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J. Anim. Sci. 2003. 81:116-121
© 2003 American Society of Animal Science

Fenceline contact of beef calves with their dams at weaning reduces the negative effects of separation on behavior and growth rate1

E. O. Price*,2, J. E. Harris*, R. E. Borgwardt*, M. L. Sween* and J. M. Connor{dagger}

* Department of Animal Science, University of California, Davis 95616 and and {dagger} University of California Sierra Foothill Research and Extension Center, Browns Valley 95918

2 Correspondence:
phone: 530/752-6789; fax: 530/752-0175; E-mail:
eoprice{at}ucdavis.edu


    Abstract
 Top
 Abstract
 Introduction
 Materials and Methods
 Results
 Discussion
 Implications
 Literature Cited
 
The purpose of this study was to examine the hypothesis that fenceline contact between beef calves and cows at weaning reduces indices of behavioral distress and associated temporary reductions in weight gain. One hundred Angus/Hereford-cross calves were randomly assigned to five treatments for 7 d in each of 3 yr to determine the effect of different weaning techniques on their behavior and subsequent growth. Treatments were 1) fenceline separation from dams on pasture (F-P), 2) total separation from dams on pasture (S-P), 3) total separation from dams in a drylot (corral) preconditioned to hay (S-D-P), 4) total separation from dams in a drylot not preconditioned to hay (S-D-NP), and 5) nonweaned controls on pasture (C-P). At the end of the 7-d postweaning period, all calves were placed on pasture in large groups. Calves were weighed weekly for 10 wk. In the days following weaning, F-P and C-P calves spent more time eating (grazing or eating hay) than S-P and S-D-NP calves (P < 0.05). The S-P calves spent more time walking (pacing) than calves in the other four treatments (P < 0.05), which did not differ. The S-P calves also spent less time lying down than C-P, F-P, and S-D-P calves (P < 0.05); S-P and S-D-NP calves did not differ in lying time. The F-P calves vocalized less than S-P and S-D-NP calves (P < 0.05). In general, treatment differences were greatest during the first 3 d following weaning with d 2 (20 to 30 h after weaning) showing the greatest disparity. The F-P calves spent approximately 60% of their time within 3 m of the fence separating them from their dams during the first 2 d following weaning, whereas F-P cows spent about 40% of their time within 3 m of the fence during this period. Postweaning cumulative body weight gains of the F-P calves were greater than the gains recorded for the calves in the three totally separated treatments (which did not differ). The F-P calves gained 95% more weight than the average calf in the three totally separated treatments in the first 2 wk and were still heavier at 10 wk (21.4 vs 11.0 kg, respectively, at 2 wk and 50.0 vs 38.2 kg, respectively, at 10 wk; P < 0.05). It was concluded that providing fenceline contact between beef calves and cows for 7 d following weaning reduces behavioral indices of distress seen in the totally separated calves. In addition, fenceline contact with dams at weaning minimizes losses in weight gain in the days following separation. Totally separated calves did not compensate for these early losses in weight gain even after 10 wk.

Key Words: Beef Cattle • Behavior • Calves • Growth • Weaning


    Introduction
 Top
 Abstract
 Introduction
 Materials and Methods
 Results
 Discussion
 Implications
 Literature Cited
 
Artificial early weaning in mammals is typically accompanied by physical separation of mother and young, termination of milk feeding, and often a change in solid feeds and living environment. These changes can result in both behavioral and physiological indices of distress in cattle (Lefcourt and Elsasser, 1995; Stookey et al., 1997; Lay et al., 1998), sheep (Orgeur et al., 1998; 1999), horses (McCall et al., 1985; 1987; Holland et al., 1996), and red deer/wapiti (Haigh et al., 1997; Church and Hudson, 1999; Pollard and Littlejohn, 2000). The purpose of this study was to determine the extent to which fenceline contact between beef cows and their calves at weaning reduces the negative effects of separation and termination of milk feeding on calf behavior and growth. Nicol (1977) and Stookey et al. (1997) studied this topic by comparing the postweaning growth and behavior of calves that had either fenceline contact with their dams or were abruptly and completely separated at weaning. In both studies, the fenceline contact calves initially gained more weight than the abruptly separated calves, but after 2 to 3 d there was no difference between treatments. Using traditional behavioral indicators of comfort, such as eating, lying down, and rumination, Stookey et al. (1997) showed that the well-being of newly weaned calves was improved if allowed some social contact with their dams at weaning. In contrast to these earlier studies, the present study employed a nonweaned control group to determine the full extent to which fenceline contact with dams at weaning mitigated the distress associated with physical separation and termination of milk feeding.


    Materials and Methods
 Top
 Abstract
 Introduction
 Materials and Methods
 Results
 Discussion
 Implications
 Literature Cited
 
Animals and Management.
One hundred Angus/Hereford-cross heifer calves and the respective dams of 40 calves were used in each of 3 yr (1998, 1999, and 2000). The calves were born and maintained on pasture at the University of California Sierra Foothill Research and Extension Center, Browns Valley, CA. Ten days prior to weaning in May of each year, the 100 calves were assigned to 10 equal-size groups so that average body weights for the calves at weaning were similar in each group (Table 1Go). Three days prior to weaning, each calf in five of the ten groups received an individual identification tag consisting of 10-cm high black numbers on each end of a 13- x 46-cm white vinyl strip glued to the animal’s hair (KaMar adhesive; KaMar, Inc., Steamboat Springs, CO) across the back, posterior to the shoulders. Calves in each group were numbered zero to nine. The dams of 20 calves (two groups) received back tags with numbers identical to their respective calves. The calves averaged 207, 228, and 203 d of age at weaning in 1998, 1999, and 2000, respectively. Eighty percent of the animals used were born within a 35-,27-, and 55-d period in 1998, 1999, and 2000, respectively.


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Table 1. Mean (±SE) body weight (kg) of calves at weaning by treatments and years
 
Treatments.
In each of the 3 yr, two groups of 10 calves were assigned to each of the following five treatments: 1) nonweaned controls on pasture (C-P), 2) fenceline separation from dams on pasture (F-P), 3) total separation from dams on pasture (S-P), 4) total separation from dams drylot (corral) preconditioned to alfalfa hay in a drylot for 10 d immediately prior to weaning (S-D-P), and 5) total separation from dams drylot not preconditioned to hay (S-D-NP). Dams were left with calves in the C-P treatment. In the F-P treatment, dams were housed in a pasture adjacent to their calves. Fences separating calves and cows in the F-P treatment were made of five strands of barbed wire overlaid with wire-mesh with openings too small to accommodate a calf’s head. Treatments were administered for only the first 7 d following weaning. From d 7 to d 28 postweaning, calves (from all treatments) were placed in two groups of 50 animals each on pasture. On d 28, calves were placed in a single group of 100 until d 70 (10 wk) when body weight data were collected for the last time. The C-P calves were weaned (total separation - on pasture) at 8 wk (56 d postweaning).

Procedure.
On the day of weaning in each year, calves were weighed and placed in 10 pastures/corrals in assigned groups of 10 animals. Pastures ranged in size from 0.69 to 4.57 ha (mean = 2.34 ha), and corrals were 12 x 24 m with dirt substrate except for a 3- x 12-m cement slab at one end. Water and alfalfa hay were available at all times. Pasture treatment groups were assigned to different areas each year.

Behavioral observations began on the day of weaning after animals had been placed in their respective enclosures (about 1200). One group of 10 calves (numbered) from each of the five treatments were observed for their behaviors on the first 3 d following weaning in 1998 and the first 5 d in 1999 and 2000. Unnumbered calf groups were not observed for behavior but were monitored for weight gain.

One or two observers were assigned to each of the five numbered groups in each year. The behaviors of individual animals in each group were recorded using instantaneous sampling (Martin and Bateson, 1993) every 60 s in round-robin fashion. Except on d 1 and d 5, animals were observed from 0800 to 1000, 1200 to 1400 and 1600 to 1800. The early morning observation was not possible on the day of weaning (d 1), and the late afternoon observation was not made on d 5. Behaviors recorded included eating (grazing or eating hay), walking (pacing), lying down, vocalizing, proximity of F-P calves and cows to the fence separating them (3 m or less), and proximity of calves in the C-P treatment to their dams (3 m or less). Except for vocalizations, behaviors were summarized for each individual on each day as the percentage of total instantaneous sampling points in which the behavior was exhibited. Vocalizations were recorded for groups rather than individuals since the close proximity of animals often made it difficult to determine which animal emitted the sound.

Calves were weighed at weaning and every 7 d following weaning for 10 wk to assess the effects of the different weaning treatments on short-term and long-term weight gain. Gathering and weighing the animals were conducted in small groups to minimize the effect of defecation and urination in the holding pens and working chutes on week-to-week body weight measurements.

Statistical Analyses.
All variables except for the proximity scores were analyzed using the General Linear Models Repeated Measures Analysis of Variance procedure (SAS Inst. Inc., Cary, NC) to test for differences due to treatments, years and days (behavior), or weeks (body weight). Bonferroni t-tests were used to test for the significance (P< 0.05) of paired mean comparisons when significant main or interaction effects were obtained. Because behavioral data were not collected for all treatments on d 4 and 5 in the first year of the study, the data were analyzed in two ways, the first 3 d in all 3 yr and all 5 d in yr 2 and 3. The behavioral data were also subjected to a third analysis using group means as the statistical unit, considering the possibility that some behaviors may have been socially facilitated within treatment groups.

Proximity data were analyzed only for a day effect since comparison of the F-P and C-P treatments was confounded by the presence of the fence in the F-P treatment, which artificially influenced the animals’ spatial relationships. The Friedman Two-Way Analysis of Variance (Siegel and Castellan, 1988) was used in these analyses, and data for the 3 yr were analyzed separately.

Body weight data were expressed as the cumulative weight gain of individual animals (over their weaning weight) at each weekly weighing. In 1999, over 85% of the calves contracted infectious keratoconjunctivitis ("pinkeye") in at least one eye during the first 9 wk postweaning. Consequently, the weight gain data in this year were artificially variable and not reported.


    Results
 Top
 Abstract
 Introduction
 Materials and Methods
 Results
 Discussion
 Implications
 Literature Cited
 
Behavior.
There was close agreement between the results of the three statistical tests used to analyze the behavioral data (first 3 d in all 3 yr, all 5 d in yr 2 and 3, and group means).

Significant main effects for treatments and days (P < 0.002) were obtained for percentage of time spent eating (grazing or eating hay) in all three analyses. Both C-P and F-P calves spent more time eating than the totally separated calves (P < 0.05). Differences during the first 3 d postweaning are presented in Table 2Go. The S-P and S-D-P calves did not differ in eating time and S-D-P calves ate more than S-D-NP calves during this period. Except for the S-D-NP calves, treatment differences in eating tended to decline on d 4 and 5 of observation. Time spent eating during behavioral observations on d 2 (20 to 30 h after weaning) was less than on d 3 and d 4 (P < 0.05) while eating times on d 1 and d 5 were intermediate and did not differ from any other day (P > 0.05). There were no year differences for this variable.


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Table 2. Mean (±SE) percentage of observations in which calves were exhibiting various behaviors on d 1 through 3. Years are combined; N = 30 animals per cell
 
Significant main effects for treatments and days (P < 0.001) were also obtained for percentage of time calves spent walking (pacing). Walking was typically around the perimeter of the groups’ pasture or enclosure. The S-P calves were observed walking more than the other four treatments (P < 0.05; Table 2Go). Walking time was intermediate for the S-D-NP calves and did not differ (P > 0.05) from the S-P calves when all 5 d were included in the analysis. Walking was greatest on d 1 and 2 (Table 2Go) and then gradually decreased over d 3, 4, and 5 (P < 0.05). A significant main effect for years (P < 0.01) resulted from calves walking more in 1998 than in 1999 or 2000.

The main effect for treatment for percentage of time spent lying down was significant (P < 0.001). In the first 3 d of observation, S-P calves spent less time lying down than calves in the C-P, F-P, and S-D-P treatments (P < 0.05; Table 2Go). The S-P and S-D-NP calves did not differ (P > 0.05). Time spent lying down also differed over days (P < 0.003). Calves were observed lying down less on d 1 and 2 than on d 4 and 5 (P < 0.05) with d 3 being intermediate. Calves laid down more in 1998 than in 1999 or 2000 (P < 0.05).

Treatments, days, and years were significant (P < 0.001) when the data for vocalization frequency were analyzed. Calves in the C-P treatment almost never vocalized (Table 2Go). The S-P and S-D-NP calves vocalized more than F-P calves (P < 0.05) with S-D-P calves intermediate (Table 2Go). The S-P, S-D-NP, and S-D-P calves did not differ (P > 0.05). Vocalizations were most frequent on d 2 and least frequent on d 4 and 5 (P < 0.05) with d 1 and 3 intermediate (Table 2Go). Calves vocalized more in 1998 than in 1999 or 2000 (P < 0.05).

On d 1 and 2, F-P calves spent 62% and 61% of their time, respectively, within 3 m of the fence separating them from their dams. In general, F-P calves moved away from the fence during this period only to graze and obtain water. Fenceline contact declined (P < 0.001) on d 3, 4, and 5 (46%, 24%, and 31%, respectively). The F-P cows spent 38% and 45% of their time within 3 m of the fence on d 1 and 2, respectively, followed by a reduction (P < 0.001) in contact time over subsequent days (26%, 12%, and 3% on d 3, 4, and 5, respectively). The C-P calves were observed within 3 m of their dams 58%, 51%, 56%, and 53% of the time on d 1, 2, 4, and 5, respectively, and 39% of the time on d 3.

Body Weight.
A severe outbreak of pinkeye after the weaning treatment period in 1999 precluded use of the body weight data in that year. Treatment differences were found for cumulative weight gain over the 10-wk postweaning period (P < 0.001), and treatment differences were generally greater in the weeks immediately following weaning than later. At 2 wk postweaning, F-P calves had gained 95% more weight than the average calf in the three totally separated groups (21.4 vs 11.0 kg, respectively; P < 0.05; Table 3Go). Weight gains of the three totally separated groups did not differ (Table 3Go). Two-week cumulative weight gains for C-P and F-P calves were similar. At 10 wk postweaning, F-P calves were still about 10 to 11 kg heavier (P < 0.05) than calves in the three totally separated treatments (50.0 vs 38.2 kg, respectively), and, again, the three totally separated groups did not differ (Table 3Go). The C-P calves had gained 23% more weight than F-P calves. Calves gained more weight in 2000 than in 1998 (P < 0.01). A treatment by year interaction (P < 0.01) was found for 10-wk weight gains. The difference in cumulative weight gain between F-P calves, and calves in the three totally separated treatments was greater in 2000 than in 1998.


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Table 3. Mean (±SE) cumulative weight gain (kg) of calves (N = 20 per treatment per yr) at 2 and 10 wk postweaning in years 1998 and 2000a
 
No treatment differences were found in the rate at which pinkeye was first observed in individual animals in 1999. On d 21 postweaning, 11 C-P (55%), 7 F-P (35%), 7 S-P (35%), 7 S-D-P (35%), and 9 S-D-NP (45%) calves showed some evidence of the virus in at least one eye.


    Discussion
 Top
 Abstract
 Introduction
 Materials and Methods
 Results
 Discussion
 Implications
 Literature Cited
 
In general, the behavior of the F-P calves in the days following weaning resembled that of the C-P calves more than the calves totally separated from their dams. Except for vocalization frequency, no statistically significant differences were found between the F-P and C-P calves in time spent eating (grazing), walking, and lying down. At the other end of the scale, the behavior of S-P and S-D-NP calves was similar. Calves in these latter treatments spent less time eating (grazing), particularly in the first 3 d following weaning. They also spent more time walking, less time lying down, and exhibited higher frequencies of vocalization than the C-P and F-P calves. The S-D-P calves tended to be intermediate to the C-P/F-P and the S-P/S-D-NP treatments with respect to most behaviors. Stookey et al. (1997) also found fenceline-weaned calves to spend proportionately less time walking (pacing) and vocalizing and more time eating and lying down than totally separated calves, especially during the first 2 d after weaning.

Eating (grazing) by the F-P calves tended to occur in synchrony during the first 3 d following weaning. Most of the 10 calves in each group would together graze away from the fence and then return to stand or lie down near the fence. Distance traveled from the fence while grazing tended to increase over days following weaning until the calves failed to consistently return to the fenceline. A similar behavior was exhibited by their dams. Veissier and LeNeindre (1989) showed that weaned calves gather and have more social encounters with one another than similar-age calves which remained with their mothers. The somewhat coordinated grazing and resting behaviors exhibited by the fenceline calves in the present study is consistent with the hypothesis that social bonds between calves are strengthened at weaning and that calf groups tend to behave as a unit (Veissier and LeNeindre, 1989).

The F-P calves in 1998 and 1999 appeared relatively independent of their mothers by d 4 after weaning, while in 2000 the F-P calves still returned to the fenceline after grazing on d 5. The 2000 calves were almost identical in age to the 1998 calves at weaning and were slightly heavier than the 1999 calves; thus, it is unlikely the more lasting attraction of the 2000 F-P calves to their dams was because of factors related to age or size.

Walking (pacing) was most evident in the S-P calves. Being on pasture, the animals in this treatment had access to a relatively large area. Much of their walking time on d 1 and 2 was spent traversing the perimeter of their enclosure. Walking by the calves in the two totally separated drylot (corral) treatments (S-D-NP and S-D-P) was minimal due to their confinement, which likely discouraged movement.

Lying down was recorded because it is commonly associated with a nonreactive state in animals. Of the five treatments, S-P calves spent the least amount of time lying down on d 1 and 2, coinciding with their relatively high rates of walking. The S-D-NP calves also spent little time lying down on d 1 and 2 in spite of their lower walking time and relative confinement. Time lying down increased for these treatments over days following weaning, suggesting a gradual adaptation to their new environments and the absence of their dams.

Vocalization frequencies were greatest in the S-P and S-D-NP treatments and least in the C-P and F-P groups. The C-P calves almost never vocalized because they were not separated from their dams. Vocalization frequencies recorded were greatest in those groups hypothesized to experience the highest levels of distress at weaning (i.e., totally separated calves).

Vocalization frequencies were greater on d 2 than on d 1 or 3. Data collection on d 1 was completed within 6 h of weaning, which may not have been a long enough period for the effects of separation to be fully manifested in the behavior of the calves. The relatively high vocalization rates on d 2 suggested that motivation to contact their dams had reached a peak at that time. Vocalization frequencies gradually declined after d 2 suggesting adaptation to being separated from their dams and the weaning environment. These results are consistent with other studies on weaning in beef calves (Stookey et al., 1997), sheep (Orgeur et al., 1998), horses (Holland et al., 1996), and wapiti (Haigh et al., 1997; Church and Hudson, 1999; Pollard and Littlejohn, 2000).

Year differences in age or body size at weaning could not explain the year differences found for some behaviors.

Decreased eating and lying down and increased walking and vocalizations associated with weaning could be related to social separation, the termination of milk feeding or a combination of these factors. Haley et al. (2001) noted that 71/2-mo-old beef calves rarely vocalized if they were abruptly prevented from sucking but remained with their dams. When totally separated from their dams after being unable to suck for 4 d, the calves vocalized 84% less, walked 79% less, and spent 24% more time eating than separated control calves, which had not been previously prevented from sucking.

Adaptation to separation was exhibited in the F-P treatment by calves and cows gradually becoming more independent of one another and spending less time along the fenceline. With few exceptions, F-P cows remained highly responsive to their calves through d 2. Udders appeared very distended on d 2, and it was assumed that cows were highly motivated to have their calves suck. Interestingly, time spent in proximity to the fenceline by the F-P calves (about 60%) during d 1 and 2 was similar to time spent by the C-P calves in proximity (within 3 m) to their dams (about 55%) during that same period. It should be noted that the 3-m proximity criterion used represents a relatively short distance (i.e., close proximity). Calves and cows were repeatedly observed near the fenceline, well within visual contact of one another, but greater than 3 m. The 3 m criterion was chosen because it was believed to reflect a strong attraction to the other member of the cow-calf pair.

To the extent that relatively lower levels of eating/grazing and lying down and higher levels of walking and vocalizations are indicative of distress associated with the weaning process, it appeared that the calves in the three totally separated treatments experienced more distress at weaning than the calves in the F-P and C-P treatments. This difference was also reflected in rate of gain in body weight over the 10 wk following weaning. Weight gains for the F-P calves exceeded weight gains of the calves in the three totally separated treatments in all ten weekly weighings. These results suggested that the difference in weight gain between F-P and totally separated calves was largely established during the weeks immediately following weaning and that compensatory growth by the totally separated calves over 10 wk did not occur to close the gap between these groups. The C-P calves had gained more than the F-P calves at 10 wk most likely because of access to mother’s milk during most of that period. In addition, the C-P calves’ dams may have led them to better forage or encouraged more efficient grazing behavior than displayed by the other treatments.

Relative to abruptly weaning calves on pasture, cumulative postweaning weight gains were not appreciably influenced by confining weaned calves in relatively small corrals and providing high-quality alfalfa hay for feed. Preconditioning calves to hay for 10 d prior to weaning in the same corrals used in this study did not improve weight gain following weaning but appeared to have reduced indices of behavioral distress in some instances.

No physiological measures were taken in this study but other physiological research has supported the hypothesis that weaning and separation represent a stressful experience for both mother and young. Lefcourt and Elsasser (1995) reported increases in peripheral catecholamine concentrations in beef calves and epinephrine concentrations in dams in response to weaning and separation. Higher neutrophile/lymphocyte ratios have been found in abruptly weaned beef calves (J. S. Church, unpublished observations) and red deer calves (Church and Hudson, 1999) compared to gradually weaned counterparts. Interestingly, plasma cortisol concentrations have not consistently increased in response to abrupt weaning (McCall et al., 1987; Lefcourt and Elsasser, 1995; Orgeur et al., 1998).

The fences used to separate calves and cows in the F-P treatment (barbed wire overlayed with woven wire) proved effective in keeping the animals separated. In no instance did the observers see either a calf or cow make a concerted effort to break through or go over the fences used. Producers in California who have used fenceline weaning have successfully employed electrified wire either as the fence, itself, or as an outrigger 30 to 40 cm from the fence on the calf side.


    Implications
 Top
 Abstract
 Introduction
 Materials and Methods
 Results
 Discussion
 Implications
 Literature Cited
 
The results of this study support the hypothesis that fenceline contact between mother and young at weaning results in fewer indices of behavioral distress than total abrupt separation of calves from their dams and minimizes reductions in weight gain often associated with weaning. This management technique not only improves animal welfare but also may economically benefit cattle producers, especially those who sell their calves in the days or weeks following weaning.


    Footnotes
 
1 The authors would like to acknowledge V. Arias, R. Austin, T. Beaucaire, C. Bench, L. Chu, S. Flack, S. Flanders, E. Gonzales, L. Gregg, T. Hoyt, S. Jeszeck, K. Johnson, J. Kang, S. Mongold, K. Nizato, K. Simpson, J. Williams, and S. Yin for their assistance in data collection and E. Coffin and the staff of the University of California Sierra Foothill Research and Extension Center for their efforts in animal handling and care. Back

Received for publication March 8, 2002. Accepted for publication September 18, 2002.


    Literature Cited
 Top
 Abstract
 Introduction
 Materials and Methods
 Results
 Discussion
 Implications
 Literature Cited
 


Church, J. S., and R. J. Hudson. 1999. Comparison of the stress of abrupt and interval weaning of farmed wapiti calves (Cervus elaphus). Small Ruminant Res. 32:119–124.

Haigh, J. C., J. M. Stookey, P. Bowman, and C. Waltz. 1997. A comparison of weaning techniques in farmed wapiti (Cervus elaphus). Anim. Welf. 6:255–264.

Haley, D. B., J. M. Stookey, J. L. Clavelle, and J. M. Watts. 2001. The simultaneous loss of milk and maternal contact compounds distress at weaning in beef calves. Page 41 in Proc. 35th Int. Congress of the Int. Soc. of Appl. Ethology, Univ. of California, Davis.

Holland, J. L., D. S. Kronfeld, R. M. Hoffman, K. M. Greiwe-Crandell, T. L. Boyd, W. L. Cooper, and P. A. Harris. 1996. Weaning stress is affected by nutrition and weaning methods. Pferdeheilkunde 12:257–260.

Lay, Jr., D. C., T. H. Friend, R. D. Randel, C. L. Bowers, K. K. Grissom, D. A. Neuendorff, and O. C. Jenkins. 1998. Effects of restricted nursing on physiological and behavioral reactions of Brahman calves to subsequent restraint and weaning. Appl. Anim. Behav. Sci. 56:109–119.

Lefcourt, A. M., and T. H. Elsasser. 1995. Adrenal responses of Angus x Hereford cattle to the stress of weaning. J. Anim. Sci. 73:2669–2676.[Abstract]

Martin, P., and P. Bateson. 1993. Measuring Behaviour. 2nd ed. Cambridge University Press, Cambridge, UK.

McCall, C. A., G. D. Potter, and J. L. Kreider. 1985. Locomotor, vocal and other behavioural responses to varying methods of weaning foals. Appl. Anim. Behav. Sci. 14:27–35.

McCall, C. A., G. D. Potter, J. L. Kreider, and W. L. Jenkins. 1987. Physiological responses in foals weaned by abrupt or gradual methods. J. Equine Vet. Sci. 7:368–374.

Nicol, A. M. 1977. Beef cattle weaning methods. N. Z. J. Agric. 134:17–18.

Orgeur, P., S. Bernard, M. Naciri, R. Nowak, B. Schaal, and F. Levy. 1999. Psychobiological consequences of two different weaning methods in sheep. Reprod. Nutr. Dev. 39:231–244.

Orgeur, P., N. Mavric, P. Yvore, S. Bernard, R. Nowak, B. Schaal, and F. Levy. 1998. Artificial weaning in sheep: Consequences on behavioural, hormonal and immuno-pathological indicators of welfare. Appl. Anim. Behav. Sci. 58:87–103.

Pollard, J. C., and R. R. Littlejohn. 2000. Effects of management at weaning on behaviour and weight gain of farmed red deer calves. Appl. Anim. Behav. Sci. 67:151–157.[Medline]

Siegel, S., and N. J. Castellan, Jr. 1988. Nonparametric Statistics for the Behavioral Sciences. 2nd ed. McGraw-Hill, New York.

Stookey, J. M., K. S. Schwartzkopf-Fenswein, C. S. Waltz, and J. M. Watts. 1997. Effects of remote and contact weaning on behaviour and weight gain of beef calves. J. Anim. Sci. 75(Suppl. 1):157 (Abstr.).

Veissier, I., and P. LeNeindre. 1989. Weaning in calves: Its effects on social organization. Appl. Anim. Behav. Sci. 24:43–54.


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