animal
Animal (2007), 1:9, pp 1283–1296 & The Animal Consortium 2007
doi: 10.1017/S175173110700050X
Prenatal and pre-weaning growth and nutrition of cattle: longterm consequences for beef production
P. L. Greenwood- and L. M. Cafe
NSW Department of Primary Industries, Beef Industry Centre of Excellence and Cooperative Research Centre for Beef Genetic Technologies, University of New
England, Australia
(Received 27 February 2007; Accepted 18 May 2007)
Severe, chronic growth retardation of cattle early in life reduces growth potential, resulting in smaller animals at any given age.
Capacity for long-term compensatory growth diminishes as the age of onset of nutritional restriction resulting in prolonged
growth retardation declines. Hence, more extreme intrauterine growth retardation can result in slower growth throughout
postnatal life. However, within the limits of beef production systems, neither severely restricted growth in utero nor from birth
to weaning influences efficiency of nutrient utilisation later in life. Retail yield from cattle severely restricted in growth during
pregnancy or from birth to weaning is reduced compared with cattle well grown early in life, when compared at the same age
later in life. However, retail yield and carcass composition of low- and high-birth-weight calves are similar at the same carcass
weight. At equivalent carcass weights, cattle grown slowly from birth to weaning have carcasses of similar or leaner
composition than those grown rapidly. However, if high energy, concentrate feed is provided following severe growth restriction
from birth to weaning, then at equivalent weights post-weaning the slowly-grown, small weaners may be fatter than their
well-grown counterparts. Restricted prenatal and pre-weaning nutrition and growth do not adversely affect measures of beef
quality. Similarly, bovine myofibre characteristics are little affected in the long term by growth in utero or from birth to
weaning. Interactions were not evident between prenatal and pre-weaning growth for subsequent growth, efficiency, carcass,
yield and beef-quality characteristics, within our pasture-based production systems. Furthermore, interactions between genotype
and nutrition early in life, studied using offspring of Piedmontese and Wagyu sired cattle, were not evident for any growth,
efficiency, carcass, yield and beef-quality parameters. We propose that within pasture-based production systems for beef cattle,
the plasticity of the carcass tissues, particularly of muscle, allows animals that are growth-retarded early in life to attain normal
composition at equivalent weights in the long term, albeit at older ages. However, the quality of nutrition during recovery
from early life growth retardation may be important in determining the subsequent composition of young, light-weight cattle
relative to their heavier counterparts. Finally, it should be emphasised that long-term consequences of more specific and/or
acute environmental influences during specific stages of embryonic, foetal and neonatal calf development remain to be
determined. This need for further research extends to consequences of nutrition and growth early in life for reproductive
capacity.
Keywords: birth weight, foetal programming, meat quality, muscle fibres, neonates
Introduction
There are numerous growth path possibilities during early
and later life that may influence productive characteristics
of cattle. These different growth paths result from factors
including climate, soil quality and pasture species, which
contribute to variable pasture and nutrient quality and
availability. Growth of the bovine foetus has well-studied
consequences for survival (Holland and Odde, 1992) and
-
E-mail: paul.greenwood@dpi.nsw.gov.au
can be slowed during the latter half of gestation by
restricted nutrition and/or inadequate placental development (Bell et al., 2005). Similarly, influences of pre-weaning
nutrition, most notably lactational performance of the dam,
on growth to market weights of cattle are well characterised (Berge, 1991). However, consequences of foetal
calf growth for subsequent growth, and of foetal and
neonatal calf growth for efficiency and carcass- and beefquality characteristics, are less well understood.
Hence, this paper reviews research on consequences of
cattle nutrition and growth during foetal and neonatal life
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Greenwood and Cafe
for subsequent growth and efficiency, and for carcass and
beef quality. It includes findings from our recent studies on
consequences of maternal nutrition (commencing between
days 30 and 90 of gestation) and growth during pregnancy
and to weaning of cattle sired by bulls of extreme genotypes for muscle and intramuscular fat development (Cafe
et al., 2006a and 2006b; Greenwood et al., 2006). Factors
affecting growth and nutrition of the bovine foetus and
milk-fed calf are also briefly described. The reader is also
referred to reviews on consequences of prenatal development in livestock species by Bell (2006) and Symonds et al.
(2007), and on consequences of bovine foetal, pre-weaning
and early post-weaning growth and nutrition by Berge
(1991) and Greenwood et al. (2005).
Normal bovine conceptus growth and metabolism
During postnatal growth, energy and nutrient availability
directly influence growth and body composition of cattle.
However, environmental influences on foetal growth and
development and, hence, birth characteristics are regulated
via the dam, and by the placenta that functions as a
nutritional conduit between the dam and the foetus.
Most growth of the bovine foetus occurs during the final
100 days or so of a gestation averaging approximately 280
days (Winters et al., 1942; Lyne, 1960; Ferrell et al., 1976;
Prior and Laster, 1979). Foetal nutrient uptake becomes a
quantitatively important contributor to maternal nutrient
requirements only after mid-gestation (Ferrell et al., 1983).
Unlike the sheep, in which the placenta attains most of its
mass of dry tissue, protein and DNA by mid-gestation
(Ehrhardt and Bell, 1995), the bovine placenta normally
continues to increase in weight until near term (Prior and
Laster, 1979; Ferrell, 1989). As a result, it has been suggested that placental growth may be less sensitive to
nutritional deficiencies in cattle than in sheep. Placental
weight and birth weight are highly correlated in cattle
(Anthony et al., 1986b; Echternkamp, 1993; Zhang et al.,
1999); however, the functional capacity of the placenta is
closely related to placental perfusion. Bovine uterine and
umbilical blood flow increases exponentially during the
second half of gestation, which equates to relatively constant rates of umbilical blood flow on a foetal weightspecific basis during this period (Reynolds et al., 1986). A
more detailed account of placental function and metabolism
in cattle is provided by Ferrell (1989), and of foetal macronutrient requirements and metabolism in cattle and
sheep, and of placental function and metabolism, by Bell
et al. (2005).
Intrauterine growth retardation
Maternal nutrition
Severe nutritional restriction for at least the last half to onethird of pregnancy is usually required to reduce bovine
foetal growth. Significant reductions in birth weight were
1284
caused by prolonged underfeeding of heifers from weaning
until parturition (Wiltbank et al., 1965), and underfeeding
of heifers and cows for the second and third trimesters
(Ryley and Gartner, 1962; Hodge and Rowan, 1970; Freetly
et al., 2000; Cafe et al., 2006b) or late pregnancy only
(Hight, 1966; Tudor, 1972; Bellows and Short, 1978; Kroker
and Cummins, 1979). The effect of nutritional restriction on
birth weight was more pronounced in calves from heifers
than those from cows when the period of restriction
encompassed mid- and late gestation (Hennessy et al.,
2002) rather than late gestation only (Tudor, 1972). However, birth weight was not significantly affected by nutritional restriction of heifers from mating to 140 days
gestation (Cooper et al., 1998) or during the final 12 weeks
of pregnancy (Hodge et al., 1976), or of mature cows for
the second trimester (Freetly et al., 2000).
During the final one-half to one-third of pregnancy, feed
energy available to the dam appears to have more influence
on birth weight than the availability of protein, although
results are variable (Holland and Odde, 1992). Variation in
feed energy available to the dam during this period resulted
in differences in birth weight, ranging from 0 to 8.2 kg
(Dunn et al., 1969; Tudor, 1972; Laster, 1974; Corah et al.,
1975; Bellows and Short, 1978; Kroker and Cummins, 1979;
Bellows et al., 1982). Similarly, variable protein supply of
the diet during the third trimester may (Bellows et al., 1978)
or may not (Anthony et al., 1986a; Holland and Odde, 1992)
alter birth weight of calves, while restricted or supplemental
dietary protein during early or mid-pregnancy had little effect
on birth weights (Perry et al., 1999 and 2002). Furthermore,
supplementation of grazing cows for 3 months pre-partum
with 0.45 kg/day of 42% crude protein supplement did
not affect calf birth weights (Stalker et al., 2006). However,
more chronic nutritional restriction of energy and/or protein
of heifers from weaning until parturition resulted in birth
weight differences of up to 10 kg due to energy supply
and up to 7.3 kg due to protein supply (Wiltbank et al.,
1965).
As described above, placental weight and birth weight
are highly correlated in cattle. However, because the bovine
placenta may continue to increase in mass until near term,
it is less clear whether the placenta regulates bovine foetal
growth to the same extent as it does in sheep (Ferrell,
1989). Placental characteristics may be altered by nutrition
during early and mid-pregnancy without significantly
affecting foetal size (Rasby et al., 1990), and protein supplementation of cows during early or mid-pregnancy may
also alter placental characteristics without necessarily
affecting birth weight (Perry et al., 1999 and 2002).
Development and growth of vital organs precede development of bone, muscle and fat (Palsson, 1955), respectively; hence the mass of the relatively late maturing
carcass tissues are generally considered more susceptible to
the effects of nutrition during later pregnancy when nutrition impacts most on foetal growth. However, more subtle
effects on organ and tissue development due to nutrition
during early pregnancy may occur, with the potential for
Consequences of growth early in life of cattle
long-term consequences for health, as shown in sheep
(Greenwood and Bell, 2003; Bell et al., 2005; Symonds
et al., 2007).
Thermal environment
Foetal growth in cattle was restricted (18% lower foetal
weight) by chronic heat stress of pregnant cows, while
provision of shade resulted in a 3.1 kg increase in birth
weight (Collier et al., 1982). In sheep, chronic heat stress in
early to mid-gestation restricts placental development, thus
imposing a limitation on subsequent foetal growth irrespective of nutrition later in pregnancy (Bell et al., 1987).
This suggests that restricted foetal calf growth due to heat
stress is probably a consequence of reduced placental
development. Severe cold stress of cattle may also reduce
foetal growth if inadequate nutrition is provided to meet
the additional metabolic requirements of cows in addition
to foetal requirements for growth and development
(Andreoli et al., 1988), although in sheep, more moderate
cold stress of ewes in late gestation increased birth weight
by 15% (Thompson et al., 1982). It is believed that temperature regulates blood flow to the periphery and lungs
in order to preserve or dissipate body heat, resulting in
increased or decreased blood flow and nutrient supply to
the gravid uterus (Reynolds et al., 1985).
Parity
Heifers give birth to smaller calves, on average, than cows
(reviewed by Holland and Odde, 1992) due at least in part
to size and nutritional requirements for growth of heifers,
limiting nutrient availability for placental and foetal growth.
Severe maternal nutritional restriction may impact more on
birth weights of calves of heifers than of cows, particularly
among male calves and those of sires with inherently high
birth weight of offspring (Hennessy et al., 2002), presumably due to their greater requirements for nutrients
compared with female calves and those of sires with
inherently low birth weight of offspring. In adolescent
sheep fed to attain excessive fatness prior to and during
gestation, placental and foetal growth and birth weight are
reduced (Wallace et al., 1996 and 1999), although the
extent to which over-nutrition of adolescent heifers
influences birth weight is not clear.
Litter size
Twin calves and higher multiples are rare in cattle unless
exogenous regulation of ovarian function or embryo
transfer is practised. Individuals within litters have reduced
foetal growth compared with singletons due to a reduced
number of placentomes and mass of placenta per foetus
(Hafez and Rajakoski, 1964; Greenwood et al., 2000b) and
because of greater total nutrient requirements of the litter.
On average, twin calves range from 7.4 to 9.8 kg lighter
than singletons (Gregory et al., 1990 and 1996; De Rose
and Wilton, 1991; Cummins, 1994; Wilkins et al., 1994).
Restricted nutrition limits foetal growth earlier and more
severely in twins or higher multiples than in singletons,
although stocking rates of pregnant cows fed on pasture
did not significantly influence the birth weight of twins
(Wilkins et al., 1994).
Foetal and maternal genotype
Foetal genotype is most important in determining foetal
growth during early and mid-pregnancy, whereas maternal
genotype is more important in determining foetal growth
during late pregnancy when most foetal growth normally
occurs and foetal growth is increasingly subject to external
influences mediated via the dam. The effect of foetal and
maternal genotype on foetal growth has been most convincingly demonstrated in cattle by Ferrell (1991) who
implanted Charolais (heavier birth weight) or Brahman
(lighter birth weight) embryos into Charolais and Brahman
cows. At 232 days of pregnancy, each foetal genotype was
similar in size, irrespective of dam breed. However, by 274
days of gestation Charolais foetuses in Brahman cows were
7 kg lighter than those in Charolais cows. In contrast,
Brahman foetuses in Charolais cows were only 2 kg heavier
than those in Brahman cows. Similar results were obtained
by Joubert and Hammond (1958) for birth weights for South
Devon and Dexter cattle and their reciprocal crosses. In this
regard, foetal growth capacity as influenced by sex and siregenotype may also influence the nutritional status of the
pregnant cow during late gestation (Greenwood et al.,
2002b), probably due to differences in foetal nutrient
uptake, contributing to maternal nutrient requirements.
Growth and development from birth to weaning
Calves undergo a transition at birth from a diet comprising
primarily glucose and amino acids to one that is quantitatively greater and is proportionately higher in fat. This is
associated with maturation of the digestive, metabolic and
endocrine systems. Evidence in sheep suggests severely
growth-retarded newborns are immature with respect to
energy metabolism and have more foetal-like metabolism
than their well-grown counterparts (Greenwood et al.,
2002a; Rhoads et al., 2000a and 2000b).
The major nutritional factors affecting pre-weaning calf
growth and composition at weaning are the lactational
performance of the dam and the quality and availability of
nutrients from pasture and/or supplementation prior to and
following parturition. Most notably, maternal genotype, age
and parity, nutrient availability and body condition and live
weight of the dam, and capacity of the calf to grow and
consume milk, interact to influence lactational output.
Calves become increasingly dependent on forage-based
diets that result in the production of volatile fatty acids that
stimulate development and maturation of the rumen
(Warner and Flatt, 1965) until weaning, when this dependence becomes complete.
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Greenwood and Cafe
Long-term consequences of altered growth during the
early life of cattle
In this section, the long-term consequences of altered
growth early in life for growth, feed efficiency, and carcass,
yield, beef quality and myofibre characteristics are discussed first for altered foetal growth, and then for altered
pre-weaning growth. Interactions between prenatal and
pre-weaning growth are then discussed.
Consequences of foetal growth and nutrition
Postnatal growth. In our recent studies of consequences of
foetal growth, which compared performance of calves differing by 10.2 kg or by 35% in birth weight (Table 1),
capacity of low-birth-weight cattle to exhibit compensatory
growth was limited. Cattle significantly growth retarded
during foetal life due to severely restricted maternal nutrition from early pregnancy (commencing between day 30
and 90 of pregnancy) to parturition remained smaller during
rearing on their dams and at any given postnatal age
after weaning compared with their well-grown or betternourished counterparts (Table 1). However, it remains
speculative whether this represents a permanent stunting
or simply a delay of attainment of mature size of cattle.
Growth of low-birth-weight cattle was significantly slower
than those of high birth weight at all stages of postnatal
growth, although about half the difference in average daily
gain (ADG) during feedlotting was explained by differences
in weight at feedlot entry between the low- and high-birthweight cattle (Table 1). However, when differences in birth
weights were less-pronounced, post weaning and feedlot
growth were not significantly affected by birth weight (Cafe
et al., 2006a). These findings are consistent with those of
Swali and Wathes (2006) who found that small size at birth
resulted in smaller cattle compared with high-birth-weight
cattle at 15 months of age, while average-birth-weight
cattle did not differ significantly in weight during postnatal
growth compared with the low- or high-birth-weight
groups.
In contrast to the above findings, artificially reared lowbirth-weight male calves grew more rapidly to weaning
than their high-birth-weight counterparts, although the
opposite occurred for female calves (Tudor and O’Rourke,
1980). Hence, it is important to recognise that an assessment of influences of foetal development on postnatal
performance requires consideration of the consequences of
nutrition during pregnancy on subsequent maternal performance when offspring remain on their dams to weaning,
due to carry-over effects on the dam. In this regard, readers
are referred to Greenwood et al. (1998) for an example of a
rearing system designed to uncouple prenatal and postnatal
influences in ruminants varying in birth weight. The net
effects of maternal nutrition during pregnancy on the calf
remain of practical significance to livestock producers, and
influences of nutrition during mid- and late pregnancy or
late pregnancy only on calf weaning weight have been
consistently shown (e.g. Hight, 1966 and 1968a; Cafe et al.,
2006b; Stalker et al., 2006), irrespective of effects on foetal
growth.
In relation to potential interactions between prenatal
and postnatal nutrition and growth, differences in weight of
calves at birth following three levels of maternal nutrition
during late pregnancy disappeared by weaning when
postnatal nutrition was of high quality and availability
Table 1 Consequences of growth in utero for growth and live-weight characteristics of beef cattle to 30 months of age (adapted from Greenwood
et al., 2005 and 2006)Prenatal growth/birth weight
Birth weight (kg)
Pre-weaning ADG (g)
Weaning (7 months) weight (kg)
Backgrounding ADG (g)
At equivalent age (26 to 30 months)
Feedlot entry (26 months) weight (kg)
Feedlot ADG (g)
Feedlot exit (30 months) weight (kg)
At equivalent feedlot entry live weight (500 kg)
Age at feedlot entry (day)Feedlot ADG (g)
Feedlot exit weight (kg)
Age at feedlot exit (day)y
Low (n 5 120)
High (n 5 120)
Significance of difference (P)
28.6
670
174
571
38.8
759
198
603
,0.001
,0.001
,0.001
481
1480
647
520
1617
703
,0.001
,0.001
,0.001
797
1515
671
914
715
1583
679
833
,0.001
0.019
0.017
,0.001
-
Abbreviation is: ADG 5 average daily gain.
Values are predicted means from residual maximum likelihood (REML) analyses including effects of birth weight, pre-weaning nutrition, sex/year cohort,
sire-genotype and their interactions.
Maternal nutritional treatments commenced between days 30 and 90 of pregnancy (refer to Cafe et al. (2006b) and Greenwood et al. (2006) for details of
pasture-based nutritional treatments and selection criteria for calves used to study long-term consequences of growth early in life).
Predicted from mean ADG during backgrounding.
y
Predicted from mean ADG during background and mean feedlotting period.
-
1286
Consequences of growth early in life of cattle
Table 2 Consequences of growth in utero for feed intake and efficiency of beef cattle during feedlotting from 26 to 30 months of age (L. M. Cafe
and P. L. Greenwood, unpublished results)
Prenatal growth/birth weight
Birth weight (kg)
At equivalent age (26 to 30 months)
Feedlot entry (26 months) weight (kg)
Feedlot ADG (g)Feed intake (kg/day)Feed efficiency (kg DM intake/kg LW gain)
Residual feed intake (kg)At equivalent feedlot entry LW (490 kg)
Feedlot ADG (g)Feed intake (kg/day)Feed efficiency (kg DM intake/kg LW gain)
Low (n 5 77)
High (n 5 77)
Significance of difference (P)
28.1
38.4
466
1279
13.21
10.00
20.005
513
1396
14.63
10.38
0.003
,0.001
0.004
,0.001
0.26
0.99
1317
13.86
10.25
1361
14.01
10.15
0.46
0.55
0.89
-
Abbreviations are: ADG 5 average daily gain, DM5dry matter; LW5live weight.
Values are predicted means from residual maximum likelihood (REML) analyses including effects of birth weight, pre-weaning nutrition, sex/year cohort,
sire-genotype and their interactions, with feedlot entry weight as a covariate (linear and, where significant, quadratic) to predict means at equivalent feedlot
entry weight. Difference in significance of feedlot ADG between Tables 1 and 2 is due to the number of cohorts studied (three cohorts in Table 1 v. two cohorts
in Table 2) and the duration of the measurement period (average of 117 days in Table 1 v. 70 days in Table 2).
During 70-day period in feed intake pens.
At mean metabolic live weight.
-
(Hight, 1968b). In this study, however, residual effects of
the previous year’s nutrition influenced calf growth, with
cows previously well nourished producing heavier calves,
and vice versa (Hight, 1968b). Similarly, effects of variable
nutrition during mid- and/or late pregnancy on weight at
birth were overcome by adequate nutrition post partum,
resulting in no difference in body weight at 58 days of age
(Freetly et al., 2000). While twin cattle are lighter at birth
and grow more slowly on their dams to weaning (Hennessy
and Wilkins, 1997), they may grow more slowly (Gregory
et al., 1996), at a similar rate (De Rose and Wilton, 1991) or
more rapidly (Wilkins et al., 1994; Clarke et al., 1994;
Hennessy and Wilkins, 1997) post-weaning than singletons,
depending on the rearing system and subsequent nutritional regimen.
Feed intake and efficiency. Slower feedlot growth by lowbirth-weight calves was associated with the consumption of
fewer nutrients in the feedlot but no difference in feed
efficiency or residual feed intake compared with high-birthweight calves at an equivalent age from 26 to 30 months
(Table 2). When compared at equivalent feedlot entry live
weights, differences in feed intake due to birth weight were
no longer apparent, consistent with findings in twin cattle,
which tended to consume less feed in feedlot than singletons, due primarily to their lower live weight (De Rose and
Wilton, 1991). Similarly, provision of supplement to cows
for 3 months pre-partum had no significant post-weaning
effects on ADG, feed intake and feed efficiency in steers
(Stalker et al., 2006) or heifers (Martin et al., 2007) that
were individually fed following weaning, although the
heifers of supplemented cows tended to have greater
absolute and residual feed intakes during individual feeding
for 84 days post-weaning.
Body and carcass composition. Few studies have examined
long-term consequences of foetal nutrition and growth for
body and carcass characteristics in cattle (Tudor et al.,
1980) prior to our more recent studies (Greenwood et al.,
2006). Our research has shown that a significant reduction
in birth weight following severe maternal nutritional
restriction did not influence indices of fatness, apart from
P8 (rump) fat, in carcasses of Wagyu- or Piedmontese-sired
steers and heifers at 30 months of age, beyond that normally attributable to differences in live or carcass weight
(Table 3). Low-birth-weight cattle had a similar intramuscular fat content, retail yield, fat trim and bone content at
equivalent carcass weight, suggesting little overall difference in carcass composition from their high-birth-weight
counterparts. However, ossification score was higher in
low- compared with high-birth-weight calves (Table 3),
suggesting an impact of prenatal growth on calcification of
bone and relative maturity. Similar to our findings, gross
compositional differences were not evident in the whole
body or in the carcass of Hereford steers or heifers grown to
370 to 400 kg live weight following restricted or adequate
nutrition of their dams from 180 days of pregnancy to
parturition with a resultant 22% or 6.8 kg difference in calf
birth weight (Tudor et al., 1980). Furthermore, pre-partum
supplementation of cows had no effects on the carcass
composition of offspring following feedlotting for 222 days
post weaning (Stalker et al., 2006).
Research on twin cattle has also demonstrated that,
despite significantly lower birth weights and reduced preweaning growth, compositional differences at equivalent
slaughter weights or ages are small and not significant,
with twins generally having similar or leaner carcasses than
singletons (De Rose and Wilton, 1991; Wilkins et al., 1994;
Clarke et al., 1994; Gregory et al., 1996).
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Greenwood and Cafe
Table 3 Consequences of growth in utero for carcass and yield characteristics of beef cattle at 30 months of age (adapted from Greenwood
et al., 2006)
Prenatal growth/birth weight
Low (n 5 120)
High (n 5 120)
Significance of difference (P)
364
239
396
257
,0.001
,0.001
90.4
21.3
10.9
1.83
447
6.8
206
249
66.9
54.6
88.9
19.6
10.5
1.86
444
7.0
195
247
67.6
56.0
0.25
0.048
0.35
0.56
0.98
0.62
0.009
0.20
0.10
0.58
At equivalent age (30 months)
Carcass weight (kg)
Retail yield (kg)
At equivalent carcass weight (380 kg)
Eye muscle area (cm2)
P8 fat depth (mm)
Rib fat depth (mm)
Aus-Meat marble score
USDA marble score
Longissimus IMF (%)
Ossification score
Retail yield (kg)
Bone (kg)
Fat trim (kg)
Abbreviation is: IMF 5 intramuscular fat.
Values are predicted means from residual maximum likelihood (REML) analyses including effects of birth weight, pre-weaning nutrition, sex/year cohort, siregenotype and their interactions, with carcass weight as a covariate (linear and, where significant, quadratic) to predict means at equivalent carcass weight.
Refer to Table 1 for growth characteristics of the cattle.
Table 4 Consequences of growth in utero for objective measurements of m. longissimus (striploin) and m. semitendinosus (eye round) quality in
beef cattle at 30 months of age (adapted from Greenwood et al., 2006)
Prenatal growth/birth weight
Longissimus
Peak force (N)Compression (N)Cooking loss (%)
Ultimate pH
Colour L (lightness)
Colour a (red/green)
Colour b (yellow/blue)
Semitendinosus
Peak force (N)Compression (N)Cooking loss (%)
Low (n 5 120)
High (n 5 120)
Significance of difference (P)
39.2
13.9
21.6
5.47
39.5
26.3
13.6
40.5
14.4
21.7
5.48
40.0
26.7
13.8
0.26
0.19
0.57
0.50
0.21
0.20
0.15
46.2
22.6
21.5
46.4
22.7
21.3
0.81
0.97
0.52
Values are predicted means from residual maximum likelihood (REML) analyses including effects of birth weight, pre-weaning nutrition, sex, sire-genotype and
their interactions. Refer to Tables 1 and 3 for growth and carcass characteristics of the cattle.
Objective measures of texture, as defined by Perry et al. (2001).
Beef quality and myofibre characteristics. There were no
adverse effects on objective measurements of beef quality
including peak force, compression, cooking loss and colour
in the longissimus (striploin) and semitendinosus (eye
round) muscles at 30 months of age due to restricted
growth in utero (Table 4).
Myofibre characteristics including number and size of
myofibres, and percentages and relative areas of myofibres
in the m. longissimus lumborum (Table 5) and semitendinosus (results not shown) muscles at 30 months of
age, were also unaffected by calf growth in utero (Table 5).
1288
In this regard, nutrition during pregnancy resulting in
divergent foetal calf growth resulted in differences at birth
in the percentages of type 1 (low 17.2 v. high 23.3%) and
type 2A (28.2 v. 23.5%) myofibres, the ratio of fast to slow
(4.8 v. 3.4) myofibres and the cross-sectional area of type
2X (673 v. 831 mm2) myofibres (Greenwood et al., 2004).
However, differences in myofibre characteristics due to
foetal growth were no longer evident by weaning (P. L.
Greenwood, unpublished results). Within the present study,
as with newborn lambs restricted in growth during mid- to
late pregnancy (Greenwood et al., 1999 and 2000a),
Consequences of growth early in life of cattle
Table 5 Consequences of growth in utero for longissimus lumborum myofibre characteristics of heifer beef cattle at 30 months of age
(P. L. Greenwood and L. M. Café, unpublished results)
Prenatal growth/birth weight
Birth weight (kg)
Live weight at 30 months (kg)
Carcass weight (kg)
Muscle weight (g)
Muscle CSA (cm2)
Myofibres per mm2
Apparent myofibre number (31026)
Average myofibre CSA (mm2)
Total myofibre area (%)
Type 1
Type 2C
Type 2A
Type 2AX
Type 2X
Low (n 5 38)
High (n 5 40)
Significance of difference (P)
27.3
594
335
4097
85.3
130.5
1.112
5708
35.9
655
370
4453
90.4
126.8
1.137
5731
,0.001
,0.001
0.002
0.029
0.62
0.78
0.86
22.5
0.55
21.9
5.61
49.5
22.5
0.71
23.4
6.27
47.2
0.99
0.52
0.43
0.53
0.44
Abbreviation is: CSA 5 cross-sectional area.
Values are predicted means from residual maximum likelihood (REML) analyses including effects of birth weight, pre-weaning nutrition, sex, sire-genotype and
their interactions.
Refer to Lehnert et al. (2006) for myofibre classification and measurement methodology. Type 1, type 1 myosin heavy chain (MHC) slow oxidative; type 2C,
intermediate between type 1 and type 2A; type 2A, type 2A MHC fast oxidative-glycolytic; type 2AX, intermediate between type 2A and type 2X; type 2X,
type 2X MHC fast glycolytic.
apparent myofibre number was not affected by divergent
growth in utero (Table 5).
Consequences of pre-weaning growth and nutrition
Postnatal growth. Consequences of nutritional restriction
from birth to weaning for subsequent growth of cattle were
reviewed by Allden (1970), Berge (1991) and Hearnshaw
(1997). It is generally recognised that severe pre-weaning
nutritional restriction limits the capacity of cattle to exhibit
compensatory growth and achieve equivalent weight for
age in later life. In reviewing a series of Australian studies
on consequences of pre-weaning nutritional systems,
Hearnshaw (1997) concluded that compensatory gain
following pre-weaning growth restriction occurred most
frequently when overall post-weaning growth rates were
less than 0.6 kg/day, whereas at higher post-weaning
growth rates compensation was less evident. However, in
feedlot the differences in growth were in the opposite
direction to differences in growth post-weaning, and when
compensation did occur among cattle restricted prior to
weaning, the gains were only small. In more recent studies,
calves reared slowly (464 g/day) compared with those
reared rapidly (872 g/day) from birth to weaning were 37 kg
lighter at weaning, but 48 kg lighter following backgrounding due to a trend towards slower backgrounding
growth among the previously restricted cattle, and
remained 46 kg lighter at slaughter at 17 months of age
(Hennessy and Morris, 2003; Hennessy and Arthur, 2004).
In our recent studies, a difference in weaning weight of
73 kg resulted in a 40 kg difference in live weight and 24 kg
in carcass weight at 30 months of age (Greenwood et al.,
2005 and 2006; Table 6). The low weaning weight cattle
grew more rapidly during backgrounding and at a similar
rate in the feedlot, resulting in more rapid growth overall
from weaning to 30 months of age. However, compensation
in live weight remained incomplete by the conclusion of the
study. Similarly, in steers restricted in growth from birth to
weaning, then backgrounded to the same feedlot entry
weight as cattle grown rapidly to weaning, some compensatory growth was observed during backgrounding but
not in the feedlot (Cafe et al., 2006a). These studies have
confirmed earlier findings that severe, chronic nutritional
restriction to weaning limits compensatory growth, which
only occurred prior to feedlot entry and not in the feedlot,
resulting in smaller cattle and carcasses and less retail yield
of beef at an equivalent age.
Feed intake and efficiency. During feedlotting from 26 to 30
months of age, feed intake was lower among cattle grown
slowly to weaning than those grown rapidly; however, this
effect of pre-weaning growth rate was not evident when
assessed at the same feedlot entry weight (Table 7).
Differences in feed efficiency or residual feed intake were
not apparent on an age- or live-weight equivalent basis.
Consistent with these findings, when variation in live
weight that contributed to differences in energy requirements for maintenance and growth were accounted for, low
pre-weaning growth rates did not influence measures of
efficiency in the feedlot of cattle of equivalent age compared with those grown more rapidly prior to weaning
(Hennessy and Arthur, 2004). Furthermore, effects of early
post partum nutrition on growth, intake and efficiency of
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Greenwood and Cafe
Table 6 Consequences of growth from birth to weaning for growth and live weight characteristics of beef cattle to 30 months of age (adapted
from Greenwood et al., 2005 and 2006)Pre-weaning growth
Birth weight (kg)
Pre-weaning ADG (g)
Weaning (7 months) weight (kg)
Backgrounding ADG (g)
At equivalent age (26 to 30 months)
Feedlot entry (26 months) weight (kg)
Feedlot ADG (g)
Feedlot exit (30 months) weight (kg)
At equivalent feedlot entry live weight (500 kg)
Age at feedlot entry (day)Feedlot ADG (g)
Feedlot exit weight (kg)
Age at feedlot exit (day)y
Low (n 5 119)
High (n 5 121)
Significance of difference (P)
33.1
554
151
615
34.2
875
221
558
,0.001
483
1527
655
517
1570
695
,0.001
0.15
,0.001
789
1558
674
907
724
1540
676
841
,0.001
0.49
0.52
,0.001
-
Abbreviation is: ADG 5 average daily gain.
Values are predicted means from residual maximum likelihood (REML) analyses including effects of birth weight, pre-weaning nutrition, sex/year cohort, siregenotype and their interactions, with feedlot entry weight as a covariate (linear and, where significant, quadratic) to predict means at equivalent feedlot entry
weight.
Refer to Cafe et al. (2006b) and Greenwood et al. (2006) for details of pasture-based nutritional treatments and selection criteria for calves used to study
long-term consequences of growth early in life.
Predicted from mean ADG during backgrounding.
y
Predicted from mean ADG during background and mean feedlotting period.
-
Table 7 Consequences of growth from birth to weaning for feed intake and efficiency of beef cattle during feedlotting from 26 to 30 months of
age (L. M. Cafe and P. L. Greenwood, unpublished results)
Pre-weaning growth
Weaning weight (kg)
At equivalent age (26–30 months)
Feedlot entry (26 months) weight (kg)
Feedlot ADG (g)Feed intake (kg/day)Feed efficiency (kg DM intake/kg LW gain)
Residual feed intake (kg)At equivalent feedlot entry LW (490 kg)
Feedlot ADG (g)Feed intake (kg/day)Feed efficiency (kg DM intake/kg LW gain)
Low (n 5 75)
High (n 5 79)
Significance of difference (P)
149
215
477
1319
13.59
10.02
20.042
502
1356
14.24
10.36
0.040
,0.001
0.36
0.002
0.33
0.51
1317
13.89
10.13
1361
13.98
10.27
0.94
0.55
0.66
-
Abbreviations are: ADG 5 average daily gain, DM 5 dry matter, LW 5 live weight.
Values are predicted means from residual maximum likelihood (REML) analyses including effects of birth weight, pre-weaning nutrition, sex/year cohort, siregenotype and their interactions, with feedlot entry weight as a covariate (linear and, where significant, quadratic) to predict means at equivalent feedlot entry
weight. Difference in significance of feedlot ADG between Tables 6 and 7 is due to the number of cohorts studied (three in Table 6 v. two in Table 7) and the
duration of the measurement period (average of 117 days in Table 6 v. 70 days in Table 7).
During 70-day period in feed intake pens.
At mean metabolic live weight.
-
steers (Stalker et al., 2006) and heifers (Martin et al., 2007)
in the feedlot soon after weaning were not evident. These
results are consistent with earlier findings, reviewed by
Berge (1991), that feed conversion efficiency is little
affected in the long term by nutrition prior to weaning.
However, following extremely severe postnatal nutritional
restriction where calves were held near their birth weights
for 200 days, compared with cattle well grown to weaning,
1290
feed efficiency was adversely affected in males during
growth from 200 kg to about 400 kg live weight, whereas
during the same period females were more efficient (Tudor
and O’Rourke, 1980).
Body and carcass composition. At equivalent carcass
weight, there was more fat trim, less retail yield and there
Consequences of growth early in life of cattle
Table 8 Consequences of growth from birth to weaning for carcass characteristics of beef cattle at 30 months of age (adapted from Greenwood
et al., 2006)
Pre-weaning growth
At equivalent age (30 months)
Carcass weight (kg)
Retail yield (kg)
At equivalent carcass weight (380 kg)
Eye muscle area (cm2)
P8 fat depth (mm)
Rib fat depth (mm)
Aus-Meat marble score
USDA marble score
Longissimus IMF (%)
Ossification score
Retail yield (kg)
Bone (kg)
Fat trim (kg)
Low (n 5 119)
High (n 5 121)
Significance of difference (P)
368
242
393
254
,0.001
,0.001
90.1
20.1
10.4
1.92
450
6.88
202
251
67.8
52.8
89.2
20.8
11.0
1.77
441
6.98
199
246
66.7
57.8
0.55
0.41
0.33
0.15
0.49
0.80
0.53
,0.001
0.053
,0.001
Abbreviation is: IMF 5 intramuscular fat.
Values are predicted means from residual maximum likelihood (REML) analyses including effects of birth weight, pre-weaning nutrition, sex/year cohort, siregenotype and their interactions, with carcass weight as a covariate (linear and, where significant, quadratic) to predict means at equivalent carcass weight.
Refer to Table 6 for growth characteristics of the cattle.
tended to be less bone in the carcass among the cattle
grown rapidly compared with those grown slowly to
weaning (Greenwood et al., 2006; Table 8). This suggests
the greater fatness at weaning of the rapidly reared cattle
persisted to 30 months of age. However, because of failure
to compensate fully in weight, the carcasses from light
weaners remained smaller and weight of retail beef was
lower compared with the heavy weaners at the same age.
When cattle grown rapidly or slowly to weaning were
backgrounded to the same feedlot entry weight and
slaughtered after 120 days in the feedlot, their carcasses
did not differ in compositional and yield characteristics
(Cafe et al., 2006a).
Earlier studies within pasture-based nutritional systems
also failed to demonstrate substantial differences in body or
carcass composition due to nutrition and growth from birth
to weaning (Berge, 1991; Hearnshaw, 1997). These authors
concluded that cattle from low pre-weaning nutrition
groups generally have less fat than those from high preweaning nutrition groups, but if compared at a constant
carcass weight, differences in fatness usually disappear. As
a result, calves with lower weaning weights take longer to
reach carcass specifications than heavier calves.
In contrast to the above findings, severe nutritional
restriction to weaning that resulted in little growth post
partum, followed by concentrate (high energy) feeding from
weaning to slaughter resulted in greater fatness at the
same live and carcass weights compared with cattle well
nourished prior to weaning (Tudor et al., 1980). Within the
same study, cattle restricted or well nourished to weaning
then grown on pasture to the same slaughter weight did
not differ in composition. Factors likely to have contributed
to increased fatness among the small compared with large
weaners, which were subsequently fed concentrates,
include the following: greater length of time on concentrate
feed to reach the slaughter weight; greater weight-specific
intake of nutrients following the nutritional restriction; a
greater requirement for protein relative to energy at
weaning and, hence, potential nutrient imbalance in the
concentrate diet during the early post-weaning phase; and
more limited capacity for lean tissue accretion post weaning
in the small compared with large weaners.
Beef quality and myofibre characteristics. Differences in
objective measurements of meat quality between cattle
grown slowly or rapidly to weaning were not evident within
our research (Greenwood et al., 2006; Table 9). Similarly, in
earlier studies, objective measures of eating quality were
not adversely affected by restricted nutrition prior to weaning (Hearnshaw, 1997; Hennessy et al., 2001; Hennessy and
Morris, 2003). When they were affected, however, meat of
cattle from low-nutrition groups was usually more tender
than that of high-nutrition groups (Hearnshaw, 1997;
Hennessy et al., 2001). However, when compared at a
constant carcass weight, in about half of the studies the
meat quality differences became non-significant (Hearnshaw, 1997). Despite these findings, meat quality may be
compromised if the slow growth of cattle alter weaning
results in them being at least 8 to 9 months older at
slaughter weight (Loxton, 1997; Purchas et al., 2002). It is
unclear if similar age differences resulting from growth
restriction earlier in life have similar effects.
As with the above findings for meat quality, growth to
weaning had little overall affect on myofibre characteristics
in the longissimus lumborum muscle at 30 months of age,
apart from a small increase in the relative area of type 1
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Greenwood and Cafe
Table 9 Consequences of growth from birth to weaning for objective measurements of m. longissimus (striploin) and m. semitendinosus
(eye round) quality in beef cattle at 30 months of age (adapted from Greenwood et al., 2006)
Pre-weaning growth
Longissimus
Peak force (N)Compression (N)Cooking loss (%)
Ultimate pH
Colour L (lightness)
Colour a (red/green)
Colour b (yellow/blue)
Semitendinosus
Peak force (N)Compression (N)Cooking loss (%)
Low (n 5 119)
High (n 5 121)
Significance of difference (P)
40.5
14.2
21.8
5.48
39.9
26.5
13.7
39.2
14.1
21.5
5.47
39.6
26.6
13.7
0.25
0.63
0.23
0.60
0.48
0.70
0.93
46.3
22.1
21.5
46.3
22.8
21.3
0.89
0.47
0.45
Values are predicted means from residual maximum likelihood (REML) analyses including effects of birth weight, pre-weaning nutrition, sex, sire-genotype and
their interactions. Refer to Tables 6 and 8 for growth and carcass characteristics of the cattle.
Objective measures of texture, as defined by Perry et al. (2001).
-
Table 10 Consequences of growth from birth to weaning for longissimus lumborum myofibre characteristics of heifer beef cattle at 30 months of
age (P. L. Greenwood and L. M. Cafe, unpublished results)
Pre-weaning growth
Weaning weight (kg)
Live weight at 30 months (kg)
Carcass weight (kg)
Muscle weight (g)
Muscle CSA (cm2)
Myofibres per mm2
Average myofibre CSA (mm2)
Total myofibre area (%)
Type 1
Type 2C
Type 2A
Type 2AX
Type 2X
Low (n 5 40)
High (n 5 38)
Significance of difference (P)
138
612
345
4191
87.3
132.9
5585
203
637
360
4358
88.4
124.4
5854
0.025
0.032
0.14
0.58
0.27
0.32
23.9
0.59
22.6
6.58
46.4
21.1
0.67
22.7
5.30
50.2
0.017
0.75
0.89
0.28
0.24
Abbreviation is: CSA 5 cross-sectional area.
Values are predicted means from residual maximum likelihood (REML) analyses including effects of birth weight, pre-weaning nutrition, sex, sire-genotype and
their interactions. Refer to Lehnert et al. (2006) for myofibre classification and measurement methodology. Type 1, type 1 myosin heavy chain (MHC) slow
oxidative; Type 2C, intermediate between type 1 and type 2A; Type 2A, type 2A MHC fast oxidative-glycolytic; Type 2AX, intermediate between type 2A and
type 2X; Type 2X, type 2X MHC fast glycolytic.
myofibres in cattle grown slowly to weaning compared with
those grown rapidly (Table 10). This is despite differences
due to pre-weaning growth in the size of each myofibre
type and the percentages and relative area of fast (type 2)
myofibres at weaning (P. L. Greenwood, unpublished
results).
Interactions between in utero and pre-weaning
growth
Among the numerous beef production characteristics that
we investigated, the only interaction between growth
1292
in utero and growth prior to weaning was for the eye
muscle (m. longissimus) cross-sectional area when compared at an equivalent carcass weight at 30 months of age
(Greenwood et al., 2006). Cattle of low birth weight had a
greater eye muscle area at slaughter than high-birth-weight
cattle within the high pre-weaning growth group (91.1 v.
87.2 cm2), suggesting, perhaps, in conjunction with the
results for subcutaneous fat depth, some long-term consequences of divergent foetal growth for distribution of
carcass tissues. However, the eye muscle area did not differ
due to birth weight within the animals that grew slowly
to weaning (89.8 v. 90.5 cm2, respectively). Similarly,
interactions between prenatal and pre-weaning nutrition for
Consequences of growth early in life of cattle
post-weaning growth, feed intake, feed efficiency and carcass characteristics were not evident in the study of Stalker
et al. (2006).
Interactions between growth early in life and gender
There appear to be few studies of interactions between
growth early in life and the gender of cattle on beef production characteristics. Growth of well nourished, artificially
reared calves (Tudor and O’Rourke, 1980) and feed efficiency after weaning have been shown to be influenced by
gender following maternal nutritional restriction (Tudor and
O’Rourke, 1980). The results of Stalker et al. (2006) and
Martin et al. (2007) also suggest differences between
genders in the efficiency of nutrient utilisation following
divergent maternal nutrition during late pregnancy. However, few interactions between gender and nutrition early in
life have been demonstrated for subsequent body and
carcass characteristics, with these relating mainly to bone
growth (Tudor et al., 1980). Within our recent studies,
interactions between birth weight and gender/year cohorts
were evident for carcass weight, eye muscle area, and
weight of bones and retail beef at 30 months of age
(Greenwood et al., 2006). However, while suggestive of
interactions between sex and growth early in life, our
experimental design did not allow for this interaction to be
specifically tested.
Interactions between growth early in life and
sire-genotype
A major objective of our research has been to determine the
extent to which genotype may interact with nutrition early
in life to influence subsequent growth, carcass, yield and
beef-quality characteristics. To achieve this objective, our
research included offspring of Piedmontese (a high muscling, high-birth-weight breed homozygous for a non-functional mutation in myostatin) and Wagyu (a high marbling
and lower birth weight breed) bulls. Perhaps surprisingly, no
interactions between sire-genotype and growth early in life
were evident for any production parameters reported here
or presented by Greenwood et al. (2006).
Conclusions
Severe, chronic growth retardation of cattle early in life is
associated with reduced growth potential, resulting in
smaller animals at any given age. The capacity for long-term
compensatory growth diminishes as the age of onset of
severe nutritional restriction resulting in prolonged growth
retardation declines, such that more-extreme intrauterine
growth retardation can result in slower growth throughout
postnatal life. However, within the normal limits of beef
production systems, neither restricted growth in utero nor
from birth to weaning influences the efficiency of nutrient
utilisation later in life.
Retail yield from cattle severely restricted in growth
during pregnancy or from birth to weaning is reduced
compared with cattle well grown early in life, when
compared at the same age later in life. However, retail yield
and carcass composition of low- and high-birth-weight
calves are similar when compared at the same carcass
weight.
At equivalent carcass weights, cattle that are grown
slowly from birth to weaning have carcasses of similar or
leaner composition than those grown rapidly. However,
there is evidence to suggest that if high energy, concentrate
feed is provided following severe growth restriction from
birth to weaning, then at equivalent weights post weaning
the slowly growing, small weaners may be fatter than their
well-grown counterparts.
Restricted prenatal and pre-weaning nutrition and
growth do not adversely affect measures of beef quality
including shear force, compression, cooking loss and colour.
Similarly, bovine myofibre characteristics are little affected
in the long term by growth in utero or from birth to
weaning, despite specific myofibre-type-related effects at
birth and weaning, respectively.
Hence, economic benefits resulting from adequate
maternal nutrition, especially during pregnancy, to optimise
growth of offspring to market weights are primarily due to
advantages in carcass weight and retail beef yield at a
given age, reduced feed costs to reach a given market
weight, stocking rates and subsequent reproductive rates of
breeding cows, but not due to differences in beef-quality
characteristics (Alford et al., 2007).
Interactions were not evident between prenatal and preweaning growth for subsequent growth, efficiency, carcass,
yield and beef-quality characteristics, within our pasturebased production systems. Interactions between genotype
and nutrition early in life studied using offspring of Piedmontese (a high muscling, higher birth weight breed,
homozygous for a mutation that produces non-functional
myostatin) and Wagyu (a high marbling, lower birth weight
breed) sires mated to Hereford cows were not evident for
any growth, efficiency, carcass, yield and beef-quality
parameters.
We propose that within pasture-based production
systems for beef cattle, the plasticity of the carcass tissues,
particularly of muscle, allows animals that are growthretarded early in life to attain normal carcass composition at
equivalent weights in the long term, albeit at older ages.
This may well relate to regulation of nutrient intake to a
level appropriate for the size and lean tissue growth
capacity of the animal, coupled with the capacity of the
myosatellite cell population to generate myonuclei in
support of muscle growth over a prolonged recovery period,
as discussed previously (Greenwood et al., 1998, 1999 and
2000a). However, the availability of feed and quality of
nutrition during recovery from severe growth retardation
early in life may be important in determining the subsequent composition of young, light-weight cattle relative
to their heavier counterparts.
1293
Greenwood and Cafe
Finally, it needs to be emphasised that long-term consequences of more specific, acute environmental influences
during specific stages of embryonic, foetal and neonatal calf
development remain to be determined. This need for further
research extends to consequences of nutrition and growth
early in life for subsequent reproductive performance, which
has been recently shown to be affected in heifers by
nutrition of their dams during late pregnancy (Martin et al.,
2007), although neither reproductive or lactational performance were affected by birth weight (Swali and Wathes,
2006).
Acknowledgements
The contributions of Ms Helen Hearnshaw, Dr David Hennessy
and Professor John Thompson in the conduct of research on
consequences of growth and nutrition during early life of cattle
described herein are most gratefully acknowledged. The
financial and in-kind support of the Cooperative Research
Centre for Cattle and Beef Quality, NSW Department of Primary Industries, CSIRO Livestock Industries and the University
of New England is gratefully acknowledged. We also
acknowledge the considerable efforts of research, technical
and/or farm staff of NSW Department of Primary Industries
Agricultural Research and Advisory Stations at Grafton and
Glenn Innes and its Beef Industry Centre of Excellence in
Armidale, at the Beef CRC ‘Tullimba’ Feedlot, at CSIRO Livestock Industries, Queensland Bioscience Precinct, St Lucia, and
at the University of New England Meat Science Complex, in
the conduct of the Beef CRC research described in this review.
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