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Asian-Aust. J. Anim. Sci.
Vol. 22, No. 5 : 756 - 764
Perinatal Nutrition of the Calf and Its Consequences for Lifelong Productivity*
P. C. Wynn1,
**, H. M. Warriach1, 2, A. Morgan3, D. M. McGill1, S. Hanif1, 2, M. Sarwar4
A. Iqbal5, P. A. Sheehy3 and R. D. Bush3
1 EH Graham Centre for Agricultural Innovation (Charles Sturt University and NSW Department of Primary Industries)
Locked Bag 588, Wagga Wagga, NSW 2678, Australia
Provision of an optimal environment for the calf is critical to establishing the patterns of growth and development
essential to allow the heifer to express its genetic potential for milk output and reproductive capacity during its productive life. Maternal
nutrition during gestation is now recognised as a key to genetic programming in utero
and this influence is extended through the
complexity of hormones, growth factors and immunostimulants incorporated into colostrum and milk consumed by the neonatal calf.
This natural process is most often disrupted as calves are weaned abruptly to maximise milk output for commercial exploitation. The key
then is to accelerate the rate of maturation of the ruminal epithelium through the provision of concentrate starter rations and high quality
forage, which promote VFA production. Management systems to promote these processes in Holstein Friesian cattle are well developed,
however, little is known of these processes with buffalo and Bos indicus
dairy cattle such as the Sahiwal. The development of methods
to program the neonate to grow faster to puberty in these species will be important to improving their productivity for the dairy
industries in tropical and sub-tropical environments in the future. (Key Words :
Perinatal Nutrition, Calf, Buffalo, Productivity)
source of dietary energy and protein through to the point
when the calf is able to be weaned. This initial phase is
The potential for growth and high health status in the most often termed the pre-ruminant period during which
newborn calf is largely influenced by the health and milk is passed directly into the abomasum through the metabolic status of their dam. Much of the focus of cow reflex closure of the oesophageal groove. This initial period management has been on the perinatal period as the calf of development is termed the pre-ruminant phase and varies prepares for delivery into a totally foreign environment in in duration from 14-21 days depending on the animal’s which placentally derived nutrition is replaced by the initial ability to initiate the intake of dry feed. During the lacteal secretion from the mammary gland, colostrum. The subsequent 3-6 weeks ruminal function develops and the composition of this secretion is extremely important in calf derives more of its nutrient substrate from this source establishing the growth potential and life-long productivity than from milk. At this point the animal is weaned and of the calf. This then reverts to normal milk which acts as a derives its nutrient substrate solely from dry feeds through
the activity of the newly established ruminal microbial
* This paper was presented at the 5th International Symposium on
population: this describes the ruminant phase of calf
Recent Advances in Animal Nutrition during the 13th Animal
Sciences Congress, Asian-Australasian Association of Animal
This process is universal for bovine species although
Production Societies held in Hanoi, Vietnam (September 22-26th,
there are many variations in procedures used in different
2008). ** Corresponding Author: P. C. Wynn. E-mail: [email protected]
2 Livestock and Dairy Development Board, 30 Tariq Block, New
3 Faculty of Veterinary Science, University of Sydney, Camden
4 Institute of Animal Nutrition and Feed Technology, University
Feeding cows to support the protein, energy, vitamin
and mineral requirements of the growing conceptus is now
Department of Livestock Management, University of
recognised as just one factor that influences the life-long
Wynn et al. (2009) Asian-Aust. J. Anim. Sci. 22(5):756-764
productivity of the calf. We are now aware of the proteins, essential and non essential amino acids, lactose, importance of environmental factors on gene expression fatty acids, vitamins and minerals all contributing to these patterns and therefore development much earlier during requirements. Other non-nutritional factors include embryogenesis. These so-called epigenetic mechanisms nucleotides, polyamines, enzymes as well as functionally challenge the very basis of Darwinian evolutionary theory important proteins such as lactoferrin (Blum and Hammon, that the variability in populations occurs exclusively 1999; 2000) serve specific roles in directing growth through random mutations. The mechanism through which processes. Lactoferrin has also been extensively this occurs is by altering gene methylation patterns (Khosla
characterized in buffalo milk (Sharma
et al., 1999). et al., 2001) which might not only regulate the growth
Cellular components are also incorporated including
potential of the calf but also alter the germline which may mammary epithelial cells, erythrocytes macrophages, then persist as mutations across subsequent generations polymorphs, lymphocytes, plasma cells and epithelial cells (Surani, 2001). Factors that initiate these changes which (Uruakpa
et al., 2002) while a number of the non-nutritional may persist differentially across different regions of DNA and bioactive molecules are sequestered from the are influenced by changes in dietary components providing circulation. These include growth hormone, prolactin, methyl groups for this mechanism. Synthetic xenobiotics oestrogen, insulin and glucagon, all of which play a role in (Danzo, 1998), the estrogenic molecules in plants the regulating protein and energy metabolism. It is often phytoestrogens and isoflavones, all of which can be found difficult to envisage a functional role for these protein in the cow’s diet contribute to these changes (Guerrero-
hormones since they will be hydrolysed extensively in the
abomasum prior to accessing functional receptors. However the key may reside in their co-secretion with the
Thus at calving the growth potential of the animal may immunoglobulins, most notably IgG1.
have already been compromised by factors other than the
These macromolecules comprise more than 90% of the
mere supply of nutrient substrate to support the growth protein content of colostrum and appear at concentrations 5
-10-fold higher than in the circulation (Larson, 1992). Their
THE IMPORTANCE OF COLOSTRUM
sequestration coincides initially with higher oestrogen levels up to 1 month pre-partum
and subsequently with
elevated corticosteroids, growth hormone and prolactin in
It is generally considered that the newborn calf should the last week and then with depressed progesterone at 48 h
receive 4 L of colostrum in the first 12 h, although up to 6 L pre-calving.
is often recommended for the first day (University of Sydney, 2007). Requirements will depend on the quality of
The functional significance of these relationships is yet
to be established. Equally intriguing is the mechanism that
colostrum usually determined by assessing its density facilitates preferential uptake of IgG1, which is typically
associated with immunoglobulin (Ig) content: the presence 10-fold higher than IgG2 in colostrum, but present in
of enzymes to form curd in the abomasum is also a rate equivalent concentrations in the circulation. This imbalance
limiting factor for Ig absorption (Gregory, 2003; Mastelloni provides a definitive characteristic of colostrum and is
The observation that over 100 hormones and growth explained by the presence of specific receptors on the basal
membrane of secretory epithelial cells which actively
factors have been identified in colostrum or milk endocytose IgG1 and pass it to the secretory lumen of the
(Koldovsky, 1995; Koldovsky, 1996) suggests that these alveolus (Butler, 1983; Barrington
et al., 2001). It is
secretions are complex biological fluids designed to extend important to note that this mechanism is only in place
the influence of the dam over developmental processes beyond the uterine environment. These include hormones of during colostrum synthesis and therefore IgG1
concentration falls markedly after the colostral phase
the hypothalamic-pituitary, the thyroid-parathyroid group, (Kemler
et al., 1975). The immune status of the buffalo
gastrointestinal regulatory hormones as well as growth calf post-colostrum feeding is influenced also by the
factors. While the physiological significance of growth vitamin status of the dam: circulating Ig levels were
hormone regulatory peptides is most apparent the increased by 80% in calves fed colostrum from their dams
importance of the gonatropin regulatory peptide GnRH at this early stage is more obscure. However the receiving bolus injections of vitamins A., D3 and E late in
pregnancy (Sikka and Lal, 2005). VitE and selenium also
developmental processes that these hormones regulate are limit the adverse effects of endotoxin from E coli
most likely important when calves are born into a associated with many calf rearing systems (Sharma et al.,
challenging environment in which ambient temperature 2005).
varies significantly from that experienced in utero
and when pathogen loads are high. The provision of key nutrients to
Further protection against pathogenic bacteria and
viruses is provided by the presence of antimicrobial
support growth is also an integral function of milk with
Wynn et al. (2009) Asian-Aust. J. Anim. Sci. 22(5):756-764
proteins, lactoferrin and lysozyme. Lactoferrin is an iron memory in the piglet (Wang
et al., 2007). There is little binding moiety that prevents microbial growth through reason to suspect that suckling behaviour in both cows and depriving microbial of this essential mineral and by binding buffalo is not influenced by these molecules, although to bacterial cell membranes thereby compromising their bovine and human milk diverges in the concentration and permeability (van Hooijdonk
et al., 2000), while lysozyme composition of their sialyloligosaccharide content: this has lyses bacterial cell walls (Lonnerdal, 2002). Interestingly implications for the development of infant formulae these two proteins are capable of acting synergistically to (Martín-Sosa
et al., 2003). Interestingly buffalo milk enhance their bacteriostatic activity (Pakkanen and Aalto, gangliosides appear to have greater toxin binding and anti-1997).
inflammatory properties than cows milk suggesting some
The immunoglobulins are accompanied by a range of potential novel applications for the product from buffalo in
protease inhibitors including trypsin inhibitor, α2-
the future (Colarow
et al., 2003). This is also important for
macroglobulin, α2-antiplasmin, antithrombin III, C1-
the survival of the buffalo calf in environments with high
inhibitor, inter-α-trypsin inhibitor, bovine plasma elastase pathogen loads. Certainly buffalo milk is highly valued as inhibitor and bovine plasma trypsin inhibitor, all of which an alternative to breast milk among Indian mothers serve to protect their functional integrity (Christensen
et al., (Kaushal
et al., 2005). 1995).
The composition of colostrum changes rapidly and its
Other multi-functional proteins are coming to light provision to the calf during the first 24 h of life is critical to
including a proline-rich polypeptide colostrinin. This was the calf’s survival. originally found as a fraction accompanying sheep colostral
immunoglobulins which promoted T cell-tropic and
COLOSTRUM AND THE DEVELOPMENT
maturational activity. It is also associated with the
OF THE GASTROINTESTINAL TRACT
development of precognitive functions which inhibit
pathological states centrally (Zimecki, 2008).
The rich mix of hormones, growth factors, cytokines
In addition to the role of the simple carbohydrate lactose and nutrients in colostrum provide the ideal developmental
in providing energy, more complex carbohydrates also add mix to initiate digestive activity in the abomasum, small to the multi-functionality of colostrum. Sialyloligosaccharides and large intestines. The initiation of the functional integrity are present in high concentrations for the first 12 h of of the intestinal epithelium is essential for the absorption of lactation in the cow and are thought to be important in nutrients and bioactive molecules to direct developmental preventing infections acting against rotavirus, rheovirus and processes in the body. In particular the apical junctional Helicobacter pylorum
et al., 2003). Interestingly complex plays an important role in maintaining the integrity they also appear to be involved in the development of of this epithelium and prevents access for pathogens to the cognitive processes in the brain, with the supplementation circulation. A wide range of cytokines and growth factors of milk for piglets with sialic acid improving learning and influence tight junction integrity, with IFN-γ, TNF-α, HGF,
The composition of colostrum and milk (Blum and Hammon, 2000; Klimes
et al., 1986)
Mature milk (5-14days postpartum
Wynn et al. (2009) Asian-Aust. J. Anim. Sci. 22(5):756-764
TGF-α, IGF-I, IGF-II, VEGF, IL-1, IL-4 and IL-13 all and shade for calves is well established yet not always decreasing barrier function while EGF, TGF-β, GDNF, adhered to in many countries. neurturin, IL-10, and IL-17 have the opposite effect
The water content of body tissues is in the vicinity of
et al., 2003). Clearly this is a closely regulated and 70% (Diaz
et al., 2001), thus the constant availability of functionally important property which can be influenced by water is mandatory in any production system. Promoting the balance of these factors present in colostrum. Again the intake of dry concentrate feeds to enhance rumen there is evidence to suggest that the intestinal epithelium in development is also dependent on constant water the buffalo calf is more resistant to some infections availability. including paratuberculosis (Sivakumar
et al., 2006).
Requirements for minerals and vitamins have also been
documented (Council, 2001). Whole milk provides an
adequate source of all minerals with the exception of iron
FOR THE NEWBORN CALF
and sometimes selenium and manganese. As most milk
starters are supplemented with minerals and fat soluble
The calf requires nutrients for both maintenance and vitamins these rarely compromise calf health and growth.
growth and it is important that the requirements for these
The only major concerns are with Vitamins A and E:
two processes are combined. Environmental factors are NRC recommendations for Vitamin A are considered to be extremely important in determining requirements with both too high leading to potential toxicity, while those for extremes of heat and cold, high pathogen loads and physical vitamin E are under-estimated (Drackley, 2008). While a and psychosocial stressors contributing to requirements. state of deficiency will rarely be found, finding the correct The activation of the immune system and the role that dose for optimal growth represents a greater challenge. maternal immunity plays in this process is important to
animals calving in sub-optimal environments (Chase
DEVELOPING THE RUMEN
As a guideline the metabolisable energy requirements
The general rule of thumb is to adopt strategies that
for a 45 kg calf under thermoneutral conditions is 7.3 allow calves to consume 700-900 g of concentrate ration by MJ/day. Since cow’s milk contains 22.5 MJ ME/kg of solids the time that they are weaned. Essentially this requires they require 2.5 L of whole milk, while the equivalent with calves to commence intake of concentrates within 14 days lower fat status milk replacers is around 3 L (Drackley, post-partum
(University of Sydney, 2007).
Animals 2008). Others recommend higher intakes up to 10 and 12% consuming 10% of their own bodyweight as milk per day of bodyweight per day to support growth (University of will in general be consuming up to 300 g of concentrate by Sydney 2007). Milk requirements will be lower in the day 25. The microbial population accumulating in the buffalo as milk fat content is higher than in cows milk, rumen will ferment carbohydrate to form predominantly although colostral fat content is the same in each species butyric acid and then propionic acid which in turn promote (Ganovski, 1979).
the differentiation of the ruminal epithelium to form the
Protein requirements for maintenance are low in the characteristic papillae (Heinrichs and Lesmeister, 2005).
neonate (30 g/d for a 45 kg calf) and reflect rates of protein Different sources of fermentable carbohydrate yield turnover in tissue. However the requirements for growth are different responses, with corn and wheat based diets approximately 6-fold higher than this, equating to 250-280 promoting ruminal development faster than oats or barley g of crude protein from milk replacer (Drackley, 2008). In (Khan
et al., 2008). Similarly processing can exert and general milk replacers containing up to 25% crude protein influence the growth response: steam flaking of corn for are recommended as long as dietary energy is not limiting. example induced ruminal epithelial development faster than Amino acid composition of replacers is also important with either dry rolling on leaving grain whole (Lesmeister and those most closely resembling the composition of cows Heinrichs, 2005). These studies showed also that processing milk being most effective.
can influence the pattern of volatile fatty acids released.
Changes in energy requirements in cold and hot Other alternative concentrate sources such as phalaris
environments are quite dramatic as body temperature is not minor
seeds have also been assessed as providing buffered by the heat of fermentation in the undeveloped appropriate carbohydrate sources to support volatile fatty rumen. Maintenance ME increases by approximately 20% acid synthesis and rumen development (Kaur et al., 2006). for each 10°C incremental decrease from 20 to -20°C
Much has been written on the role of fibre and the so
(National Research Council, 2001). In contrast the effects of called “tickle factor” in the diet of the pre-ruminant calf. heat stress have not been published although older animals Results vary widely with concentrate per se
being more require an additional 20-30% ME (National Research effective in some studies (Klein
et al., 1987), while in Council, 2001): the ready availability of water on demand others forages provided at a specific particle size (8-19 mm)
Wynn et al. (2009) Asian-Aust. J. Anim. Sci. 22(5):756-764
with concentrates gave superior results (Coverdale
et al., dam. This increment decreased to 5.5 g per kg liveweight in 2004). In further studies pelleted diets yielded superior dams weighing 576-815 kg (Usmani
et al., 1987; Usmani responses to mixed length fibre with other dietary and Inskeep, 1989). ingredients held constant (Bach
et al., 2007). Clearly our
As with the cow the first 24 h post-calving are critical
understanding of the development of the ruminal for the buffalo calf to absorb colostrum. In fact total protein environment requires further investigation, although the and Ig levels are higher in the buffalo than in cross-bred role of the volatile fatty acids in this process is well cows (Singh and Ahuja, 1993). In this study 75% of Ig and established. The inclusion of cellulolytic enzymes as feed 68% of colostral protein were absorbed within 1 h of additives also provides a beneficial growth effect if feeding a 7 h old calf. This rate of absorption declined included as a substitute for more conventional additives in rapidly after the first feed. However little other data are buffalo calf diets (El-Kady et al., 2006).
available on colostrum usage to account for high
susceptibility to infection of buffalo calves.
CALF REARING: THE PAKISTANI EXPERIENCE
Pakistan, like many developing countries has an
agrarian rural based economy. The livestock sector is a
Neonatal calf morbidity and mortality are major causes
major contributor to the national (12%) and agricultural of economic losses in livestock production. It is roughly (50%) economy (Pakistan Economic Survey, 2006). This estimated that a calf mortality of 20 percent can reduce the sector is growing quickly and provides a livelihood for net profit of an enterprise by 60% (Blood and Radostits, more than 35 million people. The productivity of livestock 1989). Ideally calf mortality should be less than 5 percent for meat and milk is low, with improper calf management with growth rates of 0.5-0.7 kg/d (Blood and Radostits, programming animals for a life of low productivity. High 1989). Mortality rates for different countries employing market prices for milk dictate that calves are weaned very different production systems are detailed in Table 2. early without appropriate quality milk replacers being used
Very high mortality rates of over 50% have been
to meet the demand of the calf for growth and development. reported in buffalo calves to one month of age. Foot and Male calves are most often sold for slaughter or left to feed mouth disease (FMD) and haemorrhagic septicaemia (HS) on poor quality roughages. The slow growth of heifer calves are endemic to Pakistan and account for up to 31 and 21.5% results in delayed puberty and age at first calving. Thus respectively of deaths in buffalo calves aged from 6-12 both the efficiency of milk and beef production are months (Ramakrishna, 2007). On other farms extreme compromised.
mortality rates of up to 80% have been recorded (Tiwari
al., 2007). Although disease contributed to this statistic, the
FEEDING THE TRANSITION BUFFALO
failure to provide colostrum, to deworm, to disinfect naval
cords and to provide an adequate milk substitute and
Late pregnant or transition Holstein cows can be appropriate shelter and water all played their role in the
affected by a range of production diseases associated with etiology of these mortalities. Other causes of calf mortality their inability to cope with the metabolic demands of high include the greater susceptibility of crossbred and production. These include hypocalcaemia, hypomagnesaemia, primiparous animals (Rao and Nagarcinkar, 1980). The ketosis, retained placenta, displacement of the abomasum failure to provide colostrum has also often been implicated and laminitis (Mulligan and Doherty, 2008). These are often (Afaq
et al., 1992). associated with an imbalance in metabolites entering key
The giving of colostrum to friends is a custom found in
biochemical pathways (Payne, 1972) which may lead to some regions and as such the calf is inevitably deprived. infertility.
Overall farmers in many regions consider calf rearing a
Similar problems are associated with buffalo production, very low priority, as the commercial value of this practice is
although the causes are most likely related to under-
nutrition. The pregnant buffalo needs to support the nutrient
demands of both lactation and growth of the foetus. Yet the
SUCKLING AND HAND FEEDING
condition score of most small-holder buffalo remains very
OF BUFFALO CALVES
low despite the need for additional nutrients to meet these
demands. Very few studies have been conducted to
The average birth weights for buffalo and Sahiwal
investigate these relationships, but a positive relationship calves are 34.85±0.46 and 21.87±0.20 kg respectively has been shown between liveweight of the calving dam and (Ahmad, 1988). The method of provision of milk for these the calf. At lower body weights (350-573 kg) calf birth calves has been shown to influence their growth efficiency. weight increased by 18 g for each kg increase in weight of For example calves reared by restricted suckling of their
Wynn et al. (2009) Asian-Aust. J. Anim. Sci. 22(5):756-764
Incidence of mortality (%) among buffalo and cow calves in different countries
dams yielded better growth rates than if the milk was in these studies are below recommendations for Holstein provided in a feeder or pale (552 vs. 370 g/d) (Khan and Friesian calves: thus productivity could be boosted further Preston, 1992) and 500 vs. 350 g/d in the study of (Gaya
et through the use of higher protein supplements. The al., 1977). If adequate milk is provided these high growth importance of offering calves a soluble concentrate ration rates are attainable: the provision of 15% of the milk high in protein and energy from calving should be production from Sahiwal cows or buffalo which equates to considered. 10% of the calf’s bodyweight has resulted in these growth rates (Ahmad, 1988). The training of buffalo calves to the
FEEDING POST WEANING
use of automatic suckling units may compromise their
growth performance by limiting intake (Rossi
et al., 2004).
Feeding strategies used post-weaning involve the use of
Given these problems it is most likely more effective to low quality crop residues, straws and stovers characterized delay the weaning of buffalo calves reared in sub-optimal by high fibre and low crude protein. Numerous studies have environments particularly as the buffalo dams display been undertaken to improve their efficiency of utilization strong maternal instincts Thus the provision of some milk to largely through their treatment or use in conjunction with calves combined with the harvest of milk from a second strategic supplements (Sarwar
et al., 2002). milking each day for commercial sale or home consumption
Urea treatment of straw is popular as it increases the N
may provide the most effective means of rearing the calves content of roughages. The addition of molasses then
of Sahiwal cattle and buffalo. This method also negates the provides a balance of N and energy for the rumen microbial
need to use oxytocin to induce milk let-down: use of the population to utilise in digesting the insoluble
carbohydrates. Wheat straw treated with varying levels of
urea (0%, 2% and 4%) and molasses (2% and 4%) ensiled
STARTER RATIONS FOR CALVES
with 30% cattle manure (on dry matter basis) for different
fermentation periods (20, 30 and 40 days) proved to be an
Poor growth rates result from the limited milk supply, ideal supplement providing linear growth responses with
low protein and energy of fodders and concentrates that are amount fed in buffalo calves (Sarwar
et al., 2006). Similar
available for calves. In order for the farmer to sell a high results were achieved by Khan et al. (1992) who reported
proportion of his milk, cost-effective milk replacers need to be developed. Starter rations containing 17% crude protein that crude protein increased by 18.4% to 22.2% in and total digestible nutrients of 75% have resulted in sugarcane bagasse ensiled with cattle manure for 30 and 60 growth rates of up to 470 g/d (Ahmad and Jabbar, 2000).
days, respectively. However, the increasing cost of urea
Similar results have been achieved through the world-wide may make this option prohibitive.
substitution of milk with soybean milk containing 1%
Caution must be used in the evaluation of supplements.
soybean oil (Matter
et al., 2005). The levels of protein used In one study sunflower meal was substituted for cottonseed
Wynn et al. (2009) Asian-Aust. J. Anim. Sci. 22(5):756-764
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