Effect of maternal nutrition on calf health and growth

Full text

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April 23 and 24, 2013 Tri-State Dairy Nutrition Conference
Effect of Maternal Nutrition on Calf Health and Growth
Jon Schoonmaker1
Department of Animal Sciences
Purdue University
Abstract
In livestock, just as in humans, there is
increasing evidence that production characteristics
in livestock may be affected by maternal diet. This
so-called metabolic or developmental
programming of postnatal growth occurs during
certain windows of growth characterized by
plasticity of metabolic regulatory systems, namely
in utero or shortly after birth. Developmental
programming in mammals occurs through
modification of the chromatin structure of
deoxyribonucleic acid (DNA), which is heritable.
A pregnant cows capacity to care for her embryo
is largely determined by the way she partitions
nutrients to support embryonic, placental, and fetal
development together with her own growth,
maintenance, and milk production. Production status
and energy balance can have a significant effect on
how nutrients are partitioned; however, diet during
pregnancy is an important modifiable factor that can
have a substantial influence on the viability and body
composition of the newborn. Nutrition (inadequate
or excess) during early gestation can effect
development of organ systems, including the liver
and pancreas, the mammary gland, gonads, and fat
tissue. The effect of nutrition during early gestation
on birth weight is variable, however, long-term
consequences have been variable. Seventy-five
percent of fetal growth occurs during the last
trimester and nutrition during late gestation can
further impact organ systems, birth weight, and long-
term growth. The first weeks of life for a neonate
seem to also have long-lasting consequences on the
physiological function of the animal. In fact, many
of the effects of fetal programming can be
ameliorated or exacerbated due to nutrition of the
neonate. Thus, management of the cow herd not
only has consequences for cow productivity but also
for productivity of the next generation of cows.
Introduction
Effective nutritional management during the
transition period has long been recognized as having
a major impact on health and productivity of lactating
dairy cows. While it is clear that the tremendous
changes that occur during the transition period
impact the cow, it has increasingly become apparent
that nutrient supply and hormonal signaling during
conception, gestation, or shortly after parturition can
exert permanent changes to metabolism affecting
lifetime performance, body composition, and
metabolic function of the offspring. Thus, it is likely
that the cow of today, with high milk yield but also
reproductive and metabolic challenges, is not only
a consequence of genetic selection, but also the result
of the way her dam was fed and the way she was
fed early after birth (Bach, 2012).
The concept that postnatal growth and
production is sensitive to direct and indirect effects
of maternal nutrition was first proposed by Dr.
Barker and co-workers (Barker et al., 2002) based
on epidemiological studies investigating low nutrient
intake by pregnant mothers experienced during the
1Contact at: 915 W. State St., 3-228 Lilly Hall, West Lafayette, IN 47907; (765) 494-4860; Email: jschoonm@purdue.edu.

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1944 Dutch Famine of World War II and the
resulting long-term health implication of their
children. The hypothesis suggests that if intrauterine
conditions are poor (e.g., poor nutrition), then the
fetus becomes adapted through altered gene
expression to maximize the uptake and utilization of
the nutrients available. Such adaptations cause the
progeny to exhibit a competitive advantage if
exposed to a similarly poor environment in
adulthood. However, favorable postnatal conditions
can challenge the individuals homeostatic
mechanisms and lead to development of deleterious
metabolic conditions, such as obesity, insulin
resistance, and type 2 diabetes (Barker et al.,
2002). Since the inception of the Barker hypothesis,
many terms have been used to categorize this
phenomenon, including fetal programming (Barker
and Clark, 2002), neonatal programming (Spencer
et al., 2011), lactational programming (Hinde and
Capitanio, 2010), metabolic imprinting (Waterland
and Garza, 1999), metabolic programming (Lucas,
2000), and developmental programming (Reynolds
et al., 2010) which encompasses all of these
developmental phases and concepts.
Developmental programming in mammals
occurs through epigenetic changes (Wu et al., 2006).
These modifications are mainly due to alterations in
the chromatin structure through acetylation of
histones or methylation of DNA. As a result, gene
expression continues to be modified throughout life
without alteration of the DNA sequence. Such a
mitotically heritable alteration of gene expression
induces a non-genomic tuning of phenotype through
developmental plasticity (Crews and McLachlan,
2006). These effects can even be passed on to more
than one succeeding generation.
In livestock, just as in humans, there is
increasing evidence that production characteristics
in livestock may be affected by maternal diet (Wu
et al., 2006). Compromised fetal or neonatal growth
in livestock has been shown to lead to increased
neonatal morbidity and mortality, slow postnatal
growth, increased fat deposition, insulin resistance,
and dysfunction of specific organs, including the
ovaries, testes, mammary gland, liver, and small
intestine (Wu et al., 2006). Potential causes and
consequences in dairy cattle have been reviewed
recently (Bach, 2012); however, the potential effects
on metabolic, immune, and reproductive function,
as well as milk performance, of pre- and early
postnatal nutrition and management in dairy cattle
has not been extensively studied.
Maternal Environment and Fetal Growth
In the United States, poor intrauterine
conditions can occur in the cow due to: 1)
competition for nutrients between maternal growth
and metabolic needs and fetal growth, particularly
in peripubertal dams, 2) selection for increased milk
production, in which the increased energy demand
of lactation competes with the increased energy
demand of fetal and placental growth, 3) pregnancy
occurring during periods of high environmental
temperature (e.g., summer to early fall), or 4) over-
nutrition of the dam during gestation during times of
supplemental feeding resulting in rapid maternal
growth and accumulation of maternal adipose tissue.
A pregnant cows capacity to care for her
embryo is largely determined by the way she
partitions nutrients to support embryonic, placental,
and fetal development together with her own growth,
maintenance, and milk production (Banos et al.,
2007). Development of the fetus through gestation
is presented in Figure 1.The peri-conceptual period
typically coincides with peak lactation in a mature
cow and with development of peri-pubertal dams.
Thus, the embryo must compete for nutrients with
the cows nutrient demands for lactation and the
peripubertal animals demands for growth. Changing
nutrient supply during the peri-conceptual period
can have profound impacts on quality of oocytes as
well as development of the embryo. Although
nutrient requirements for embryonic growth are low
during the first trimester, critical events for normal

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fetal development occur, including establishment of
functional fetal and uteroplacental circulation,
organogenesis, myogenesis, and adipogenesis
(Robinson et al., 1999; Du et al., 2010). The first
trimester typically coincides with early to mid-
lactation in a mature cow and with development of
peripubertal dams. A poor uterine environment
during the first trimester can have a significant impact
on development of the mammary gland and
subsequent milk production, development of the
ovaries and reproductive capability, development
of fat tissue stores and ability to maintain body
condition, and development of homeostatic
mechanisms in the liver and pancreas, with an impact
on capacity of the progeny to utilize and metabolize
nutrients (Symonds et al., 2010). Examples of
developmental programming in livestock models are
evident for heart, lung, pancreas, kidney, placenta,
perirenal fat, small intestine, muscle, weaning weight,
and reproductive performance (Caton et al., 2007).
During the second trimester, the fetus
continues to develop and grow, but it will only reach
about 25% of the size it will be at birth (Symonds et
al., 2010); therefore, the nutritional requirements of
the cow do not increase substantially above lactation
requirements during this period. During the second
trimester, the lactating cow may be in positive energy
balance and will be better able to sustain the fetus.
The second trimester coincides with mid-to-late-
lactation and a poor uterine environment may have
an impact on further development of organ systems
(Symonds et al., 2010). The majority of fetal
development has already occurred by the beginning
of the final trimester. However, 75% of fetal growth
(increase in tissue size) occurs during this phase
(Robinson et al., 1977). The final trimester coincides
with late-lactation and the dry period, and a poor
uterine environment during the final trimester
typically impacts birth weight, with the potential to
impact long-term health and growth of the offspring.
Production Status
Dam age
When pregnancy coincides with the
continued growth of the first parity mother, the fetus
may face intense competition for nutrients from its
mothers own metabolic needs. Thus, the normal
hierarchy of nutrient (i.e. protein and energy)
partitioning between body growth and fetal growth
may be altered (Wallace et al., 2006). Age at calving
(parity) manifests the state of development of the
dam during gestation regarding her own growth. For
example, in sheep, progeny from first parity ewes
are lighter than progeny from second parity ewes
(Bradford, 1972; Bradford et al., 1974), but
whether this effect continues with increasing parity
(i.e. > 2 pregnancies) was not clear. Fuerst-Waltl
et al. (2004) reported in Austrian dual-purpose
Simmental cows that as maternal age increased, milk
production and longevity were decreased and days
to first service were increased in female progeny.
Similarly, Banos et al. (2007) observed that
daughters of older dams had lower body condition
score (BCS), produced less daily milk, and needed
more days to first service. Banos et al. (2007) also
observed that daughters from late calving first parity
cows (30 to 36 mo) had a 7% lower BCS, produced
4.5% less milk, and had their first service almost 3
days later than daughters from early calving first
parity cows (18 to 23 mo). In contrast, cows whose
first parity dams calved later needed fewer
inseminations per conception and had fewer returns
to estrus (i.e., a higher proportion conceived at first
insemination). Early-calving cows, therefore, may
produce progeny that mature early, exhibiting the
characteristics of high-producing Holsteins cows,
but they cannot conceive as easily as cows born to
older first-calving dams. Offspring resulting from
early second calvings were associated with
increased production and improved BCS and
fertility (Banos et al., 2007).

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Dam age can also impact progeny immunity.
Research conducted at Colorado State University
(reviewed by Odde, 1988) investigated the
relationship between pre-calving nutrition in beef
cows and disease susceptibility in the neonatal calf.
First-calf heifers produced calves that had lower
levels of serum immunoglobulins at 24 hr of age
than calves born to cows greater than or equal to 3
years of age. This occurred even though colostral
immunoglobulin concentrations were similar for
these 2 age groups. The increased disease
susceptibility observed in calves born to first-calf
heifers was likely due to lower volumes of colostrum
produced by first-calf heifers, although decreased
calf vigor as a result of dystocia may have also
contributed.
Dam milk production
Milk yield is the key competitor to the fetus
for nutrients during gestation and may influence
progeny development indirectly in early gestation,
when fetal nutrient requirements are low, and then
directly later in gestation when they are high. Banos
et al. (2007) reported no significant effects of
maternal milk production during pregnancy on
subsequent offspring milk production in the first
lactation. However, Berry et al. (2008) found a
negative relationship between milk production of
the dam and lactation performance of the offspring
in the first and third, but not second, lactations and
concluded that the majority of the maternal effects
in progeny performance were due to factors other
than maternal milk production. The lack of an
association between milk production and long-term
effects on the offspring may be related to the fact
that cows could have been in various states of energy
balance (i.e. negative, neutral, or positive). Thus, it
may be energy balance, rather than milk yield, that
impacts metabolic function of the offspring (Bach,
2012).
The BCS of a pregnant cow is associated
with the amount of energy available to her to sustain
growth, maintenance, milk production, and fetal
development. A cows BCS level and change during
gestation can be associated with energy balance and
her ability to meet the needs of maintenance,
lactation, and fetal growth, which can potentially
affect the offsprings future performance as a milk-
producing cow. Calves born to thin (<5 body
condition score) two-year-old beef heifers were less
vigorous and had reduced serum immunoglobulin
levels at 24 hr of age (Odde, 1988). Banos et al.
(2007) observed that as dam BCS increased,
daughter BCS and fertility increased, but milk
production was slightly reduced. Thus, it is important
to avoid BCS losses of the dam or rapid increases
in BCS, especially during the second and third
trimesters of gestation. Appropriate nutritional
strategies in late lactation and the dry period become
crucial factors in this respect.
Nutritional Status
Diet during pregnancy is one important
modifiable factor that can have a substantial influence
on the viability and body composition of the
newborn (Symonds et al., 2010). Significant
research in livestock has demonstrated that maternal
energy under- and over-nutrition can impact
adiposity, insulin response, and development of
progeny. Protein status of the mother can also impact
progeny development but has not been as well
studied. With the exception of selenium, which can
increase blood flow in key nutrient-transferring
tissues, including intestine, placenta, and mammary
gland (Grazul-Bilska et al., 2009; Meyer et al.,
2010), information concerning the effect of individual
nutrients, such as amino acids, fatty acids, vitamins,
and minerals, in the maternal diet and their impact
on progeny development is scarce.
Early gestation
It is possible that developmental
programming of offspring begins as early as the peri-
conceptual period. Nutritional status has been

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correlated with oocyte quality and embryo survival
and are key factors influencing reproductive
efficiency (Papadopoulos et al., 2001; Armstrong
et al., 2003; Webb et al., 2004). Conflicting results
have been reported for the effects of dietary energy
density on oocyte quality and early embryonic
development in ruminants. A higher proportion of
ova from ewes on low energy diets were considered
viable compared with those from ewes on high
energy diets (McEvoy et al., 1995). However, sheep
fed low energy diets had embryos with decreased
cleavage rates compared with high energy diets
(Papadopoulos et al., 2001). Grazul-Bilska et al.
(2006) observed no differences in the number of
healthy oocytes collected from superovulated ewes
fed at maintenance energy requirements, ewes fed
at 60% of maintenance energy requirements, and
ewes overfed energy. However, at fertilization, both
the overfed and the restricted ewes had reduced
successful fertilizations, morulas, and blastocysts,
indicating that maternal nutrition (both inadequate
and excess) before mating can have profound
impacts on oocyte quality and fertilization rates.
These observations suggest that nutritional status
during the peri-conceptual period is highly important
to fertility and that some fetal programming may
actually begin in the oocyte before mating occurs.
In fact, lambs of ewes fed at 70% of nutritional
requirements from 30 days prior to mating until 100
days of pregnancy had higher neonatal mortality
rates, were lighter at birth, and grew more slowly
after weaning than those fed at 100% of estimated
requirements (Nordby et al., 1986, 1987). For
cows, positive (Kendrick et al., 1999; Boland et
al., 2001), negative (Armstrong et al., 2001), or no
effects (Tripp et al., 2000) of plane of nutrition (high
vs. low energy diets) on oocyte quality, fertilization
rate, and early embryonic development have been
reported.
The placenta plays a major role in the
regulation of fetal growth. Establishment of functional
fetal and uteroplacental circulation is one of the
earliest events during embryonic and placental
development (Patten, 1964; Ramsey, 1982). It has
been shown that the large increase in transplacental
exchange, which supports the exponential increase
in fetal growth during the last one-half of gestation,
depends primarily on the dramatic growth of the
uteroplacental vascular beds during the first one-
half of pregnancy (Meschia,1983; Reynolds and
Redmer, 1995). In beef cattle, under-nutrition of
~75% of recommended allowance during early
stages of pregnancy compromises placental
angiogenesis, cotyledon weight, and thus, fetal
development (Vonnahme et al., 2007). Upon
realimentation, Vonnahme et al. (2007) observed
that fetuses in both groups were similar in weight
near term. Previous studies with sheep have
demonstrated that an extended period of maternal
nutrient restriction during the first half of gestation
results in relatively normal birth weights but leads to
increases in the length and thinness of the neonate,
increases in adiposity, and suppressed glucose
tolerance (Whorwood et al., 2001; Bispham et al.,
2003; Ford et al., 2007). Gardner et al. (2005)
demonstrated that increased adiposity as a result of
early fetal restriction can persist to 1 year of age.
Overnutrition can also impact placental
development and development of progeny. It has
been reported that over nourishing singleton-bearing
adolescent ewes results in rapid maternal growth
and accumulation of maternal adipose tissue
(Wallace et al., 2006). As a result, placental growth
was restricted 30 to 40%, and ewes delivered lambs
prematurely that were 20 to 30% lighter at birth
compared with moderately nourished adolescents
of similar age. Furthermore, over-nourished ewes
produced less colostrum and progeny had lower
growth from birth to weaning (Da Silva et al., 2001).
The first trimester of fetal life coincides with
the peak in the number of follicles and oocytes in
fetal ovaries (Erickson, 1966), although ovarian
folliculogenesis in the bovine fetus is not completed
until late gestation (Rhind et al., 2001). Borwick et
al. (1997) reported retarded ovarian development

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in fetuses from ewes fed 50% of energy requirements
from breeding through mid-gestation when
compared with dams fed 150% of energy
requirements during that same period. Similarly,
Grazul-Bilska et al. (2009) identified that fetuses
collected at 135 days of gestation from ewes that
were nutritionally restricted from 50 to 135 days of
gestation had reduced proliferation of ovarian
primordial follicles when compared to fetuses of
adequately nourished ewes. This compliments data
in beef cattle which have illustrated a reduced
number of antral follicles present in heifers born to
nutritionally restricted dams during the first trimester
of gestation (Mossa et al., 2009; Echternkamp et
al., 2012). While reproductive development and
efficiency may be influenced in the offspring of
nutritionally altered dams, we know that epigenetic
modifications to the genome are heritable, and
therefore, alterations in reproductive capacity may
not be relegated to the first generation. Blair et al.
(2010) reported a trans-generational effect of
maternal nutrition during early pregnancy on
reproductive capacity of descendants. Fewer
grand-offspring resulting from over-nourished,
lightweight ewes reached puberty prior to their first
breeding season when compared with limit-fed,
lightweight grand-dams or heavyweight grand-dams
regardless of nutrition.
Studies investigating protein supple-
mentation of cows during early gestation have
reported alterations in the placental environment that
can impact embryonic development. Research in
dairy cattle has reported that high protein diets
increased concentrations of blood urea nitrogen
(BUN) and milk urea nitrogen (MUN), resulting in
decreased pregnancy rates, conception rates, and
embryonic development (Blanchard et al., 1990;
Butler et al., 1996; Rajala-Schultz et al., 2001).
Specifically, a BUN above 19 mg/dL decreased
preimplantation embryo development and survival
in sheep (Butler et al., 1996); whereas, BUN
concentrations below 10 mg/dL did not (Rajala-
Schultz et al., 2001). Sullivan et al. (2009) identified
in Bos indicus-cross beef cows that high crude
protein (CP) concentrations during mid-gestation
reduced primordial and primary follicle density in
female progeny at 23 months of age. Furthermore,
Sullivan et al. (2010) reported that excess maternal
dietary CP during the first trimester of gestation
resulted in decreased follicle stimulating hormone
(FSH) concentrations, seminiferous tubule diameter,
and testicular volume in male offspring at 5 months
of age. This decrease in fertility may be mediated
through changes in the uterine environment.
Research by Elrod et al. (1993) and Elrod and
Butler (1993) found that excess protein decreased
uterine pH on day 7 of the estrous cycle compared
to those fed a balanced diet. Changes in uterine pH
are important due to the effect pH has on embryo
development and survival. Ocon and Hansen (2003)
reported that in vitro culture of bovine embryos
below a pH of 7.0 resulted in decreased cleavage
rates and development to blastocyst stage.
Although excess protein may negatively
impact early embryonic development, moderate
increases in protein during the later stages of early
gestation may be beneficial. The amino acid arginine
is the substrate for nitric oxide (NO) synthesis (Flynn
et al., 2002), which is a known vasodilator, and
when used as a therapeutic agent, has increased
birth litter weight in pigs (Mateo et al., 2007) and
birth weight in humans (Vosatka et al., 1998).
Therefore, it is conceivable that increased dietary
protein could result in greater amounts of arginine
in the diet, resulting in increased uteroplacental
blood flow and subsequent birth weights. In fact,
Sullivan et al. (2009) reported that increased dietary
protein during the first 2 trimesters of gestation
increased the number of placental cotyledons in first-
parity Bos indicus heifers, and Vonnahme et al.
(2011) reported increased uterine blood flow in
protein supplemented ewes when compared with a
control.

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Late gestation
Energy demand by the gravid uterus is
greatest during the last trimester (Ferrell et al., 1976),
and supply of glucose and amino acids to the gravid
uterus are important substrates for fetal growth.
Providing an adequate supply of nutrients to the fetus
has the potential to increase birth weights and has
significant implications for subsequent growth, as
birth weights are positively correlated with calf
growth rate to weaning and live weight at weaning
when calves are reared on their dams (reviewed in
Greenwood et al., 2010). Energy deficiency (70%
of recommendations) in primiparous beef cows
during the last 100 days of pregnancy in beef cattle
was reported to decrease calf birth weight, weaning
weight, and delayed age at puberty of the progeny
compared to heifers from dams adequately fed
during the last 100 days (Corah et al., 1975).
Furthermore, Corah et al. (1975) observed that
progeny born to energy-restricted cows had
increased rates of morbidity and mortality. In sheep,
energy restriction during the last trimester has been
generally shown to decrease birth weights, but a
reduction in postnatal growth may or may not persist
through weaning and 1 year of age, depending on
adequacy of postnatal diets (reviewed in
Greenwood et al., 2010). When ewes are
undernourished during lactation, growth restriction
of progeny persists, but when ewes are adequately
nourished during lactation, growth restriction of
progeny does not persist. Thus, it appears that
direct prenatal effects on capacity for growth of the
progeny are somewhat ameliorated or exacerbated
depending upon the postnatal environment into which
they are born (Greenwood et al., 2010).
As lactation and gestation progresses in the
dairy cow, she moves from a negative to a positive
energy balance. Excess energy during this period
has the potential to increase glucose supply to the
fetus and lead to increased risk of glucose
intolerance and obesity in the offspring, as seen in
other species (Buchanan and Kjos, 1999).
Increased maternal body weight (BW) and plasma
triglycerides have been linked to an increased fatty
acid transfer to the fetus in humans, thereby inducing
increased birth weight and adiposity (Heerwagen
et al., 2010). In sheep, decreased glucose
effectiveness and insulin sensitivity, increased
appetite and gain, increased carcass adiposity, and
increased weight at 19 months of age were
observed in progeny from gestating ewes fed 150%
of energy requirements to an obese body mass index
(Long et al., 2010).
Source of energy in late-gestation beef cow
diets also appears to impact fetal growth and
development. Loerch (1996) reported that limit
feeding a high grain diet to beef cows during late
gestation resulted in heavier birth weight calves
compared to cows fed an isocaloric hay based diet;
however, inconsistent results were reported for
weaning weight (Loerch, 1996). In contrast, Susin
et al. (1995) reported similar birth weights and
postnatal growth of lambs from ewes limit-fed a
corn-based gestation diet similar to the studies
conducted in cows (Loerch, 1996).
Puberty and lifetime reproductive efficiency
may also be altered by maternal energy because
ovarian folliculogenesis in the bovine fetus is not
completed until late gestation (Rhind et al., 2001).
Da Silva et al. (2002; 2003) observed a decrease
in antral follicle counts of ewe lambs resulting from
dams fed a high plane of nutrition during late
gestation. Long et al. (2010) reported that ewes
born to dams that were nutrient restricted during
mid-gestation had reduced circulating
concentrations of progesterone during the luteal
phase of their estrous cycles, and fewer of those
offspring gave birth to a lamb during their first
parturition. Cushman et al. (2012) demonstrated
that heifers born to mature cows (>3 years of age)
fed 125% of nutritional maintenance requirements
during the third trimester of gestation conceived
earlier in their first breeding season than those born
to dams fed either 75 or 100% of nutritional
maintenance requirements during that period.

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Studies investigating protein
supplementation of cows during late gestation have
reported long-term impacts on their progeny. In a
recent review, Vonnahme and Lemley (2012)
reported that providing protein supplementation to
cows beginning on day 190 of gestation resulted in
a doubling of uterine blood flow when compared
with non-supplemented cows. It is hypothesized that
increased uterine blood flow may increase progeny
performance, owing to increased nutrient transfer
to the fetus. Gestating ewes fed a diet high in urea
had progeny who went on to exhibit increased rates
of metabolism and growth compared to progeny
from those ewes not supplemented with urea
(McEvoy et al., 1997). Singleton fetuses from ewes
consuming a high-protein diet during late gestation
were heavier on day 130 of gestation compared
with fetuses from ewes consuming the low-protein
diet, with no differences in placental weight
(Camacho et al., 2010). Heifer progeny from the
dams supplemented with protein had heavier BW
at weaning and breeding compared with progeny
from dams not supplemented with protein (Martin
et al., 2007).
With regard to maternal protein and
reproductive development of the progeny, maternal
protein supplementation during late gestation has
been shown to increase the probability of female
progeny reaching puberty prior to the first year of
breeding (Funston et al., 2010). Specifically, more
heifers born to cows that grazed winter range and
were supplemented with a 28% CP supplement
during late gestation were pubertal at initiation of
the first year of breeding when compared with
progeny of cows not receiving a supplement. In
another study, protein supplementation to cows
grazing winter range during late gestation produced
heifer progeny that had greater pregnancy rates
during their first year of breeding and calved earlier
during their first year of calving (Martin et al., 2007).
Collectively, these results provide evidence that an
individual component of the maternal diet, such as
protein, can impact heifer progeny reproductive
development as well as steer progeny growth and
body composition.
Distillers dried grains with solubles
(DDGS), a byproduct of ethanol fermentation,
contains elevated concentrations of fat and protein
and is an alternative energy source used by beef
cow producers. Although feeding fat as an energy
source to beef cows in the form of sunflower or
soybean oil (1.87 lb/day) did not impact progeny
birth weights or performance (Alexander et al.,
2002). Research that we have conducted at Purdue
University (Gunn et al., 2011a), as well as research
conducted by others (Radunz et al., 2010; Radunz
et al., 2012; Wilson et al., 2012) have demonstrated
that when DDGS is fed to gestating (third trimester)
and lactating beef cows to meet energy requirements,
increased progeny birth weights result compared
to beef cows fed hay-based or corn-based diets
with similar energy concentrations. However,
problems with dystocia have been observed (Gunn
et al., 2011a). The effect of maternal DDGS on
postnatal progeny weight gain has been variable. In
male progeny fed in the feedlot, weaning and yearling
weights did not differ (Radunz et al., 2010; Gunn et
al., 2012a; Radunz et al., 2012 ), however, female
progeny from DDGS treated dams tended to be
heavier than progeny from corn silage/hay treated
dams throughout the developmental post-weaning
period (Gunn et al., 2012b). In addition, DDGS
progeny had a greater frame score than control
progeny throughout the developmental period.
Progeny from DDGS fed cows have not differed in
blood glucose or insulin concentrations, glucose
clearance, morbidity, or immunoglobulin
concentrations (Radunz et al., 2010; Gunn et al.,
2012a; Radunz et al. 2012).
The effect of DDGS on reproduction is
similar to the effect that protein has had on
reproduction. Gunn et al. (2011b) observed that
ovarian size, antral follicle count, follicular growth
parameters, and age at puberty in progeny did not
differ due to maternal supplementation of DDGS.

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However, BW at puberty and pregnancy rates to
artificial insemination (AI) were greater in DDGS
progeny compared to non-DDGS fed progeny
(70.6 vs. 33.3%). Overall breeding season
pregnancy rate did not differ. Rate of dystocia in
female progeny at first parturition and grand-
offspring birth BW did not differ due to treatment.
It is unclear if source of energy or overfeeding of
protein caused these effects.
Neonatal Programming
The first weeks of life for a neonate seem
to also have long-lasting consequences on the
physiological function of the animal. In fact, many
of the effects of fetal programming can be
ameliorated or exacerbated due to nutrition of the
neonate. In beef cows, Freetly et al. (2000)
demonstrated that effects of variable nutrition during
mid and/or late pregnancy on weight at birth are
overcome when adequate, high quality postnatal
nutrition is available and resulted in no differences
in calf BW at 58 days of age. Right after birth, the
first nutrients consumed by calves are from
colostrum. In addition to nutrients, colostrum is also
rich in immunoglobulins (Ig), growth factors, and
hormones. DeNise et al. (1989) demonstrated a
positive and significant relationship between plasma
IgG concentrations and future milk production of
calves that were allowed to suckle their dams for
the first 24 hr of life. Other factors in colostrum
could also influence subsequent production. Hough
et al. (1990) observed that although maternal under
nutrition did not affect colostral IgG concentrations,
calves born to nonrestricted cows that received
colostrum from restricted dams tended to have
lower serum IgG concentrations at 24 hr of life than
those receiving colostrum from well-nourished
cows.
In modern dairy calf rearing systems, the
restricted feeding of milk replacer and the early
weaning of calves are practiced to lower costs.
There is increasing evidence, however, that intensified
early nutrition may not only improve the health, well-
being, and growth of the calf in the short term, but
may also have a long-term positive impact on
performance (Jasper and Weary, 2002). Findings
of Tudor and ORourke (1980) with calves reared
on milk replacer, coupled with studies in which
calves are reared on their dams (Café et al., 2006),
show that the postnatal maternal contribution to
preweaning growth of calves is substantial. A
positive relationship between milk intake and dry
feed intake exists when considering the entire
preweaning period, indicating that the influence is
through increasing the capacity of the calf to consume
feed (Christian et al., 1965). In fact, some have
reported that increased gain in dairy calves during
the first 2 mo of life results in significantly greater
BW at 24 mo of age (Robelin and Chilliard, 1989;
Moallem et al., 2010), reduced breeding age
(Raeth-Knight et al., 2009), and higher milk yield
when the calf matures (Moallem et al., 2010). One
possible explanation for increased milk yields could
be the influence of early feeding levels on the
development of the mammary gland. A high feeding
level during the first 8 weeks of life was
accompanied by an intensified development of the
mammary parenchymal mass, DNA, and RNA of
Holstein heifer calves compared to moderately fed
controls (Brown et al., 2005). Interestingly, the first
weeks of life were found to be most influential
because a higher feeding intensity between weeks
8 and 14 of life did not affect the development of
the mammary gland (Brown et al., 2005).
Composition of milk has also been suggested as an
important source of variation in calf performance.
Christian et al. (1965) observed that maternal milk
composition accounted for more of the variability
in weaning weight and pre-weaning gain than total
milk intake during this period. With respect to body
composition, increasing the percentage of milk
replacer protein has been demonstrated to increase
body protein and decrease adiposity compared with
lower levels of protein (Donnelly and Hutton, 1976;
Diaz et al., 2001). Additionally, decreasing milk
replacer fat content decreases body fat percentage
with no effect on protein (Tikofsky et al., 2001).

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Research conducted at Purdue University
(Shee et al., 2012) has demonstrated that feeding
DDGS to cows from calving to mid lactation (129
DIM) increased male progeny gain and weight at
weaning (219 DIM) compared to male progeny
from cows fed a corn silage based diet. The diets
were isocaloric but differed in fat and protein
contents. As a result, milk composition of the cows
was altered and may have been responsible for
changes in growth. Feeding DDGS to cows from
calving to mid lactation did not affect milk production
but decreased milk fat and protein, increased the
percentage of long-chain, monounsaturated and
polyunsaturated fatty acids (FA) in milk, including
conjugated linoleic acid (CLA), and decreased the
percentage of medium-chain and saturated FA.
Specific FA and level of unsaturation of the FA in
maternal milk can impact growth of progeny. In rats,
maternal consumption of a highly unsaturated or
highly saturated FA diet results in lower birth weights
and decreased postnatal weight gain (Siemelink et
al., 2002). Shee et al. (2012) observed that the
weight advantage for male progeny whose dams
were fed DDGS was not maintained through
slaughter; however, intramuscular fat content was
decreased in the male progeny of DDGS-fed cows,
indicating that maternal DDGS could impact
progeny body composition.
Conclusion
Unfortunately, there is a lack of studies in
dairy cattle that demonstrate direct effects of
maternal nutrition on postnatal performance of
offspring; however, numerous studies in beef cattle
and sheep demonstrate that both energy and protein
(inadequate or excess) can effect long-term
productivity of progeny. Thus, management of the
cow herd not only has consequences for cow
productivity but also for productivity of the next
generation of cows. Fortunately, management of the
neonatal calf can ameliorate many of the negative
consequences of poor fetal nutrition. Data are
lacking on the effect of specific nutrients (FA, amino
acids, vitamins, and minerals) on long-term
performance of offspring and further research is
needed to determine these effects and understand
the underlying mechanisms.
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Figure 1. Sheep conceptus growth and development in gestation and periods of specific organ development
that can be influenced by maternal nutrition (CV = cardiovascular system; Symonds et al., 2010).