNamara, J.P., Hillers, J.K., 1993. Relationships of body condition score to production variables
in high producing Holstein dairy cattle. J. Dairy Sci. 76, 34103419.
Weaver, L.D., 1992. Condicin corporal, produccin y reproduccin. Carta Ganadera Marzo, pp. 1318.
Weesner, G.D., Norris, R.A., Forrest, D.W., Harms, P.G., 1987. Biological activity of luteinizing hormone in the
peripartum cow: least activity at parturition with an increase throughout the postpartum interval. Biol. Reprod.
37, 851858.
Wells, P.L., Holness, D.H., Reymark, P.J., McCabe, C.T., Lishman, A.W., 1985. Fertility in the Afrikander cow. 2.
Ovarian recovery and conception in suckled and non-suckled cows postpartum. Anim. Reprod. Sci. 8, 315326.
Wettemann, R.P., 1994. Precalving nutrition/birth weight interaction and rebreeding efficiency. Oklahoma State
University (Ed.), Anim. Sci. Res. Report.
Wettemann, R.P., Hill, G.N., Boyd, M.E., Spitzer, J.C., Forrest, D.W., Beal, W.E., 1986. Reproductive performance
of postpartum beef cows after short-term calf separtion and dietary energy and protein suplementation.
Theriogenology 26, 433443.
Wettemann, R.P., Turman, E.J., Wyatt, R.D., Totusek, R., 1978. Influence of suckling intensity on reproductive
performance of range cows. J. Anim. Sci. 47, 342346.
Whisnant, C.S., Kiser, T.E., Thompson, F.N., Barb, C.R., 1986. Naloxone infusion increases pulsatile luteinizing
hormone release in postpartum beef cows. Dom. Anim. Endocrinol. 3, 4954.
Wildman, E.E., Jones, G.M., Wagner, P.E., Bowman, R.L., Troutt, H.F., Lesch, T.N., 1982. A dairy cow body
condition scoring system and its relationship to selected production characteristics. J. Dairy Sci. 65, 495.
Williams, G.L., 1989. Modulation of luteal activity in postpartum beef cows through changes in dietary lipid. J.
Anim. Sci. 67, 785793.
Williams, G.L., 1990. Suckling as a regulator of postpartum rebreeding in cattle: a review. J. Anim. Sci. 68,
831852.
Williams, G.L., Griffith, M.K., 1995. Sensory and behavioural control of gonadotrophin secretion during
suckling-mediated anovulation in cows. J. Reprod. Fertil. 49 (Suppl.), 463475.
Williams, G.L., Kirsch, J.D., Post, G.R., Tilton, J.E., Slanger, W.D., 1984. Evidence against chronic teat stimulation
as an autonomous effector of diminished gonadotrophin release in beef cows. J. Anim. Sci. 59, 10601069.
Williams, G.L., Kotwica, J., Slanger, W.D., Olson, D.K., Tilton, J.E., Johnson, L.J., 1982. Effect of suckling on
pituitary responsiveness to gonadotropin-releasing hormone throughout the early postpartum period of beef
cows. J. Anim. Sci. 54, 592602.
Williams, G.L., Koziorowski, J., Osborn, R.G., Kirsch, J.D., Slanger, W.D., 1987. The postweaning rise of tonic
luteinizing hormone secretion in anestrous cows is not prevented by chronic milking or the physical presence
of the calf. Biol. Reprod. 36, 10791084.
Williams, G.L., McVey, W.R., Hunter, J.F., 1993. Mammary somatosensory pathways are not required for
suckling-mediated inhibition of luteinizing hormone secretion and ovulation in cows. Biol. Reprod. 49, 1328
1337.
Williams, G.L., Talavera, F., Petersen, B.J., Kirsch, J.D., Tilton, J.E., 1983. Coincident secretion of
follicle-stimulating hormone and luteinizing hormone in early postpartum beef cows: effects of suckling and
low-level increases of systemic progesterone. Biol. Reprod. 29, 362373.
Wiltbank, J.N., Rowden, W.W., Ingalls, J.E., Zimmerman, D.R., 1964. Influence of postpartum energy level on
reproductive performance of Hereford cows restricted in energy intake before calving. J. Anim. Sci. 23, 1049
1053.
Wright, I.A., Rhind, S.M., Russel, A.J.F., Whyte, T.K., McBean, A.J., McMillen, S.R., 1987. Effects of body
condition, food intake and temporary calf separation on the duration of the post-partum anoestrus period and
associated LH, FSH and prolactin concentrations in beef cows. Anim. Prod. 45, 395402.
Wright, I.A., Rhind, S.M., Whyte, T.K., Smith, A.J., 1992. Effects of body condition at calving and feeding level
after calving on LH profiles and the duration of the post-partum anoestrous period in beef cows. Anim. Prod.
55, 4146.
Wright, I.A., Rhind, S.M., Whyte, T.K., Smith, A.J., McMillen, S.R., Prado, R., 1990. Circulating concentrations
of LH and FSH and pituitary responsiveness to GnRH in intact and ovariectomized suckled beef cows in two
levels of body condition. Anim. Prod. 51, 93101.
Wright, P.J., Malmo, J., 1992. Pharmacologic manipulation of fertility. Vet. Clin. North. Am.: Food Anim. Pract.
8, 5789.
Yelich, J.V., Holland, M.D., Schutz, D.N., Odde, K.G., 1995. Synchronization of estrus in suckled postpartum
beef cows with melengestrol acetate, 48-h calf removal and PGF2 . Theriogenology 43, 401410.
Zalesky, D.D., Forrest, D.W., McArthur, N.H., Wilson, J.M., Morris, D.L., Harms, P.G., 1990. Suckling inhibits
release of luteinizing hormone-releasing hormone from the bovine median eminence following ovariectomy.
J. Anim. Sci. 68, 444448.
Zurek, E., Foxcroft, O.R., Kennelly, J.J., 1995. Metabolic status and interval to first ovulation in postpartum dairy
cows. J. Dairy Sci. 78, 19091920.

Body Condition and Suckling As Factor Influencing The Duration of Postpartum Anestrum in Cattle.

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Animal Reproduction Science 85 (2005) 126

Body condition and suckling as factors influencing

Corresponding author. Tel.: +52-29-9342075; fax: +52-29-9344053.

E-mail address: fmontielp@hotmail.com (F. Montiel).

The existence of a relationship between reproduction and nutritional status in cattle

of nutrients relative to metabolic demands, is a major factor contributing to prolonged

2.1. Body condition score evaluation

2.2. Energy balance

2.3. Pre- and postpartum nutrition

2.4. Secretion of hypothalamic and hypophyseal hormones

2.5. Postpartum follicular dynamics

2.6. Pregnancy rate

Prolonged postpartum anestrus is a major limitation to achieve a optimal reproductive

The primary endocrine deficit resulting in the establishment of postpartum anovulation

3.2. Restricted and unrestricted suckling

3.3. Suckling own or unrelated calves

3.4. Role of vision and olfaction

The role of vision and olfaction in suckling-mediated inhibition of LH secretion in cattle

In summary, reproduction is a major component of the productive efficiency for a cowcalf

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