J. Dairy Sci. 87:1380-1388
© American Dairy Science Association, 2004.
The Effects of Forage Provision and Group Size on the Behavior of Calves
C. J. C. Phillips
School of Veterinary Science, University of Queensland, Australia
Corresponding author: C.J.C. Phillips; e-mail: c.phillips{at}uq.edu.au.
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ABSTRACT
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The effects of providing alternative forages to individual or group-reared calves on their behavior were examined in 2 experiments. In experiment 1, 24 calves were reared in groups of 3 or individually in straw-bedded pens from age 1 to 7 wk. One-half of the calves in each treatment were provided with ad libitum cut perennial ryegrass herbage. Grass intakes and time spent eating grass were greater for grouped calves than for individual calves. Providing grass reduced concentrate intake of grouped calves and reduced the time that all calves, but particularly individual calves, spent eating straw bedding. Ruminating time was increased by offering grass to grouped calves compared with individual calves. Grass reduced the frequency of calves licking their buckets and their pen, vocalizing, and investigating their pen. Particularly for grouped calves provision of grass reduced all grooming. Group rearing reduced the frequency of calves licking their bucket, vocalizing, and investigating their pen, but had no effect on the frequency of pen licking. Calves were weaned at wk 7 and transferred to indoor silage feeding or grazing. Most effects of group rearing and grass provision were not maintained after weaning, but calves that had received grass ate for longer periods when turned out to pasture. In experiment 2, 72 calves were offered a mixture of straw, molasses, and pot ale syrup or grass hay. Calves offered the straw mixture ate more forage and concentrates and grew faster than calves offered hay. It was concluded that nontraditional forages, such as fresh grass and straw mixtures, could benefit the behavior and growth of calves compared with hay and straw.
Key Words: calf • grass • group size • isolation
Abbreviation key: CAP = calves at pasture, CI = calves inside
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INTRODUCTION
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The young calf has a strong motivation to suckle, which is not satiated by providing milk in buckets (Margerison et al., 2002). As a result, calves develop oral vices, such as excessive licking, sucking, and chewing of objects in their immediate environment, which can include the body parts of other calves, leading to rapid spread of infection in a group of calves (Smits and de Wilt, 1991). Such vices sometimes persist into adulthood, such as udder sucking in the female or prepuce sucking and tongue rolling in the male (Albright et al., 1991). Normally, the motivation for suckling declines once the calf starts ruminating (Margerison et al., 2002); therefore, it is important to provide palatable food that is sufficiently fibrous to stimulate rumination in young calves (Sambraus et al., 1984). Hay or straw is usually provided, but fresh grass may be more palatable and may be eaten at a younger age and in larger quantities, thus allowing earlier weaning of the calves or reduction of concentrate allocation. Alternatively, a straw mixture can be offered with low digestible straw enhanced by ingredients that are more digestible and palatable. European Union regulations now require that all calves over 2 wk of age have access to fibrous food. (At least 100 g/d per calf must be available at 2 wk of age, increasing to 250 g/d per calf at 20 wk of age.)
The young ruminant naturally learns to take solid feed by allelomimicry, using its mother as a model, if available, or peers (Key and MacIver, 1980; Phillips and Youssef, 2003). Calves in individual pens with solid sides have little opportunity for allelomimicry, and solid feed consumption may be delayed as a result. Such partial isolation increases the motivation to explore novel environments compared with group rearing (Arave et al., 1985), even though the calves are more fearful when isolated in a strange environment (Jensen et al., 1997). Calves in individual pens are more likely to develop oral vices, such as tongue playing, particularly if the pens have solid sides (Seo et al., 1998). Isolation also reduces the social abilities of calves (Jensen et al., 1999[Au: 1997? or new ref.?]), and this, coupled with fear of other calves, may reduce their adaptation to a novel environment when they are eventually mixed. Alternatively, conditioning calves to social stress at an early age may condition them to accept it more readily later in life (Albright and Stricklin, 1989; Arave et al., 1992).
The objective of the 2 experiments reported here was to investigate whether providing palatable forage can stimulate intake and reduce the incidence of abnormal oral behavior in calves and whether this is more likely in isolated calves, which have a greater motivation to explore, or grouped calves, which can learn from each other.
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MATERIALS AND METHODS
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Calves in experiments 1 and 2 were kept in accordance with the UK Code for the Welfare of Cattle.
Experiment 1
The effects on behavior of providing fresh cut grass and rearing in groups or individually were examined in 24 female calves. The calves were Friesian crosses with a mixture of beef breeds: 11 Hereford, 5 Charolais, 2 Simmental, 2 Limousin, 1 Belgian Blue, 1 Piemontese, 1 Blonde d’Aquitaine, and 1 Aberdeen Angus (mean BW = 49.0; SE = 0.81 kg). The calves were purchased at 7 to 10 d of age and were allocated by BW to pairs; one calf from each pair was reared individually, the other calf was reared in a group of 3. Individual pens were 0.9 x 1.8 m, and group pens were 1.8 x 2.7 m; both providing 1.6 m2 per calf. They were arranged in 2 rows, so that calves faced each other with a 2-m passage in between. The sides of the pens were of solid wooden boarding, except to the front, providing partial visual isolation for calves in individual pens. Individual and group pens were accommodated in separate, identical halves of a portal-framed building that prevented the transfer of vocal or olfactory signals between calves in the 2 treatments. All pens were provided with straw bedding, which provided the only form of forage for calves without fresh herbage. The straw was replenished daily if soiled. The calves remained in these pens from May 15 to July 6, a total of 53 d, at the end of which they were weighed. Weight gain of individual calves was calculated from end BW minus initial BW.
Feeding Management
One-half of the group and individual pens were allocated at random to receive ad libitum fresh cut herbage in racks (45-cm length per calf), with offered and refused herbage weighed daily. Herbage was from a perennial ryegrass (Lolium perenne, L.) pasture cut twice daily to a height of approximately 10 cm with a reciprocating blade plot harvester (Haldrup, Lovstock, Denmark). The other one-half received no herbage, but calves in both groups were able to consume straw bedding, complying with EU legislation requiring the provision of forage to calves. All calves were individually fed 0.5 kg of milk replacer (Llaeth y Felin; Bolac, UK) in 4 L of warm water in 2 feedings at 0900 and 1600 h. Milk was fed in buckets; after feeding, the buckets were washed, and fresh water was provided during the time between milk feedings. A pelleted concentrate feed (Denkavit Calf Cudlets 255; Dalgety Agriculture Ltd., Almondsbury, Bristol [CP, 180 g/kg; oil, 45 g/kg; fiber, 100 g/kg; and Na, 4 g/kg]) was offered ad libitum in a second bucket, and the offered and refused weights recorded daily.
Behavior Recording
The behavior of each calf was recorded at 10-min intervals for a continuous period of 24 h at the end of each week. Recording was by 4 experienced observers recording 6 h each. The duration of the following main behaviors were recorded at the end of each 10-min period: time spent eating grass, concentrate, and straw bedding; lying (with and without ruminating); and standing (with and without ruminating). In addition, whether there had been any incidences of jumping or kicking, tail swishing, drinking, grooming self or others, licking the bucket or pen, sniffing, calling, and rubbing (maximum 1 recording per 10-min period) was recorded.
Residual Effects
Following the treatment period, all calves were weaned at 7 wk of age and, within treatments, were allocated to pairs that were similar in BW. Within pairs, one calf was allocated at random to grazing, and the other was moved to a portal-framed building similar to that from which they had been housed in up to 7 wk of age. In this building, they were offered grass silage ad libitum at a feeding barrier. The grazing calves were kept in 4 separate paddocks of 35 x 35 m, each with 3 calves from a single treatment, and in the portal-framed building the calves were kept in 4 pens, again each containing the 3 calves from a single treatment. Grazing was on Italian ryegrass (Lolium multiflorum) pasture, which had previously been grazed by sheep to a height of approximately 4 cm and then rested until it reached the recommended height (10 cm) for cattle on this sort of pasture to achieve maximum intake (Phillips, 2001). Both groups were offered 1 kg/d of the same concentrate feed as offered pre-weaning. This second period of the experiment lasted for 4 wk, with one 24-h behavior study at the end of each week. Behavior was recorded by a team of observers, who moved between the 2 groups of calves at 10-min intervals. Within each 10-min period, the observer recorded the duration of feeding, lying (with and without ruminating), standing (with and without ruminating) times, and the frequency of drinking, rubbing themselves against posts in the enclosure, licking the enclosure, licking the navel of other calves, vocalization, tail swishing, grooming self and other calves, changes in lying position, walking, butting, and licking their teeth. In addition, for the outside calves, the development of grazing and ruminating behavior was recorded as follows by a single observer. The rate of biting while grazing and the ruminating chewing rate were recorded for each calf each week. The inter bolus interval during ruminating was recorded for 1 min per calf each week. After familiarizing the calves to a person’s presence behind them, the distance walked in 60 s was recorded when the calf was grazing. Calves were individually followed at a fixed distance with a surveyor’s wheel.
At weekly intervals during this 4-wk residual period, each calf was subjected to an open field test, monitoring their movement using a videocamera (National Panasonic WV-1450/B; Mitsushita, Uxbridge, Middlesex). The camera was installed 2.4 m above the floor of the 3- x 3-m pen and connected to a videodigitizing modem (Sallinen and Hatunen, 1993) installed on a computer (Matmos 486; Middlesex, United Kingdom). The modem recorded each occasion (termed a ‘movement’) that 15% of the integer-valued picture elements (pixels) changed brightness, reflecting major calf movements in the pen. For each ‘movement,’ the proportion of pixels changing intensity, which was termed the movement strength, was recorded. This modem recorded the number of times that 
4% of the pixels on the video screen changed brightness because of movement of the calf (calves were dark on a white floor). Measurements were made every 5 s over a 5-min period, and the order of testing was randomized for calves within treatments.
Statistical Analysis
Grass intakes could not be analysed statistically, as the group intakes could not be apportioned to calves within group. Other data were tested for normality by the Anderson-Darling test of the statistical package Minitab (1995), and several were transformed by taking the square root, logarithm, or other transformations, which are detailed in the units for each parameter given in the tables. Some could not be rendered normal by mathematical transformation and were analyzed by the Kruskal-Wallace test for non-parametric data.
It is recognized that social facilitation of certain behaviors could exist, which may make it unwise to use individual calves as replicates; however, there is currently insufficient information on which behaviors are affected. The use of individual cattle within grazing groups as replicates has been previously discussed (e.g., Phillips, 1998, 2002).
Experiment 2
The provision of grass hay or a straw mixture on the behavior of Friesian calves was examined in 72 male and female calves of mean BW 43.1 + 0.50 kg. Calves were allocated to the 2 treatments in alternate pairs in their order of calving, each pair sharing a feeding rack for the forage. All calves were penned from birth until abrupt weaning at 10 wk of age and were fed 500 g/d of warm milk replacer (BOCM Pauls; 17% oil, 23% CP, 7.2% ash, and 0.25% fiber). In addition, a concentrate mix containing 88% rolled barley and 12% high CP feed (soybean-based; Masterfeed, Chapman and Pearson, UK) was fed ad libitum twice daily, and intakes were measured once weekly. The straw mixture was mixed thoroughly by hand and contained 70% barley straw (chopped to lengths of approximately 10 cm), 15% cane molasses (Farmblend; Rumenco, Stretton, UK), 15% pot ale syrup (Rumenco, UK; contained 60% DM and 13.4 mJ metabolizable energy and 16% CP per kg of DM), and 0.5% propionic acid. Forage was weighed and offered twice weekly in sufficient quantities to ensure that at least 10% remained as a residue. Both forage and concentrate feeds were sampled weekly for analysis of CP, modified ADF, OM, and DM contents by the procedures of AOAC (1984).
Calves were weighed upon starting the experiment at 2 wk of age and at weaning, when the experiment terminated. A 24-h behavior study was conducted when the last calves entered the experiment, by which time the first calves were in their final week before weaning. Thirty-eight randomly selected calves were recorded by 3 observers in rotation. Every 15 min, an instantaneous scan sample was made, recording whether the calves were lying, drinking, ruminating, feeding on forage or concentrate, drinking, grooming themselves, grooming another calf, or licking the pen.
Results of the randomized block experiment were analyzed using the General Linear Models procedure, with forage type and pairs (block) as factors, using the statistical package Genstat (LAT, 1985). Data were tested for normality by the normal score of the statistical package Minitab (1995).
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RESULTS
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Experiment 1
There were no significant effects of treatment on the BW gain of calves (Table 1
). Grass intakes were greater for grouped calves than for individual calves (2.1 vs. 0.7 kg DM/d per head), and the grouped calves spent longer eating grass. Concentrate intakes were reduced in grouped calves compared with individual calves. Providing grass reduced concentrate intake and concentrate eating time in the individual calves, but not in grouped calves. Time spent eating straw bedding was greater for grouped calves than for individual calves and was reduced by offering grass, particularly for the grouped calves. The time spent ruminating was increased by a similar amount for grouped calves vs. individual calves (333 vs. 248 min/d) and by offering grass (+ grass, 330 min/d; – grass, 252 min/d). The frequency of drinking was decreased both by group rearing and by offering grass. The grouped calves increased urination and defecation frequency.
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Table 1. Body weight gain, and ingestive and eliminative behavior of calves housed individually (n = 12) or in groups (n = 12) and offered grass (n = 12) or no grass (n = 12) in experiment 1. Values in parentheses indicate means, where transformed values were used for statistical analysis.
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The provision of grass reduced the total time that grouped calves spent lying, but did not affect time that individual calves spent lying (Table 2). This result was due to a reduction in time spent lying doing nothing, rather than lying while ruminating or lying while eating straw. The latter was more common in individual calves, and, in that situation, it was reduced by the provision of grass. Although grass reduced the time that calves spent lying doing nothing, it did not affect the number of lying bouts. The provision of grass also reduced the time spent standing doing nothing (not feeding or ruminating). Grouped calves tended to spend less time than individual calves standing doing nothing. Little time was spent standing ruminating, and it was not affected by treatment. The total standing time, including time spent feeding, increased in grouped calves fed grass, which was principally a reflection of their increased eating time.
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Table 2. Lying, standing, and other activity behaviors of calves housed individually (n = 12) or in groups (n = 12) and offered grass (n = 12) or no grass (n = 12) in experiment 1.
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The frequency of calves grooming themselves was increased by providing grass to the grouped calves but did not change when grass was provided to the individual calves; also, frequency of the grouped calves grooming others was reduced (Table 3). The frequency of licking the bucket and sniffing was substantially reduced by providing grass and was greater for individual calves than for grouped calves. Providing grass also reduced the frequency of licking the pen, but group size had no effect. Vocalization only occurred in individual calves and tended to be reduced by grass provision. There was no difference between treatments in jumping or kicking behavior, but tail swishing was increased by offering grass and by group housing.
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Table 3. Grooming, licking and other interactions with the environment in the behavior of calves housed individually (n = 12) or in groups (n = 12) and offered grass (n = 12) or no grass (n = 12) in experiment 1. Values in parentheses indicate means, where transformed values were used for statistical analysis.
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The residual effects demonstrated that there tended to be an increase in eating time and time spent self grooming for calves that had received grass, compared with calves that had not received grass, when they were outside but not when they were inside (Table 4). The lying ruminating time was longest for those fed inside, especially when they had not received grass. Those calves that received grass inside had fewer drinking, rubbing, and tail swishing bouts per day in the first 4 wk of the residual period (Table 5). In addition, when compared by the Kruskal-Wallace test, calves at pasture (CAP) performed more of the following incidences per day than did calves inside (CI): changing lying position (CAP = 0.72, CI = 0.39; P = 0.05), vocalizing (CAP = 1.0, CI = 0.4; P < 0.01), butting (CAP = 0.27, CI = 0.10; P = 0.10), and licking the navel of other calves (CAP = 0.12, CI = 0.02; P < 0.001). Also when compared by the Kruskal-Wallace test, CAP performed fewer of the following incidences per day than did CI: walking (CAP = 0.3, CI = 1.5; P < 0.001) and licking their enclosure (CAP = 0.3, CI = 1.5; P < 0.001).
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Table 4. Effects of the provision of grass and keeping cattle inside or outside on the behaviors exhibited during the residual period of experiment 1 (indoor or outdoor housing), analyzed as the differences between 4 treatments (n = 6) by the Kruskal-Wallace test for non-parametric data. Only effects where P < 0.10 are presented.
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Table 5. Effects of providing grass (n = 12) vs. no grass (n = 12) during the indoor housing period on the behavior of calves in the residual period of experiment 1, analyzed by the Kruskal-Wallace test for non-parametric data. Only effects where P < 0.10 are shown.
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Experiment 2
Straw mixture was of higher fiber and lower CP content than hay, with similar DM and OM contents (Table 6). The calves offered straw mixture ate more forage and concentrate and grew faster than those offered hay (Table 7). The difference in concentrate intake only occurred in the last 2 wk of the experiment. The calves offered straw mixture also spent less time lying not ruminating and more time lying ruminating, with increased total time spent ruminating. There was no difference in the licking behavior of the calves between treatments.
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DISCUSSION
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Both of the novel forages for calves, fresh grass and straw mixture, were consumed in significant quantities, compared with the intake of concentrate. The palatability of these feeds may be due to their high soluble carbohydrate content, intrinsic to the fresh grass and added in the form of molasses and pot ale syrup in the straw mixture. The high intake of straw mixture stimulated growth, compared with hay, which stimulated concentrate consumption, as has been previously recorded (Kellaway et al., 1973; Thomas and Hinks, 1982). The maximum feed intakes of calves has previously been recorded at 20 to 23% ADF (Kang and Leibholz, 1973), which is closer to the fiber concentration in the straw mixture diet. The low CP content of the straw mixture, compared with hay, might be expected to reduce straw mixture intake, and taste characteristics might have had a greater effect on intake than post-digestive feedback of nutritional value. The increased intake of calves offered the straw mixture, compared with hay, probably led to the increase in rumination time.
The first experiment demonstrated that several behaviors were statistically affected by group size, but it does not necessarily follow that individual animals cannot be used as replicates in statistical analysis, as they could be affected by a herd effect, but still behave independently of other individuals. Hence, animals may be aware of the number of conspecifics present, and this affects their behavior, but the behavior of an individual may or may not be affected by any other individual. A group size effect was noticeable in the consumption of grass, with much greater intakes for grouped calves. This group size effect was accompanied by a reduction in the consumption of concentrates and the time spent eating concentrates. Time spent eating straw was also socially facilitated, and the reduction in straw consumption time was greater in the grouped calves than in the individual calves. Therefore, it is likely that there was a greater substitution of grass for concentrates in the individual calves and for straw in the grouped calves. It may be that the individual calves ate extra concentrates because of boredom, which was reduced if another food or grass was offered. This could be because grass is more palatable than concentrates, which might derive from an increased content of water-soluble carbohydrates, protein, or more natural feeding behavior or taste.
The increase in grass intake for grouped calves compared with individual calves suggests social facilitation of this behavior, but not subsequent concentrate intake. Increased grass intake might have been influenced by the method of presentation, as the act of taking food into the mouth was only observable by other calves for grass and straw, not when the calves consumed concentrates from a bucket. It may be important that individual calves in the grass-fed group could see others consuming grass and were able to join in feeding from the same source, whereas with concentrates fed from individual buckets this was not the case. In other species, an ability to join with other animals in performing the behavior at the same site is important for social facilitation (Olsson et al., 2002). Hence, straw consumption time, not concentrate, was reduced for grouped calves when they were offered grass. This result suggests that the sight of feed being taken into the mouth is the relevant stimulus for social facilitation of feeding behavior in calves, rather than the feeding posture of other calves. The facilitative effect was on eating time not rate of intake. (Mean grass intake rates of grouped and individual calves were 12 and 13 g DM/min, respectively.) Matching the color of a novel feed consumed by monkeys in the presence of conspecifics with that of their main feed increased eating time, but not the rate of intake (Visalberghi and Addessi, 2001). Hence, visual stimuli from feed are important for social facilitation to synchronize eating behavior, and these were not present during concentrate feeding.
An alternative hypothesis is that the calves were more easily socially facilitated to take fibrous foods, such as hay or straw, than high-energy feeds, such as concentrate, because of a greater need for fiber than for energy or protein. However, it is unlikely that such young animals would have a major need for fiber when they would not be consuming much forage if their nutritional requirements were still being met from a cow’s milk, as would be the case for a calf suckling a beef cow. Conceivably, fibrous feeds, which are slower to be consumed than concentrates, redirect the suckling need more effectively than concentrates, but evidence is not available in support of this.
The lack of substitution of concentrates for grass in grouped calves might be expected to increase BW gain, as occurred when calves consuming straw mixture in experiment 2 saw increased BW, rather than to decrease their concentrate intake. However, the inclusion of fresh grass in the diet could reduce the gut contents (Balch and Line, 1957; Tayler and Wilkinson, 1972) compared with more fibrous feeds, such as the straw bedding, masking any increase in BW gain. Diurnal variation in BW might have been affected by the provision of grass, thus confounding BW gain of the different treatments. The stimulation of rumination by forage consumption would be expected to accelerate rumen development (Williams et al., 1985) and allow the calves to make use of forages earlier. This was demonstrated by the increase in grazing time of calves that had received grass. The absence of any increase in time spent eating silage (Table 4
) demonstrates that there is probably a psychological effect relating consumption of fresh grass to that of grazing pasture that does not extend to silage, which is less similar than fresh grass in taste, odor, pH, and texture to pasture.
The reduction in drinking frequency with provision of grass and grouped housing, which increased grass intake, reflects the effect of both on water intake from grass. Therefore, the total water intake was likely to have been similar between treatments (Castle and Watson, 1973). However, the urination and defaecation frequency were increased in grouped calves, but were unaffected by the provision of grass, which suggests that these 2 behaviors were socially facilitated, with more regular and perhaps smaller excretions by grouped calves.
Individual calves spent more time than grouped calves licking their buckets and sniffing their pen, but not licking their pen. Pathological mouthing of inedible environmental items is common in individually penned calves (Margerison et al., 2002). Such mouthing is believed in other species to function as a temporary physiological stress-coping strategy (Wood-Gush, 1973; Cronin et al., 1986), reflecting a failure to adapt to environmental conditions (McBride, 1984; Tennessen, 1988). It has also been attributed to the inability of individually penned calves to lick and suck each other (Fraser, 1980). However, it should be noted that the amount of licking activity between grouped calves was low in relation to the amount of self-grooming activity. The frequency of licking bouts between grouped calves was more than the reduction in the frequency of bucket licking as a result of group housing, lending support to this hypothesis.
The increase in lying ruminating time for CI that had not received grass before may represent a slower development of rumination processes for these calves. This behavior only related to calves consuming fibrous silage rather than the more easily digested fresh grass. The increase in tail swishing in those calves that had not had grass when they went outside might have been demonstrative of satisfaction of having access to fresh grass, but it is also possible that the increase in all 3 behaviors (drinking, rubbing, and tail swishing) was because these calves were spending less time grazing than those that had received grass before. Evidence from young rabbits suggests that the provision of fresh grass is valued for just a few weeks, after which the rabbits are as willing to consume concentrates as grass (Leslie et al., 2003).
A final consideration in relation to the advantages of providing fresh grass on the welfare of a young calf is that these advantages will be negated if the grass is heavily contaminated with parasites, as the young calf cannot mount an effective immune response.
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CONCLUSIONS
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Fresh grass provided to calves before weaning was eaten in substantial quantities, especially when the calves were in housed in groups. There was a reduction in concentrate intake in individually housed calves offered grass as well as a reduction in time spent consuming straw bedding in group-housed calves offered grass. A straw mixture was also consumed in considerable quantities, in comparison with hay, and this stimulated concentrate intake and enhanced growth. The provision of grass and the straw mixture stimulated rumination, and grass reduced the amount of behaviors that are in some circumstances considered problems (oral stereotypies, e.g., bucket and pen licking; searching behaviors, e.g., calling, sniffing; and appeasement behavior, e.g., allogrooming (grooming others)). Grass consumption increased behaviors that may indicate satisfaction (self-grooming, tail swishing, and rubbing themselves).
Group-reared calves ate more grass and ruminated for longer than individually reared calves. Bucket licking and searching behaviors (calling and sniffing) were less frequent, and total grooming (allogrooming plus grooming self) and tail swishing were more frequent, in the group-reared calves.
Offering fresh grass tended to increase grazing time and reduce the biting rate when the calves were turned out to pasture. Most other behavioral changes were not maintained after the treatments had ended, although there was evidence of an increased level of satisfaction behaviors (tail swishing, rubbing) when calves that had not received grass were turned out to pasture.
The results emphasize the importance of providing adequate palatable forage to young calves to promote growth, normal behavior, and rapid adjustment to grazing following turnout to pasture.
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ACKNOWLEDGEMENTS
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M. Rouse, R. Lambert, I. Meneer, A. Nabels, S. Timsane, and L. Veide helped in collection of data for Experiment 1, and R. A. Njei and L. Weiguo helped in Experiment 2. The assistance of J. B. Thomas, J. Pilling, and J. Ffridd in managing the calves is gratefully acknowledged. The University of Wales, Bangor, provided facilities for this experiment.
Received for publication June 23, 2003.
Accepted for publication December 18, 2003.
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ABSTRACT
INTRODUCTION
