Long-term effects of feeding during calfhood on subsequent performance in beef cattle (a review)

Livestock Production Science, 28 ( 1991 ) 179-201 Elsevier Science Publishers B.V., A m s t e r d a m

179

Long-term effects of feeding during calfhood on subsequent performance in beef cattle (a review) P. Berge Station de Recherches sur la ~)ande, 1. N.R.A. Centre de Recherches de (Termont-Ferrand/Theix, 63122 St-Genes-Champanelle, France (Accepted 23 July 1990)

ABSTRACT Berge, P.. 1991. Long-term effects of feeding during calfhood on subsequent perlbrmance in beef cattle (a review). Livest. Prod. Sci., 28: 179-201. A review of the literature was undertaken to relate the fattening performance and the carcass quality traits in beef cattle to the feeding strategy applied during the first year of the calf's life. In all cases an early growth limitation causes major changes in body composition. Once allowed subsequently to recover on a liberal regime, the calves have a greater food dr)' matter intake and a greater food conversion efficiency than unrestricted calves. But contrary to that of animals restricted at a more advanced age, their potential for subsequent compensatory growth, particularly when restricted before weaning, is low and practically independent of the severity of the restriction. At slaughter at a fixed commercial carcass weight, the initial differences in body composition are generally no longer perceptible, thus suggesting that the feeding treatments during calflaood have no long-term effect on this parameter. It is concluded that there is no evidence that the composition of beef carcasses at slaughter can be manipulated at an early stage of development by means of feeding techniques. Keywords: beef cattle; calf feeding: compensatory growth: carcass composition.

INTRODUCTION

The price of beef carcasses in Europe is traditionally determined both by conformation and fat scores. The best classification is given to well conformed and heavy carcasses containing a moderate proportion of fat such as those produced by animals of late-maturing breeds (e.g. Charolais, Limousin). On the other hand, carcasses of early-maturing breeds (e.g. dairy breeds such as Holstein, Friesian, Normand ) are given lower prices due to their lower muscular development and their greater adiposity (Robelin, 1986a). So the limitation of fat deposition has become one of the major concerns of producers as a mean of increasing the weight and the commercial value of beef carcasses.

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© 1991 - - Elsevier Science Publishers B.V.

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Many works have demonstrated that a reduction in the feeding level during the late productive phase of beef cattle contributes to limit body fat deposition, particularly in dairy breeds (Geay et al., 1976; Geay and Robelin, 1979; Robelin, 1979; Price et al., 1984). During the fattening period, the rate of fat deposition after 250-300 kg live weight is consistently greater in this type of cattle than that in animals of late-maturing breeds (Robelin, 1986b). The advantage of a feed restriction, however, is questionable as a longer fattening period is required to reach a fixed carcass fat proportion at slaughter while the carcass weight is generally little increased. Less information is available on the long-term effects of a manipulation of beef cattle at an early stage of life on their subsequent performance until slaughter. Most of the works carried out in the past have investigated the shortterm effects of level of feeding a given basal feed (milk or forage) or those of level of supplementation by concentrates. The interaction of the various treatments applied throughout the successive phases of growth, fattening and finishing actually makes more difficult the understanding of their respective effects and the comparison of the results in the literature. The aim of this paper is to review the most relevant works undertaken in beef cattle in the field of growth, voluntary feed intake, feed conversion efficiency and body composition as affected by the feeding strategy applied during calfhood. It will refer to the results obtained in animals of less than 1 year of age at the start of the experiments and will consider successively the three main periods that can be defined after the calf's birth (milk feeding, weaning and post-weaning). 1. P E R I O D O F M I L K F E E D I N G

During the pre-weaning phase, a limitation of the quantity of milk fed is known to reduce the calf's live weight gain. But its effect on body composition is much more variable. This can be partly explained by the fact that calves of early-maturing breeds are generally weaned early (2 to 6 months) after intensive milk feeding while calves of late-maturing breeds are suckled at pasture and weaned at an age of 8 months or more. Since one may suspect the experimental results of being biased by an interaction between breed maturity and production system, these two situations will be studied separately.

1. I. Earl.v-weaned calves Body composition and weight gain of calves fed milk artificially and weaned before 6 months of age are known to depend upon the level of milk intake. Hooven et al. (1972) and Robelin and Chilliard (1989) found that a 40% fall in weight gain due to a restriction on milk was followed by a 50% decrease in body fat at 3 to 6 months of age in calves of various dairy breeds. But when

EFFECTS OF FEEDINO DURING CALFHOODON PERFORMANCE IN BEEF CATTLE

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Butterfield et al. ( 1971 ) compared calves at the same live weight, carcass composition was not dependent upon the milk feeding treatment. After a milk restriction, calves generally exhibit little or no compensatory growth when turned out to pasture (Wardrop, 1966; Kaiser, 1976; Giovanni, 1982 ) or fed fattening diets of concentrate and hay (Bond et al., 1972; Meadowcraft and Yule, 1976 ). Some authors even found a positive correlation between the growth rate during a milk-feeding period from birth to 2 or 4 months of age and the subsequent live weight gain (Everitt and Jury, 1977; Folman, 1977; Harte and Fallon, 1984). Over a longer recuperation period such as in the 20- to 30-month beef systems, no more compensation can be expected (Lonsdale and Tayler, 1969: Everitt, 1972; Reardon and Everitt, 1972; Harte et al., 1981; Harte and FalIon, 1982, 1983 ). However, mention should be made of the results obtained by Tudor and O'Rourcke (1980) suggesting a possible interaction between the pre-weaning level of feeding and the type of feed during recuperation. A difference of 12~5 kg live weight (or 76% in relative terms) at weaning persisted in animals subsequently fed a high-quality regime over a 9-month fattening period while it was 40% compensated in comparable animals turned out to pasture. In the absence of a significant compensatory growth, the voluntary food intake (expressed per unit live weight°-75), and consequently feed conversion efficiency (daily weight gain per unit food intake), during the fattening period or during the first winter following a pasture season are little affected by previous milk restriction (Reardon and Everitt, 1972; Folman et al., 1974: Meadowcroft and Yule, 1976). In some cases feed conversion efficiency was increased but this effect did not exceed 10 to 30% (Bond et al., 1972; Tudor and O'Rourcke, 1980). The effect of a growth limitation during the milk-feeding period on final carcass composition again depends on the type of feeding following weaning. After one or several pasture seasons, the fat proportion in the carcass of previously restricted animals is lower than that of unrestricted animals at the age of 12 months ( - 3 points; Wardrop, 1966 ) and still at the age of 21 months ( - 0 . 5 point; Reardon and Everitt, 1972; Everitt and Jury, 1977). This is obviously a consequence of their lower live weight at slaughter since no difference was reported when animals were slaughtered at the same live weight (Bond et al., 1972; Tudor et al., 1980). But when concentrate-based diets are fed during the finishing period, the carcass of restricted animals generally contains more fat and less protein than that of control animals (+4.0, and - 1.3 to - 2 . 4 points respectively; Bond et al.. 1972; Meadowcroft and Yule, 1976; Tudor et al., 1980). 1.2. Late-weaned calves In suckled calves, the variations in milk allowance are generally smaller than those obtained in early-weaned calves. In addition, their effects are to

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some extent compensated by the substitution of milk by grass, the latter being consumed in greater quantities as milk intake decreases (Le D u e t al., 1976; Le Neindre and P6catte, 1987; Wright and Russel, 1987 ). This could explain why Le Neindre and P6catte ( 1987 ) and Stuedemann et al. ( 1968 ) found in Normand and Hereford suckled calves that body composition at weaning at an age of 6 to 8 months was little affected by milk restriction despite live weight differences of around 45 kg with control animals (less than 1 percentage point of variation in carcass fat depots). But when the growth rate of suckled calves was increased by concentrate supplementation ( + 10 to 15% ), carcass fat proportion was increased in one case ( + 3 percentage points; Stuedemann et al., 1968) and remained unchanged in another (Le Neindre and P6catte, 1987) after an intensive finishing period. In the 15- to 20-month young bulls and young steers systems, the pre-weaning treatment (milk allowance and concentrate level) has no practical consequence for the daily live weight gain during the subsequent fattening period (Stuedemann et al., 1968; Clutter and Nielsen, 1987; Le Neindre and.P6catte, 1987; Osoro and Wright, 1988; Lewis et al., 1989). In a 20- to 24-month beef system, the compensatory growth is also negligible during the first winter and until slaughter after a period of milk restriction (Drennan and Harte, 1970; Wright et al., 1987; Lewis et al., 1989; Lowry et al., 1989). Several authors, however, reported that high levels of feeding during the first winter tended to promote higher daily live weight gains during the following pasture season, thus limiting the effects of restriction during the milk phase (Brookes and Hodges, 1959; Keane and Drennan, 1978; Drennan, 1979; Drennan and Harte, 1979). The voluntary food intake and the food conversion efficiency during the first and second winters are little changed by a milk restriction to 8 months of age (variations of less than 10%) when the animals are fed a fattening regime (Stuedemann et al., 1968; Lewis et al., 1989) or a grass silage-based diet (Keane and Drennan, 1978; Drennan and Harte, 1979). Finally, the carcass composition of young bulls and 24-month steers at slaughter at a given live weight is practically independent of the previous level of milk feeding (Drennan, 1979; Le Neindre and P6catte, 1987; Lewis et al., 1989). But when slaughtered at the same age, the carcasses of steers previously restricted during calfhood weigh less and have a lower proportion of fat than those of continuously fed steers (Drennan and Harte, 1979; Lowry et al., 1989). 2. PERIOD OF WEANING

Weaning leads to important modifications in the internal anatomy and physiology of the calf due to a more or less abrupt change in the feeds con-

EFFECTS OF FEEDING DURING CALFHOOD ON PERFORMANCE IN BEEF CATTLE

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sumed, and marks the transition between the monogastric and the pre-ruminant stages. The substitution of liquid milk by solid feeds initiates the rumen cellulolytic activity and the absorption of the volatile fatty acids produced (Candeau, 1972). The rumen then changes both in morphology (weight and volume) and in histological composition (weights per surface unit ofmucosa and muscle fractions) (Warner et al., 1953, 1956; Harrison et al., 1960; Warner, 1961 ; Stobo et al., 1966a,b; Hodgson, 1971; Kaiser, 1976; Ohtani et al., 1976; Le Neindre, personal communication, 1986 ). The extent of these modifications, however, depends upon the type of solid feeds consumed (e.g. forage v. concentrates; Fig. 1 ). However, as suggested by Warner ( 1961 ), there is no evidence that the variations in rumen capacity and activity at the calf stage due to the type of feeds consumed at weaning are still perceptible when the animal Volume of the reticulo-rumen (I/kg LW) 0.8 ~

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is mature. The long-term effect of this factor on feed intake, live weight gain and body composition still needs to be investigated. At weaning the calf body mass suffers a drastic reduction in lipids as a result of important modifications in the energy metabolism. At the same time, the protein fraction is affected but to a much lesser extent. These effects are more marked as the calf's growth rate, and consequently its fat depots, during the period of milk feeding increase (Robelin and Chilliard, 1989). After weaning, the growth of weaned calves is generally greater than or equal to that of unweaned calves (Harvey et al., 1975; Kubisch and Makarechian, 1987) unless the animals are weaned very early (viz. less than 1 month of age; Butterfield et al., 1966). From the experiments carried out by several authors on Friesian, Salers and Jersey calves fed milk artificially, the age at weaning has practically no influence on the calf's subsequent feed intake capacity, feed conversion efficiency and growth capacity: in most cases a difference in live weight due to the age at weaning was not compensated later (Swart and Swart, 1965; Butterfield et al., 1966; Castle et al., 1967; Lonsdale and Taylor, 1969; Geay, 1970; Richardson et al., 1978; Le Neindre, personal communication, 1986). When a compensatory growth occurred, only 50 to 60% of the initial live weight differences were compensated after 6 to 12 months of fattening (Aitken et al., 1963; Harvey et al., 1975; Kubisch and Makarechian, 1987). However, observations made by Muller and P6catte (personal communication, 1987) indicate that suckled calves exhibit better performances the following year at pasture (lower weight loss at turnout, + 50 g mean daily live weight gain) when compared to early-weaned calves fed conserved forages and grown at a similar rate from weaning to turnout. On the other hand, the age at weaning has a marked short-term effect on body composition. Geay (1970) reported that, in Friesian calves growing at a similar rate, those fed milk at pasture and weaned at 8 months of age had a lower carcass fat proportion ( - 4 points or - 30%) at 9 months of age than those weaned at 3.5 months and fed concentrates up to 8 months of age. However, such differences must progressively vanish during the subsequent finishing period since animals weaned at different ages and slaughtered at the same age have practically the same carcass composition (Aitken et al., 1963; Geay, 1970; Harvey et al., 1975; Kubisch and Makarechian, 1987; Le Neindre, personal communication, 1987 ). 3. P O S T - W E A N I N G

PERIOD

3.1. Compensatory growth and food conversion efficiency 3. I. 1. Effect of growth level during the first winter After weaning, calves subjected to a food restriction during their first winter can generally compensate part of the growth delay during the subsequent

18 5

EFFECTS OF FEEDING DURING CALFHOOD ON PERFORMANCE IN BEEF CATTLE

pasture season. Their average daily live weight gain at grass is 35 to 70% higher than that of unrestricted control animals (Bohman, 1955; Horton and Holmes, 1978; Drennan and Harte, 1979; Baker et al., 1985; Keane et al., 1988; Lewis et al., 1988; Lowman et al., 1988 ). Though not so active as in older animals (see O'Donovan, 1984), the compensatory growth is sometimes still perceptible during the second winter (Owen and Ochoa, 1982; Micol, 1986) or not (Heinemann and Van Keuren, 1956; Meyer et al., 1965; Broadbent et al., 1969; Drennan et Harte, 1979; Lewis et al., 1988 ). Such a wide variation in the compensation rates reported in the literature is not apparently related to variations in the age of the animals at the start of the winter restriction period (Fig. 2 ). In the case of calves ranging in age from 3 months (autumn-born) to 10 months (spring-born), the effect of other factors may have predominated. The compensation rate at pasture and also during fattening tends to increase as the severity of winter restriction, and consequently the live weight difference at the end of winter, increases (Heineman and Van Keuren, 1956; Lawrence and Pearce, 1964a; Hironaka and Kozub, 1973; Saubidet and Verde, 1976; Drennan, 1979; Keane and Drennan, 1980; Williams and Macdearmid, 1983; Micol, 1986; Wright et al., 1986). As also reported in older animals of 18 months of age (Conway, 1967; O'Donovan et al., 1972 ), the compensation at pasture varies little in absolute terms with the Compensation rate at pasture (%) 220 200 180 160

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stocking rate ( + 2 0 0 g increase in daily live weight gain; Meyer et al., 1965; Baker et al., 1985). A compromise must then be found between the growth level during winter, the calf's compensatory growth capacity (which also depends on its ability to utilize grass or high-energy fattening diets) and its propensity to deposit fat. In his review on beef production from grassland, Baker (1975) stated that a moderate winter gain (300 to 600 g/day) for calves from suckler herds ensures maximum compensation at pasture and highest overall weight gains over first winter and second summer seasons. Micol (1986), however, suggested that the optimum live weight gains during the first winter for grass beef should be adapted to the breed maturity, i.e. 600 to 800 and 900 to 1000 g/day for calves of early-maturing type (Normand) and latematuring type (Charolais) respectively. The type of feeds subjected to restriction is also a factor of compensatory growth, especially when restriction takes place at an early stage of growth. Former works, recently confirmed by those of Owen and Ochoa ( 1982 ), Abdalla et al. ( 1988 ) and Drouillard et al. (1989), proposed that a growth limitation during calt'hood was better compensated when it was obtained through a food restriction on energy rather than on protein (Winchester et al., 1957 ). From the results obtained in animals slaughtered at 24 or 30 months of age one may not expect an overall compensation during summer of more than 30 to 50% of a live weight difference due to a protein restriction during the first winter, though in some cases an almost complete compensation was found and in others a negative compensation (Table 1). When nitrogen-supplemented and unsupplemented calves were grown at the same rate during winter, Wilkinson and Mackie (1988) obtained the same performances during the following pasture season. This contradiction between the experimental results highlights the importance of the interaction between winter and summer feeding in young cattle. Its origin is still unexplained but probably lies in the modifications induced in the composition of body gain (Beever and Baker, 1986). The voluntary feed intake (expressed per unit live weight °-75) is always greater after a growth limitation suffered during the first winter. The difference is generally more marked as the degree of restriction increases. It is maximum during the first weeks of realimentation (up to 50%) and then decreases more or less rapidly. A good description of these variations is given by the works of Hironaka and Kozub (1973), Saubidet and Verde (1976), and Williams and Macdearmid ( 1987 ) in calves fed concentrate-based diets (50 to 80% cereals) during recuperation (Fig. 3 ). At pasture one may expect a similar variation of grass intake despite the greater imprecision of its measurement by indirect methods. After restricting 1-year-old calves during winter, Horton and Holmes (1978) registered a 50 and 7% increase in grass intake at pasture respectively 1 and 4 months after turnout. From the data given by Meyer et al. (196-5), Baker et al. ( 1 9 8 5 ) a n d Wright et al. (1986, 1987)

EFFECTS OF FEEDING DURING CALFHOOD ON PERFORMANCE IN BEEF CATTLE

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TABLE 1 Effect of a restriction on food protein supply to calves during the first winter period on the subsequent live weight compensation during the following summer Authors and type of calves a

BohmanandTorell(1956) nm, HE KeaneandDrennan(1980) S, nm Hennessy et al. (1981) S, HE Kirby and Chalmers ( 1981 ) S and H, FR and x H E Kirby and Chalmers (1982) S, FR Baker and Gibb (1982) S, FR Kirby and Chalmers (1983) S, FR Mackie el al. (1983) B, FR Sleen (1986) S, FR Chapple (1987) H, nm Abdalla et al. (1988) S, FR

Winter restriction period

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Initial Duration Final LW difference LW (months) control-restricted ( kg ) (kg) (%)

Duration Compensation of (months) the LW difference

Compensation rateb

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(%)

193

4

8

4

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15

188

1.27

242

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4 13 30

3

170

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3 18 30

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1.03 1.15 1.24

265 208 183

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48 121 75

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aBull calf (B), heifer calf (H) or steer calf (S); Friesian (FR), Hereford (HE), Hereford-cross ( x H E ) . hExpressed as daily live weight gain (DLWG) of previously restricted calves/DLWG of control calves. nm: not'mentioned.

the mean increase in grass intake over the complete pasture season can be estimated to be approximately 10 to 20%. The feed conversion efficiency is also markedly increased after a winter restriction when calves are fed liberally a concentrate-based diet (up to 40%; Hironaka and Kozub, 1973; Saubidet and Verde, 1976), or turned out to pasture (from 30 to 90%; Meyer et al., 1965; Baker et al., 1985; Wright et al., 1986 ). During the second winter this advantage may still be perceptible ( + 15 to 20%; Micol, 1986 ) or not (Meyer et al., 1965; Keane and Drennan, 1978 ).

3.1.2. Effect of growth level during the first pasture season The effects of early growth limitation have been less investigated in the case of a restriction applied during the grazing season than during the winter indoor feeding period. Actually young calves are seldom turned out to pasture

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Fig. 3. Effect of a food restriction from 8 to 12 months of age on the subsequent voluntary food intake in steer calves (control: unrestricted calves; moderate and high restrictions: - 1 7 and - 4 4 % live weight difference from controls at the end of restriction respectively; data adapted from Saubidet and Verde, 1976).

before 5-6 months of age. When growth is limited at pasture, a compensation generally takes place during the following winter. But compensatory growth is less active than that exhibited at pasture by calves previously restricted in winter (Moran and Holmes, 1978 ), probably because winter restrictions are generally more severe than those imposed at pasture by varying the stocking rate and the level of concentrate supplementation (O'Donovan et al., 1972). The nature of the restriction again affects the calf's compensation capacity. Denham ( 1977 ) compared the respective effects of energy and protein supplementations in grazing Hereford steer calves. The two types of supplement had a positive effect on the growth rate of calves at pasture but had opposite effects on weight gain during the subsequent winter fattening period, negative in the case of the energy supplement and positive in the case of the protein supplement. The compensation during winter is also more active when the diet energy concentration - - or the concentrate proportion - - increases (Fox et al., 1972; Verde et al., 1974; Moran and Holmes, 1978). This could explain why a partial compensation can be observed in calves fed cereal- or maize silage-based diets (Perry et al., 1971; Denham, 1977; Moran and Holmes, 1978; Giovanni, 1982; Keane and Drennan, 1983 ) and practically no compensation when

EFFE{ 'TS OF FEEDING D U R I N G CALFHOOD ON PERFORMANCE IN BEEF CATTLE

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they are fed grass silage-based diets (Wilkinson and Prescott, 1970; Drennan et al., 1982). During the recuperation period the voluntary feed intake (expressed per unit live weight °75) and the feed conversion efficiency of previously restricted calves are greater than those of their counterparts continuously fed. Differences of 10 to 15% in feed intake and 11% in feed conversion efficiency were reported by Keane and Drennan (1983) and Denham (1977) respectively. These values are much lower than those observed in calves restricted during their first winter (see above) but they need to be confirmed.

3.2. Body composition and composition of body gain 3.2.1. At the end of the differential growth period Though many works have been carried out with beef cattle on the effect of feed restriction on subsequent performance, little information is available on body composition of young calves after a period of growth limitation. The adipose tissues are by far the most sensitive to feeding variations though they only represent a relatively small proportion of the calf's body mass (less than 10% - - against 40% for muscles - - in Friesian and Charolais calves of up to 250 and 350 kg empty body weight respectively; Robelin, 1986a). Calves whose growth was limited during their first year, and weighing 20 to 30% less than control animals, present evident signs of a lower adiposity at the end of winter at 15 months of age: - 2 to - 7 percentage points (or - 10 to - 5 0 % ) in carcass lipids (Meyer et al., 1965; Hancock et al., 1987, 1988 ), - 90% in fat cover depth over the 10th rib (Morgan and Everitt, 1968 ), - 40% in cross-sectional subcutaneous fat area (Wright et al., 1986). On the other hand, their carcasses contain higher proportions of protein ( + 1.5 point or + 8 % ) , saleable meat ( + 5 . 5 points or + 9 % ) , and bone ( + 6 . 6 points or + 29% ). From these results it is difficult to understand why Baker et al. ( 1985 ) could observe the opposite effect in 7-month-old Friesian steer calves restricted to 65% of the live weight of control calves from the age of 4 months on grass silage-based diet, that is to say a greater proportion of lipids ( + 0.9 point) and a lower proportion of protein ( - 0 . 7 point) in the body mass.

3.2.2. At slaughter after a recuperation period When the animals are slaughtered at the same age, those whose growth rate was limited during calfhood generally have lower body weight and carcass weight than their unrestricted counterparts. Their carcass also contains less fat at 24 months of age after a winter finishing period (Steen, 1986) and even at 12 or 24 months of age after finishing at pasture (Morgan and Everitt, 1968; Baker et al., 1985). In the latter case, the difference is greater as the daily live weight gain increases or the stocking rate decreases during the pasture season (Meyer et al., 1965 ). Some other workers, however, did not find

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any difference in the carcass composition of steers due to an early restriction from 3 to 7 months of age and slaughtered at 16 or 24 months, though differ ences in carcass weight reached 7 to 21 kg respectively (Broadbent et al., 1969; Keane and Drennan, 1983 ). When a similar slaughter weight is achieved, the previously restricted animals have the same body composition or are a little leaner than unrestricted animals (Lawrence and Pearce, 1964b; Micol, 1986; Carstens et al., 1988). In the pi-oduction of 19- to 24-month Charolais crossbred steers, Wright et al. (1986) pointed out that those animals whose growth was limited during the first winter were not finished at the end of the second grazing season and required a further winter period of intensive feeding to reach the same level of fatness as that of control steers slaughtered at 19 months. Their carcass weight at 24 months then was 9 to 13% greater, which confirms that at the same age of 19 months their live weight and their carcass fat proportion were lower than those of the unrestricted steers. A better understanding of these differences in body composition due to an early growth limitation is given by the results obtained by Rompala et al. ( 1985 ) in steer calves restricted from 7 to 9 months of age and refed until the age of 19 months. At the start of the realimentation period, the growth of body lipids in calves previously restricted was negative or nil and caused a temporary reduction of their adiposity. At the same time, control calves of similar empty body weight (EBW) and achieving the same weight gain were depositing fat at an increasing rate. Further on, the fat proportion in the weight gain of previously restricted calves increased very rapidly with live weight and exceeded that observed in control calves. This effect was detected earlier as weight gain increased. Finally the initial difference in body lipids at 250 kg EBW ( - 6 points ) was pra~ically resorbed at 550 kg EBW ( - 0 . 5 point ). On the other hand, the end of restriction was immediately followed by an increase in the proportion of protein in the body gain of previously restricted calves when compared to control calves. But the difference varied with the level of daily live weight gain. At the lowest level of gain the difference was greatest at the start of the realimentation period ( + 4 points) at 200-250 kg EBW and then was more or less maintained to 500 kg EBW, while at the highest level of gain it was almost negligible throughout the recuperation period ( + 0.4 point). These results agree well with those of Waldman et al. ( 1971 ) and Fox et al. ( 1972 ), obtained in Friesian and Hereford steer calves. These authors found that differences in carcass composition caused by a growth limitation from birth to 90 kg live weight or by a 6 months restriction from 230 kg EBW were no longer perceptible at 350-450 kg live weight. The reason for this compensation is that the initial difference in the composition of body gain during recuperation (more protein and less lipids) is progressively reduced and even reversed at the end of the recuperation period (Fox et al., 1972; Carstens et

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191

al., 1988). Confirmation was provided by Morgan and Everitt (1968) who found that at slaughter at 22 months of age the weight of carcass components (edible meat, excess fat, bone) of Jersey steers restricted or not from 3 to 15 months of age was highly related to the age and the carcass weight at slaughter but was independent of their nutritional past. 4. D I S C U S S I O N A N D C O N C L U S I O N

4.1. Growth From the results presented in this review, it may be stated that calves have only a limited capacity to compensate for growth delays. The rate of compensatory growth tends to increase with the severity of restriction. However, this trend was not demonstrated in the case of very early food restrictions, i.e. during the milk feeding phase. Though it is not possible from the available literature to give either a precise description of the effect of age on compensatory growth (see Fig. 2), or a limit of age under which the calf's compensation capacity is definitively altered, there is evidence that compensation is lower in calves than in cattle restricted at a more advanced age, as previously suggested by Winchester and Ellis (1956), Lawrence and Pearce (1964a), Levy et al. ( 1971 ) and Morgan (1972). The growth delays imposed before weaning are actually the least likely to be subsequently compensated. Similarly, the consequences for growth of a restriction of feed protein supply during the first months of life are generally irreversible. This may be the case during the first winter when calves are fed grass silage-based diets or during summer at the end of the pasture season. The overall effects of age and severity of restriction are summarized in Fig. 4 from a wide set of data from the literature, obtained with beef cattle ranging in age from 0 to 25 months. The animals were divided into two populations according to their age at the start of the restriction period (less and more than 6 months) regardless of the type of cattle and the type of feeding. The figure shows clear interaction between these two factors. On average, weight compensation in calves restricted during calfhood is low and practically independent of the severity of restriction, while in cattle restricted at a later stage of development it is greater and increases with the severity of restriction. Compensatory growth, however, should not be considered negligible in young cattle but in most cases a significant partial compensation of a growth delay will be obtained at the cost of a long period of recuperation. A demonstration is given by comparing the calves studied by Harte ( 1968 ), Bond et al. ( 1972 ) and Keane and Drennan (1983) restricted at less than 7 months of age with those of McCarrick et al. (1963) and Lawrence and Pearce (1964a) restricted at ages ranging from 10 to 22 months. The former required 14 to 18 months to compensate 70 to 80% of their growth delay while the latter required only 4 to 7 months for the same result.

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Compensation rate (%) 220

Age at the start of the restriction period

200

• Lessthan 6 months

<> O

180

O Morethan6 months

O OO


160 .

140 OO 0

~ ¢~

O©<> OO

© •

•

120

t

•

100 8O 6O

L

0

10

20

~

~

i

30

40

50

_ _

k

60

_

.

i

J

70

80

Severity of the restriction (%) Fig. 4. Interaction between the severity of food restriction and the age at the start of the restriction period on the subsequent compensatory growth in cattle (severity of food restriction ex-

pressed as the percent live weight difference between restricted and control animals at the end of the restriction period; compensation rate expressed as the ratio of the daily live weight gain of previously restricted calves on the daily live weight gain of control calves × 100; data from 74 experiments, each of the 177 points corresponding to a restricted group mean value).

Fig. 4 also shows the great variability in the response of cattle to live weight gain limitation at all ages. This reflects the diversity of the experimental conditions, and more generally the high number of factors involved in compensatory growth.

4.2. Voluntary food intake The increase in voluntary food intake after a period of restriction is certainly one of the main factors determining the compensatory growth in cattle. It demonstrates that the growth of the different constituents of the digestive tract, in particular that of the stomachs, is little affected in absolute terms by the variations in level of feeding (Saubidet and Verde, 1976; Le Neindre and P6catte, 1987 ). During the realimentation period, those calves whose growth rate was limited have a lower live weight and consequently have lower maintenance requirements than calves fed normally. A greater fraction of the metabolizable energy intake then is available for growth (Saubidet and Verde,

1976). The effect of an early growth limitation on subsequent efficiency of converting food energy in body tissues, however, is still controversial. Carstens

EFFECTS OF FEEDING DURING CALFHOOD ON PERFORMANCE IN BEEF CATTLE

| 93

et al. (1987) pointed out that the kr (efficiency of utilization of metabolizable energy above maintenance ) reaches high values during early realimentation and then decreases with time. In steers fed the same level of energy per unit live weight after a period of food restriction, Thomson et al. ( 1982 ) actually observed a compensatory growth, thus indicating that the animals utilized food energy more efficiently than did their non-restricted counterparts. On the contrary, under the same conditions of energy supply, Robelin and Chilliard ( 1989 ) did not observe any compensatory growth in young Friesian bulls after a milk restriction from birth to 3 months of age. The difference in calf age during the restriction period could explain the discrepancy between the results of these two experiments. But the respective contribution of voluntary food intake and efficiency of food energy utilization in the expression of compensatory growth is still unknown.

4.3. Body and carcass composition The most remarkable short-term response of body anatomical composition of cattle to a growth limitation during calfhood is the reduction in fat deposition rate and, to a lesser extent, the increase in muscular growth. These variations are even observed in calves restricted just after birth. Their implication in the subsequent increase in the voluntary food intake is certainly of major importance (Wright et al., 1986). The modifications in the relative growth of the various constituents of the digestive tract caused by a food restriction should also be considered but they have been little investigated. Though the reduction in the calf's adiposity is spectacular, it generally does not persist long after the end of the restriction period whatever the calf's age at the start of the restriction. The differences in body composition from unrestricted control calves rapidly vanish as soon as the restricted calves are allowed to exhibit compensatory growth. Thus it is quite improbable that these differences are maintained throughout the recuperation period until slaughter at a given commercial carcass weight. However, one may expect the systems of beef production from grass to be somewhat better adapted than intensive systems when the objective is to limit the development of adipose tissues after a period of restriction during calfhood. It can be concluded from this review that the main lines of the long-term effects of a growth manipulation during the first year of the calf's life are well known but many points still need to be investigated if a wide practical application is to be given in the situations now found in the various systems of beef production in Europe. At this stage it can be suggested that in the future priority should be given to the study of the following factors: - - the breed effect has not been clearly demonstrated. A few attempts were made to compare the main Anglo-Saxon breeds (Angus, Friesian, Holstein, Jersey, Shorthorn, Hereford; Brookes and Hodges, 1959; Bond et al., 1972;

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Meadowcroft and Yule, 1976). These early-maturing breeds have not yet been compared to the late-maturing continental breeds such as Charolais and Limousin. But such a comparison must take into account the possible interaction of this factor with the rearing method which differs between breeds (early weaning in dairy breeds, late weaning in suckler herds ). - - the method of weaning is certainly a factor that has been little studied. The effect of age at weaning should be evaluated with calves growing at similar rates between the weaning dates, as well as the long-term effect of type of feeds during the weaning phase. - - the effect of type of feeds on the relative growth of the constituents of the digestive tract is well known, but not that of a variation of growth rate during calfhood on this parameter. A better knowledge of the latter would be helpful in understanding the relationship between the growth level and the subsequent voluntary feed intake. - - the effects of sex and castration are still undetermined. One comparison only was undertaken by Tudor and O'Rourcke (1980) between male and female calves, and it showed no difference in performance due to the sex of the animals. the measurements of the rate of compensation seldom took into account the variations in the weight of the digestive tract content. As pointed out by Lawrence and Pearce (1964a) and Carstens et al. (1988), these variations can in fact explain part of the live weight variations in cattle, and particularly during the major alimentary transitions (i.e. weaning, turnout to pasture, housing in autumn ). Little attention has been paid in the past to these critical periods which also could account for part of the variation observed in the animal performance in response to a growth delay imposed during calfhood. -

-

REFERENCES Abdalla, H.O., Fox, D.G. and Thonney, M.L., 1988. Compensatory gain by Holstein calves after underfeeding protein. J. Anim. Sci., 66: 2687-2695. Aitken, J,N., Preston, T.R., Whitelaw, F.G., Macdearmid, A. and Charleson, E., 1963. Intensive beef production. 2. The effect of three, twelve or sixteen week weening on the performance of Aberdeen-Angus crossbred cattle. Anita. Prod., 5: 53-56. Baker, R.D., 1975. Grassland systems for beef production from dairy bred and beef calves. Livest. Prod. Sci., 2: 121-136. Baker, R,D. and Gibb, M.J., 1982. Compensatory growth studies. Grassland Research Institute, Hurley, Annual Report 1982, pp. 104-105. Baker, R.D., Young, N.E. and Laws, J.A,, 1985. Changes in the body composition of cattle exhibiting compensatory growth and the modifying effects of grazing management. Anita. Prod., 41: 309-321. Beever, D.E. and Baker, R.D., 1986. Improving the utilization of forage for lean beef production. Agricultural and Grassland Research Institute, Hurley, Annual Report, pp. 102-105. Bohman, V.R., 1955. Compensatory growth of beef cattle: the effect of hay maturity. J. Anim. Sci., 14: 249-255.

EFFECTS OF FEEDING DURING CALFHOOD ON PERFORMANCE IN BEEF CATTLE

195

Bohman, V.R. and Torell, C., 1956. Compensatory growth of beef cattle: the effect of protein supplements. J. Anim. Sci., 15: 1089-1096. Bond, J., Hooven, N.W., Warwick, E.J., Hiner, R.L. and Richardson, G.V., 1972. Influence of breed and plane of nutrition on performance of dairy, dual-purpose and beef steers. II. From 180 days of age to slaughter. J. Anim. Sci., 34:1046-1053. Broadbent, P.J., Ball, C. and Dodsworth, T.L., 1969. The effect of plane of nutrition during calfhood on the subsequent performance of Hereford × Ayrshire steers. Anita. Prod.. 11:155160. Brookes, A.J. and Hodges, J., 1959. Studies in beef production. I. The effect of level of feeding and of breed on the growth and fattening of spring born cattle. J. Agric. Sci. Camb., 53: 78. Butterfield, R.M., Pryor, W.J. and Berg, R.T., 1966. A study of carcase growth in cattle. Res. Vet. Sci., 7: 417-423. Butterfield, R.M., Johnson, E.R. and Pryor, W.J., 1971. A study of growth in calves. I. Carcass tissues. J. Agric. Sci. Camb., 76: 453-458. Candeau, M., 1972. Stimulation physico-chimique et d6veloppement du rumen. Thbse de Doctorat d'Etat, Univ. Paris VI, 223 pp. Carstens, G.E., Johnson, D.E. and Ellenberger, M.A., 1987. The energetics of compensatory growth in beef cattle. J. Anita. Sci., 65 (Supplement 1 ): 263-264. Carstens, G.E., Johnson, D.E., Ellenberger, M.A. and Tatum, J.D., 1988. Composition of carcass and non-carcass tissues in beef steers exhibiting normal and compensatory growth. J. Anita. Sci., 66 (Supplement 1 ): 491-492. Castle, M.E., Campbell, A.G. and Clayton, D.G., 1967. Rearing Jersey calves on concentrates. Proc. Ruakura Farmer's Conf. Week, pp. 181-185. Chapple, D.G., 1987. Patterns of winter feeding for spring-born suckled calves in relation to their subsequent performance at grass. Anim. Prod., 44:474 (abstr.). Clutter, A.C. and Nielsen, M.K., 1987. Effect of level of beef cow milk production on pre- and postweaning calf growth. J. Anim. Sci., 64:1313-1322. Conway, A., 1967. Compensatory growth of beef cattle under different stocking rates. An Foras Tahintais, Dublin, Research Report 1967, p. 11. Denham, A.H., 1977. Influence of energy and protein supplements on grazing and feedlot performance of steers. J. Anita. Sci., 45: 1-7. Drennan, M.J., 1979. Compensatory growth in cattle. 1. Influence of feeding level during the first winter (9 to 14 months of age) on subsequent performance and carcass composition. Ir. J. Agric. Res., 18: 131-143. Drennan, M.J. and Harte, F.J., 1970. Effect of plane of nutrition in early life on subsequent performance of beef cattle. An Foras Taltintais, Dublin, Animal Production, Research Report 1970, p. 10. Drennan, M.J. and Harte, F.J., 1979. Compensatory growth in cattle. 2. Influence of growth rate in the calf stage (birth to 8 months) and during the first winter (8 to 13 months) on subsequent performance and carcass composition. Ir. J. Agric. Res., 18:145-156. Drennan, M.J., Conway, A. and O'Donovan, R., 1982. Compensatory growth in cattle. 3. Effect of stocking rate at pasture in summer on animal performance at pasture and during the subsequent winter period. Ir. J. Agric. Res., 21 : 1-11. Drouillard, J., Ferrell, C. and Klopfenstein, T., 1989. Compensatory growth following protein or energy restriction. Beef Cattle Report 1989. The Agricultural Research Division, I.A.N.R., Univ. Nebraska, Lincoln, pp. 35-38. Everitt, G.C., 1972. Calf growth and lifetime performance of beef cattle. N.Z. Soc. Anita. Prod., 32: 20-25. Everitt, G.C. and Jury, K.E., 1977. Growth of cattle in relation to nutrition in early life. N.Z.J. Agric. Res., 20: 129-137. Folman, Y., 1977. A note on the relationship between liveweight gain during calfhood and sub-

196

P. BERGE

sequent performance of intact male calves of the Israeli-Friesian breed. Anita. Prod., 24: 283-286. Folman, Y., Drori, D., Holzer, Z. and Levy, D.. 1974. Compensatory growth in intensively raised bull calves. J. Anita. Sci., 39: 788-795. Fox, D.G., Johnson, R,R., Preston, R i . , Dockerty, T.R. and Klosterman, E.W., 19"72. Protein and energy utilization during compensatory growth in beef cattle. J. Anita. Sci,, 34:310318. Geay, Y., 1970. Adaptation du mode d'elevage et d'alimentation au type g6n6tique. In: S.E.I.C.N.R.A. (Editors), La Production de Viande par les Jeunes Bovins. Etude no. 46. pp. 95109. Geay, Y. and Robelin, J., 1979. Variation of meat production capacity in cattle due to genotype and level of feeding: genotype-nutrition interaction. Livest. Prod. Sci., 6: 263-276. Gcay, Y.. Robelin, J. and Beranger, C., 1976. Influence du niveau atimentaire sur le poids vifet la composition de la carcasse de taurillons de diff6rentes races. Ann. Zootech., 25: 287-298. Giovanni, R., 1982. Mise a l'herbe aprbs sevrage des veaux laitiers n6s en fin d'hiver. 1. Influence du niveau d'alimentation avant la mise a l'herbe et de la compl6mentation au p~turage. Bull. Tech. C.R.Z.V. Theix, I.N.R.A., 50: 33-45. Hancock, D.L., Williams, J.E., Hedrick, H.B., Beaver, E.E., Larrick, D.K., Ellersieck, M.R., Garner, G.B.. Morrow. R.E., Paterson, J.A. and Gerrish, J.R., 1987. Performance, body composition and carcass characteristics of finishing steers as influenced by previous forage system. J. Anim. Sci., 65:1381-1391, Hancock, D.L., Williams, J.E., Hedrick, H.B.. Beaver, E.E., Hannah, S.M., Miller, S.J., Ellersleek, M.R,, Garner, G.B., Morrow, R.E. and Gerrish, J.R., 1988. Effect of previous forage system on feedlot performance, body composition and plasma variables of finishing steers. J. Anim. Sci., 66: 2272-2283. Harrison, H.N., Warner, R.G., Sander, E.G. and Loosli, J.K., 1960. Changes in the tissue and volume of the stomachs of calves following the removal of dry feed or consumption of inert bulk. J. Dairy Sci., 43: 1301-1312. Harte, F.J., 1968. Effects of plane of nutrition on calves for beef production. I. Growth rate, feed conversion efficiency, carcass yield and offals. It. J. Agric. Res., 7:137-148. Harte, F.J. and Fallon, R.J., 1982. Effect of plane of nutrition in the period 0-8 weeks on subsequent calf performance. An Foras Talt~ntais, Dublin, Animal Production, Research Report 1982, p. 12. Havre, F.J. and Fallon, R.J., 1983. Effect of plane of nutrition in calfhood on subsequent performance. An Foras Taluntais, Dublin, Animal Production, Research Report 1983, p. 7. Havre, F.J. and Fallon, R.J., 1984. Effect of level of feeding in the calf stage on the life time performance of Friesian steers. Proc. Br. Soc. Anim. Prod., 38:533 (abstr.). Harte, F.J., Fallon, R.J. and Drennan, M.J., 1981. Effect of plane of nutrition in the period 0-8 weeks on subsequent calf performance. An Foras Talfintais, Dublin, Research Report 1981, p. 15. Harvey, R.W., Burns, J.C., Blumer, T.N. and Linnerud, A.C., 1975. Influence of early weaning on calf and pasture productivity. J. Anim. Sci,, 41: 740-746. Heinemann, W.W. and Van Keuren, R.W., 1956. The effect of wintering plane of nutrition on subsequent gains of beef yearling steers on irrigated pastures. J. Anim. Sci., 15:1097-1102. Hennessy, D.W., Williamson, P.J., Lowe, R.F. and Baigent, D.R., 1981. The role of protein supplements in nutrition of young grazing cattle and their subsequent productivity. J. Agric. Sci. Camb., 96: 205-212. Hironaka, R. and Kozub, G.C., 1973. Compensatory growth of beef cattle restricted at two energy levels for two periods. Can. J. Anim. Sci., 53:709-715. Hodgson, J., 1971. The development of solid food intake in calves. 3. The relation between solid food intake and the development of the alimentary tract. Anita. Prod., 13: 449-460.

EFFECTS OF FEEDING DUR|NG CALFHOOD ON PERFORMANCE IN BEEF CATTLE

197

Hooven, N.W., Jr., Bond, J., Warwick, E.J., Hiner, R.L. and Richardson, G.V., 1972. Influence of breed and plane of nutrition on the performance of dairy, daily-purpose and beef steers. I. Birth to 180 days of age. J. Anita. Sci., 34: 1037-1045. Horton, G.M.J. and Holmes, W., 1978. Compensatory growth by beef cattle at grassland or on an alfalfa-based diet. J. Anita. Sci., 46: 297-303. Kaiser, A.G., 1976. The effect of milk feeding on the pre- and post-weaning growth of calves, and on stomach development at weaning. J. Agric. Sci. Camb., 87: 357-363. Keane, M.G. and Drennan, M.J., 1978. Effect of level of feeding in winter on performance of weanlings previously reared on a high or moderate plane of nutrition. An Foras Taltintais, Dublin, Animal Production, Research Report 1978, p. 26. Keane, M G . and Drennan, M.J., 1980. Response of weanlings to low level of protein supplementation. An Foras Talfintais, Dublin, Animal Production, Research Report 1980, p. 13. Keane, M.G. and Drennan, M.J., 1983. Supplementation of grassfed calves. 3. Effects of concentrate level and protein content, and subsequent performance to slaughter. It. J. Agric. Res., 22: 113-125. Keane, M.G., Drennan, M.J. and Sherington, J., 1988. Effects of nutrition and anabolic agents in the growing and finishing phases on growth rate and carcass weight of Friesian steers. Ir. J. Agric. Res., 25:11-21. Kirby, P.S. and Chalmers, A.J., 1981. Effect of protein supplementation of silage diets on performance of growing cattle. Proc. Br. Soc. Anita. Prod., 32:387-388 (abstr.). Kirbx, P.S. and Chalmers, A.J., 1982. A comparison of extracted soya bean meal and fish meal as protein supplements for growing beef cattle given grass silage ad libitum. Proc. Br. Soc. Anim. Prod., 34:386 (abstr.). Kirby, P.S. and Chalmers, A.J., 1983. A comparison of formaldehyde-treated soya bean meal and two types of fish meal as protein supplements for growing beef cattle given grass silage ad libitum. Proc. Br. Soc. Anim. Prod., 36:538 (abstr.). Kubisch, H.M. and Makarechian, M,, 1987. Effect of date of weaning on post-weaning performance of bull calves of three breed groups. Can. J. Anita. Sci., 67: 941-949. Lawrence, T.L.J. and Pearce, J., 1964a. Some effects of wintering yearling beef cattle on different planes of nutrition. I. Liveweight gain, feed consumption and body measurement changes during the winter period and subsequent grazing period. J. Agric. Sci. Camb., 63:5-21. Lawrence, T.L.J. and Pearce, J., 1964b. Some effects of wintering yearling beef cattle on different planes of nutrition. It. Slaughter data and carcass evaluation. J. Agric. Sci. Camb.. 63: 23-24. Le Du, Y i . P . , Baker, R.D. and Barker, J.M., 1976. Milk-fed calves. 2. The effect of length of milk feeding period and milk intake upon herbage intake and performance of grazing calves. J. Agric. Sci. Camb., 87: 197-204. Le Ncindre, P. and Pecatte, G., 1987. Consequences des quantites de lait et d'aliment concentr¢5 consomm6es avant sevrage sur les performances de bovins mfiles entiers de 3 mois/t l'abattage. Ann. Zootech., 36: 387-410. Levy, D., Folman, Y., Holzer, Z. and Drori, D., 1971. Compensatory growth in intensively raised bull calves. J. Anim. Sci., 33: 1078-1085. Lewis, J.M., Klopfenstein, T.J., Stock, R.A. and Nielsen, M.K., 1988. Effect of wintering and growing systems on cattle finishing performance. J. Anita. Sci., 66 (Supplement 1 ): 479480 (abstr.). Lewis, J.M., Klopfenstein, T.J., Nielsen, M.K., Stock, R.A. and Hunt, C., 1989. Forage versus grain finishing systems and the fate of increased weaning weight due to an increased level of milk. Beef Cattle Report 1989. The Agricultural Research Division, I.A.N.R., Univ. Nebraska-Lincoln, pp. 29-31. Lonsdale, C.R. and Tayler, J.C., 1969. The artificial rearing of calves and their growth on grass

198

P. BERGE

diets. II. The effect of length of period of feeding cold milk substitute to spring-born calves at pasture. J. Agric. Sci. Camb., 73: 483-488. Lowman, B.G., Hinks, C.E., Swift, G., Neilson, D.R. and Hunter, E.A., 1988. Factors affecting the performance and carcass characteristics of finishing grazed cattle within a 20-month beef system. Anim. Prod., 46:500 (abstr.). Lowry, S.R., Woods, B.L. and Bradley, N.W., 1989. Compensatory growth and development during different phases of growing cattle. J. Anita. Sci., 67 (Supplement 2): 28. McCarrick, R.B., Harrington, D. and Conway, A., 1963. The effect of energy restriction between 20-24 months of age on subsequent growth and carcass development of beef cattle. Ir. J. Agric. Res., 2: 131-147. Mackie, C.K., King, M.E. and Kay, M., 1983. A comparison of foods for Friesian bulls slaughtered at 15 to 16 months old. Anim. Prod., 36:540-541 (abstr.). Meadowcroft, S.C. and Yule, A.H., 1976. A comparison of two planes of nutrition for beef calves. Exp. Husb., 31: 24-32. Meyer, J.H., Hull, J.L., Weitkamp, W.H. and Bonilla, S., 1965. Compensatory growth responses of fattening steers following various low energy intake regimes on hay or irrigated pasture. J. Anim. Sci., 24: 29-37. Micol, D., 1986. Production de viande de boeufs et de jeunes taureaux. In: D. Micol (Editor), Production de Viande Bovine. I.N.R.A. Publications, Paris, pp. 169-200. Moran, J.B. and Holmes, W., 1978. The application of compensatory growth in grass/cereal beef production systems in the United Kingdom. World Rev. Anita. Prod., 14: 65-73. Morgan, J.H.L., 1972. Effect of plane of nutrition in early life on subsequent live-weight gain, carcass and muscle characteristics and eating quality of meat in cattle. J. Agric. Sci. Camb., 78: 417-423. Morgan, J.H.L. and Everitt, G.C., 1968. Beef production from Jersey steers grazed in three environments. N.Z. Soc. Anim. Prod., 28:158-176. O'Donovan, P.B., 1984. Compensatory gain in cattle and sheep. Nutr. Abst. Rev. Set. B, 54: 389-410. O'Donovan, P.B., Conway, A. and O'Shea, J., 1972. A study of the herbage intake and efficiency of feed utilization of grazing cattle previously fed two winter planes of nutrition. J. Agric. Sci. Camb., 78: 87-95. Ohtani, S., Asahida, Y. and Hirose, Y., 1976. The effect of grazing on the development of the alimentary tract in the young calf. Jpn. J. Zootech. Sci., 47:175-180. Osoro, K. and Wright, I.A., 1988. Post-weaning compensatory growth in autumn-born suckled calves. Anim. Prod., 46:524 (abstr.). Owen, E. and Ochoa, C., 1982. Energy and protein supplementation of grass silage for Friesian steers: effect on performance from 4 to 7 months of age and on subsequent compensatory growth. Proc. Br. Soc. Anim. Prod., 34:388 (abstr.). Perry, T.W., Huber, D.A., Mort, G.O. and Rhykerd, C i . , 1971. Effect of level of pasture supplementation on pasture drylot and total performance of beef cattle. I. Spring pasture. J. Anim. Sci., 32: 744-748. Price, M.A., Butson, S. and Makarechian, M., 1984. The influence of feed energy level on growth and carcass traits in bulls of two breed types. Can. J. Anim. Sci., 64: 323-332. Reardon, T.F. and Everitt, G.C., 1972. Effect of pre-weaning nutrition on subsequent growth rate, feed conversion efficiency and carcass composition of identical twin steers. N.Z. Soc. Anim. Prod., 32: 26-38. Richardson, A.T., Martin, T.G. and Hunsley, S.E., 1978. Weaning age of Angus heifer calves as a factor influencing calf and cow performance. J. Anim. Sci., 47: 6-14. Robelin, J., 1979. Influence de la vitesse de croissance sur la composition du gain de poids des bovins: variations selon la race et le sexe. Ann. Zootech., 28:209-218. Robelin, J., 1986a. Composition corporelle des bovins: 6volution au cours du ddveloppement

EFFECTS OF FEEDING DURING CALFHOOD ON PERFORMANCE IN BEEF CATTLE

199

el diff6rences entre races. Thbse Doctorat d'Etat, Universit6 de Clermont-Ferrand II, 391 PP. Robelin, J., 1986b. Bases physiologiques de la production de viande: croissance et d6veloppemerit des bovins. In: D. Micol (Editor), Production de Viande Bovine. I.N.R.A. Publicalions, Paris, pp. 169-200. Robelin, J. and Chilliard, Y., 1989. Short-term and long-term effects of early nutritional deprivation on adipose tissue growth and metabolism in calves. J. Dairy Sci., 72:505-513. Rompala, R.E., ,Jones, S.D.M., Buchanan-Smith, J.G. and Bayley, H.S.. 1985. Feedlot performance and composition of gain in late-maturing steers exhibiting normal and compensatory growth. J. Anim. Sci., 61: 637-646. Saubidet, C.L. and Verde, L.S., 1976. Relationship between live weight, age and dry-matter intake for beef cattle after different levels of food restriction. Anim. Prod., 22: 61-69. Steen, R.W., 1986. The effect of plane of nutrition and type of diet offered to yearling Friesian steers during a winter store period on subsequent performance. Anita. Prod., 42: 29-37. Stobo, I.J.F., Roy, J.H.B. and Gaston, H.J., 1966a. Rumen development in the calf. 1. The effect of diets containing different proportions of concentrates to hay on rumen development. Br. J. Nutr.. 20: 171-188. Stobo, I.J.F., Roy, J.H.B. and Gaston, H.J., 1966b. Rumen development in the calf. 2. The effect of diets containing different proportions of concentrates to hay on digestive efficiency. Br. J. Nutr., 20:189-215. Stuedemann, J.A., Guenther, J.J., Ewing, S.A. and Morrison, R.D., 1968. Effect of nutritional level imposed from birth to eight months of age on subsequent growth and development patterns of full-fed beef calves. J. Anita. Sci., 27: 234-241. Swart, J.A. and Swart, J.C., 1965. The influence of early weaning on the subsequent growth of calves. Proc. S. Aft. Soc. Anim. Prod., 4:213-216. Thomson, E.F., Bickel, H. and Schfich, A., 1982. Growth performance and metabolic changes in lambs and steers after mild nutritional restriction. J. Agric. Sci. Camb., 98:183-194. Tudor, G.D. and O'Rourke, P.K., 1980. The effect ofpre- and post-natal nutrition on the growth of beef cattle. It. The effect of severe restriction in early post-natal life growth and feed efficiency during recovery. Aust. J. Agric. Res., 31:179-189. Tudor, G.D., Utting, D.W. and O'Rourke, P.K., 1980. The effect of pre- and post-natal nutrition on the growth of beef cattle. III. The effect of severe restriction in early post-natal life on the development of the body components and chemical composition. Aust. J. Agric. Res.. 31: 191-204. Verde, L.S., Joandet, G.E., Gill, E.A. and Torres, F., 1974. Level of restriction and sire effect on compensatory growth in Aberden Angus steers. Prod. Anim. (Argentina), 3:463-471. Waldman, R.C., Tyler, W.J. and Brungardt, V.H., 1971. Changes in the carcass composition of Holstein steers associated with ration energy levels and growth. J. Anim. Sci., 32:611-619. Wardrop, I.D., 1966. The effect of the plane of nutrition in early post-natal life on the subsequent growth and development of cattle. Aust. J. Agric. Res., 17: 375-385. Warner, R.G., 1961. Is hay required to develop rumen capacity? J. Dairy Sci., 44:1177-1178. Warner, R.G., Bernholdt, H.F., Grippin, C.H. and Loosli, J.K., 1953. The influence of diet on the development of the ruminant stomach. J. Dairy Sci., 36: 599-600. Warner, R.G., Flatt, W.P. and Loosli, J.K., 1956. Dietary factors influencing the development of the ruminant stomach. Agric. Food Chem., 4: 788-792. Wilkinson, S.C. and Mackie, C.K., 1988. The use of long chain n-alkanes for estimating intake and digestibility of herbage in growing cattle. Research Meeting No. 1, Br. Grassl. Soc., September 1988. Wilkinson, J.M. and Prescott, J.H.D., 1970. Beef production from grazing and silage with autumn-born calves. 2. The effects on the performance of cattle fed on silage of barley supplementation and of previous grazing. Anita. Prod., 12: 443-450.

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P. BERGE

Williams, P.V. and Macdearmid, A., 1983. Effect of overwintering plane of nutrition on subsequent growth at grass of beef steers. Anim. Prod., 36:515 (abstr.). Williams, P.V. and Macdearmid, A., 1987. Effects of severely restricted food intake and growth on subsequent appetite, growth and nitrogen balance of Friesian steers. Anita. Prod., 44:474 (abstr.). Winchester, C.F. and Ellis, N.R., 1956. Delayed growth in beef cattle. U.S.D.A. Tech. Bull., 1159. Winchester, C.F., Hiner, R.L. and Scarborough, V.C., 1957. Some effects on beef cattle of protein and energy restriction. J. Anita. Sci., 16: 426-436. Wright, I.A. and Russel, J.F., 1987. The effect of sward height on beef cow performance and on the relationship between calf milk and herbage intakes. Anim. Prod., 44: 363-370. Wright, I.A., Russel, J.F. and Hunter, E.A., 1986. The effect of winter food level on compensatory growth of weaned, suckled calves grazed at two sward heights. Anita. Prod., 43:211223. Wright, I.A., Russel, J.F. and Hunter, E.A., 1987. The effects of genotype and post-weaning nutrition on compensatory growth in cattle reared as singles or twins. Anita. Prod., 45: 423432.

RESUME

Berge, P., 1991. Effets a long terme de l'alimentation dans le jeune fige sur les performances ult6rieures des bovins a viande (revue bibliographique). Liw~st. Prod. Sci., 28:179-201 (en anglais). Une revue de la litt6rature a 6re effectu6e afin de relier les performances d'6ngraissement el les caract6ristiques de la carcasse des bovins ~. la strat~gie d'alimentation appliqu6e au cours de la premibre ann6e de la vie du veau. Une limitation pr6coce de la croissance entraine dans tous les cas un changement notable de la composition corporelle du veau. Ult6rieurement, en p6riode de r6cup6ration, les veaux ant& rieurement restreints ont une capacit6 d'ingestion d'aliment et une efficacit6 alimentaire sup& rieures a celles de leurs homologues non restreints. Contrairement aux bovins restreints plus tardivement, leur potentiel de croissance compensatrice est faible, notamment aprbs une restriction avant sevrage, et pratiquement ind~pendante de la s6v6rit6 de la restriction. A l'abattage m6me poids de carcasse, les differences initiales de composition corporelle ne sont en g6n6ral plus perceptibles, preuve que les traitements alimentaires appliqu6s dans le jeune fige n'ont pas d'effet ~ long terme sur ce param6tre. IIen a 6t6 conclu que la technique d'alimentation b, un stade precoce de d~veloppement ne peut 6tre envisag~e comme un moyen efficace de modifier a long terme la composition de la carcasse des bovins.

KURZFASSUNG

Berge, P., 1991. Langzeiteffekte der Ffitterung von K~ilbern auf nachfolgende Leistungen beim Rindvieh (Eine Uebersicht). Livest. Prod. Sci., 28:179-201 (aufenglisch). Eine Literaturstudie wurde durchgeffihrt um die Merkmale der Mastleistung und Schlacht-

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k6rperqualit~it beim Rindvieh in Verbindung zu setzen mit Ftitterungsstrategien w~ihrend des ersten Lebensjahres eines Kalbes. In allen Fallen bewirkt eine frfihe Wachstumsteinschr~nkung erhebliche Ver~inderungen in der K6rperzusammensetzung. Sobald die K~ilber sich bei freiem Ffitterungsregime wieder erholen k6nnen, wird ihre Trockensubstanzaufnahme h6her und die Futterumsatzrate ist gr6sser als bei restriktiv geftitterten Tieren. Das nachfolgende kompensatorische Wachstumspotential hingegen ist tief, vor allem wenn sie vor dem Absetzen restriktiv geftittert wurden. Es ist praktisch unabh~ingig yon der Restriktionsintersit~it im Gegensatz zu dem von Tieren, welche erst zu einem sp~iteren Zeitpunkt restriktiv geftittert werden. Bei der Schlachtung mit fixem kommerziellem Schlachtk6rpergewicht sind die fdiheren Unterschiede in der KiSrperzusammensetzung meist nicht mehr erkennbar. Dies weist daraufhin, dass die Ffitterungsmethoden im Kalbesalter keiner Langzeiteffekt auf dieses Merkmal haben. Daraus wird geschlossen, dass kein Beweis zur Beeinflussung der Schlachtk6rperzusammensetzung bei der Schlachtung dutch unterschiedliche FiJtterungsmethoden im friJhen Entwicklungsstadium, vorliegt.