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J. Dairy Sci. 89:2126-2131
© American Dairy Science Association, 2006.

Effects of Continuous Versus Periodic Milk Availability on Behavior and Performance of Dairy Calves

M. A. G. von Keyserlingk*,1, F. Wolf{dagger}, M. Hötzel{dagger} and D. M. Weary*

* Animal Welfare Program, University of British Columbia, Vancouver, BC, Canada V6T 1Z4
{dagger} Lab. Etologia Aplicada - Depto. de Zootecnia e Des. Rural, Universidade Federal de Santa Catarina, Florianópolis, SC, Brazil 88.040-900

1 Corresponding author: nina{at}interchange.ubc.ca


   ABSTRACT
TOP
 ABSTRACT
INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 ACKNOWLEDGEMENTS
 REFERENCES
 
Dairy calves fed milk ad libitum engage in feeding activity throughout the day and night. However, providing calves with continuous access to milk can present practical problems with milk quality, particularly during warm weather. One way to maintain milk quality in ad libitum feeding regimens is to limit the time that milk is available, but no research to date has addressed how this might affect calves. The objectives of this study were to quantify the effects of continuous access to milk (24 h/d) vs. access during only part of the day on milk feeding behavior and performance. Twenty-eight female Holstein calves were randomly assigned to treatments at 5 ± 3 d of age and monitored for 28 d. All calves had milk delivered through a teat. Treatments were 1) access to milk for 24 h/d; 2) access to milk for 2 feedings per day (each of 2 h), with water available through the teat the remainder of the day (4-h wet treatment); and, 3) access to milk for 2 feedings per day (each of 2 h), but with no water available through the teat (4-h dry treatment). Calves given 24-h access to milk consumed 11.2 ± 0.5 kg/d on average. Milk intake tended to be lower for the 4-h wet and 4-h dry treatments, averaging 10.0 ± 0.3 kg/d, with no difference between these 2 treatments in this or any other variable. Average daily gain over the course of the experiment did not vary with treatment and averaged 1.1 ± 0.04 kg/d. Calves in the 24-h treatment spent more time on the teat than those calves on the 4-h treatments, but during the 4 h/d when milk was available to all calves, calves in the 4-h treatments spent more time on the teat. These results show that individually housed calves fed milk ad libitum for only 4 h/d compensate by changing their milk feeding behavior and are able to achieve similar weight gains to animals fed milk continuously.

Key Words: milk intake • weight gain • feeding behavior


   INTRODUCTION
TOP
 ABSTRACT
 INTRODUCTION
MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 ACKNOWLEDGEMENTS
 REFERENCES
 
Under conventional management, preweaned dairy calves are commonly fed milk twice daily by bucket a total of 10% of their BW. However, recent studies have shown improvements in calf weight gains and health associated with feeding more milk. For example, Diaz et al. (2001) were able to achieve dramatically improved BW gains by feeding calves increased amounts of milk by bucket 3 times per day. Other work has shown that providing calves with continuous access to milk through a teat-based system also allows for much improved milk intakes and weight gains compared with conventionally fed calves (Appleby et al., 2001; Jasper and Weary, 2002).

Providing milk through teat-based systems also allows calves to perform natural sucking behavior (Hammell et al., 1988), increasing secretion of insulin and cholecystokinin (de Passillé et al., 1993; Lupoli et al., 2001). In addition, calves fed from an artificial teat tend not to suck on each other or on objects, unlike calves fed from a bucket (e.g., Bøe and Havrevoll, 1993; de Passillé, 2001). The reduction in nonnutritive sucking occurs partly because sucking milk from a teat increases overall feeding time, especially if a teat with a small orifice is used to reduce flow rate (Haley et al., 1998). Calves fed ad libitum by nipple spend approximately 45 min/d drinking milk, compared with just a few minutes per day for bucket-fed calves (Appleby et al., 2001).

Dairy calves fed milk ad libitum engage in feeding activity throughout the day and night (Hammell et al., 1988; Appleby et al., 2001). However, providing calves with access to milk 24 h/d can present practical problems with milk quality, particularly during warm weather. One method of avoiding problems with milk quality is to limit the time that milk is available to just a few hours a day. Although the impact of limited access to milk has not been evaluated, Appleby et al. (2001) demonstrated considerable diurnal variation in milk intake, with the largest meals occurring after delivery of fresh milk twice a day, suggesting that calves may be able to adjust to reduced access times. The primary objective of this study was to quantify the effects of ad libitum access vs. limited access to milk on milk intake, weight gain, and behavior in Holstein dairy calves. A possible additional advantage to intermittent milk feeding is that the same feeding apparatus can also be used to provide water to calves when milk is not available. Thus a secondary objective was to determine the effects of providing or not providing water through the nipple feeding system when milk was not available.


   MATERIALS AND METHODS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
RESULTS
 DISCUSSION
 ACKNOWLEDGEMENTS
 REFERENCES
 
This study was conducted at the University of British Columbia Dairy Education and Research Center in Agassiz (British Columbia, Canada). All animals were cared for according to the guidelines outlined by the Canadian Council on Animal Care (1993). Holstein female calves (n = 28), weighing 46.1 ± 1.2 kg at birth, were separated from their mothers within 24 h of birth, moved to individual pens (1.2 x 1.2 x 1.7 m), and fed 5 L of colostrum over 2 feedings. At d 1 of age calves were trained to drink from an artificial teat (Milk-Flow Peach teats, Skellerup Industries Ltd., Christchurch, New Zealand) positioned at a right angle to the wall at a height of 0.6 m. The black rubber teat was 84 mm long and 32 mm in diameter, with two 22-mm slits in the tapered end, and was attached by a polyethylene tube (8 mm diameter) to a covered plastic bucket of milk outside the pen. The milk feeding apparatuses were cleaned with hot water containing bleach and rinsed with warm water every morning. Calves also had ad libitum access to a calf starter consisting of 53.7% concentrate pellet (25% CP), 14.6% steam-rolled barley, 14.5% steam-rolled corn, 14.0% steam-flatted oats, 2.0% molasses spray, and 1.2% oil spray on a DM basis (Unifeed Ltd., Chilliwack, BC, Canada) in a bucket adjacent to the nipple, and water from a water bowl. All pens were cleaned and fresh bedding provided once a week. The composition of the starter was 47.1% DM and contained, on a DM basis, 20.4% CP, 7.0% crude fiber, 82.0% total digestible nutrients, and 1.13% Ca, and 0.68% P.

At approximately 5 d of age (5 ± 3 d of age; mean ± SD), calves were alternatively assigned to 1 of the 3 milk-feeding treatments and monitored until 32 d of age. In each condition, calves had ad libitum access to whole milk, including milk from fresh cows and from cows recovering from treatment with antibiotics, and free access to water from a bowl drinker.

Treatments were: 1) access to milk for 24 h/d (24h; n = 10); 2) access to milk for 2 daily feedings each of 2 h, with water available through the teat the remainder of the day (4hW; n = 11) and, 3) access as described in 4hW, but with no water available through the teat (4hD; n = 7). Fresh milk obtained directly from the parlor was distributed twice daily immediately following the morning and afternoon milking (0800 h ± 30 min and 1800 h ± 30 min).

Milk consumption was measured twice daily on 4 separate days each week. Body weight and starter intake were measured twice a week. Fecal scores were recorded daily for each calf according to established guidelines (Larson et al., 1977) and calves having scores greater than 3 were treated according to standard protocols developed in conjunction with the herd veterinarian. The standard protocol followed a treatment flowchart based on calf age, size, strength, temperature, and appearance of diarrhea. When treatment was required, emphasis was placed on rehydration therapy using an electrolyte solution with antibiotics used when necessary.

Feeding and lying behavior was recorded for 24 h during 2 consecutive d beginning at 32 ± 4 d of age using time-lapse video equipment. Two animals in adjacent pens were videotaped at a time using a video camera (Panasonic WV-BP330, Osaka, Japan) placed 2.6 m over the pens, a time-lapse videocassette recorder (Panasonic AG-6540), and a video multiplexer (Panasonic WJ-FS 216). Red lights (100 W), hung adjacent to the cameras, were used to facilitate recording at night. We measured the amount of time spent sucking (mouth covering at least half the nipple and head aligned no more than 45° to the nipple), time spent with the head over the water bowl (animal’s head is lowered in direction of and covers at least half of the water surface), and time spent lying down. Behaviors were scored from the videotape by scan sampling 1 frame every 20 s, providing 8,640 observations per calf.

To determine the interobserver reliability of these measures, 2 trained observers scored 3 h of tape from 16 calves independently. Estimates of sucking time, time at water bowl, and lying time from these observations were highly correlated (Pearson r = 0.98, r = 0.99, r = 1.0, respectively).

Statistical Methods
For the analysis of feeding behavior, milk and starter intake, average daily gain, and lying behavior the individual calf was considered as the experimental unit, with measures from the 2 d averaged to create 1 observation per calf per treatment. In the case of the behavioral measures data from 4 calves each from the 24h and 4hW treatment groups were not obtained, resulting in 6 and 7 calves, respectively, in each group. Analysis was by 1-way ANOVA with specified contrasts. First, we tested if the 2 limited-access treatments (4hW and 4hD) differed from one another. Because these treatments never differed we then compared the 24h treatment with the mean of the two 4-h treatments using a second contrast statement. Contrasts (1 df) were tested against an error term with 27 df (for production measures) or 19 df (for behavioral measures).


   RESULTS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
DISCUSSION
 ACKNOWLEDGEMENTS
 REFERENCES
 
Production
Calves in the 4hW and 4hD treatments drank, on average 10.0 ± 0.3 kg of milk each day, with no significant difference between these treatments (Table 1Go; P > 0.1). However, calves given 24 h access to milk tended to drink about 10% more milk daily (P = 0.07). Calves in the 3 treatment groups showed similar weight gains (P > 0.1 for both comparisons), with animals gaining on average 1.1 ± 0.04 kg/d over the study period (Table 1Go). Starter intake was negligible until calves were 14 d of age but increased during the last 2 wk of the trial for all 3 groups (data not shown). Overall, starter intakes did not differ among treatment groups (Table 1Go), with calves consuming approximately 2 kg of starter over the course of the experiment.


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  		Table 1. Mean ( ± SEM) milk and starter consumption and average daily gain (ADG) of calves provided access to milk for 24 h/d or access to milk for 4 h/d, with (4-h wet) or without (4-h dry) water available through the teat for the remainder of the day
 
Behavior
Calves given continuous access to milk spent more time on the teat than those on the 4h treatments (1.1 ± 0.14 vs. 0.8 ± 0.04 min/h, respectively). However, during the 4 h/d when milk was available to all calves, calves in the 4h treatments spent more time on the teat than did the calves given continuous access to milk (4.0 ± 0.2 vs. 2.8 ± 0.4 min/h, respectively; Figure 1aGo). There was also no change in the amount of time spent drinking from the water bowl (Figure 1bGo). Calves spent, on average, 44.7 ± 0.4 min/h lying down, with no difference among the treatment groups and no effect of treatment on the diurnal pattern of lying behavior (Figure 1cGo).


Figure 1
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  		Figure 1. Mean ( ± SE) time spent on the teat, at the water bowl, and lying down, measured in min/h. Values are shown separately for calves provided access to milk for 24 h/d (ad libitum) or access to milk for 2 daily feedings each of 2 h, with water available through the teat the remainder of the day (4-h wet), or access to milk for 2 daily feedings each of 2 h, with no water available through the teat (4-h dry).

 

   DISCUSSION
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
ACKNOWLEDGEMENTS
 REFERENCES
 
Feeding ad libitum by teat allows calves to perform natural sucking behaviors (Chua et al., 2002) but creates challenges in maintaining milk quality throughout the day, especially during warm weather. In the current study, calves that were provided access to milk for only 4 h/d (2 h after the morning and afternoon deliveries of fresh milk) still gained weight rapidly. Indeed, there were no differences in weight gains between treatment groups, despite a modest decline in milk consumption in the calves provided limited access to milk. Weight gains (about 1.1 kg/d) and milk intake (about 10 kg/d) in the current study were slightly higher than previously reported values for calves fed milk ad libitum (0.8 and 8 kg/d in Appleby et al., 2001; 0.8 and 9 kg/d in Jasper and Weary, 2002; 0.8 and 8 kg/d in Chua et al., 2002).

Our work is the first to show that calves provided access to milk for only a limited portion of the day are able to compensate by changing their milk feeding behavior. During the 4 h/d that milk was available to all calves, those on the 4h treatments spent 45% more time on the teat that did calves allowed 24-h access. Calves on the 4h treatments appeared to have no trouble predicting when milk was accessible to them, likely because of the many environmental cues associated with milk feeding (e.g., entrance of staff). In contrast, Jensen (2004) proposed that calves fed with a computer-controlled milk feeder might experience difficulties in predicting when milk is accessible to them.

Bornett et al. (2000) studied flexibility in feeding patterns of growing pigs by comparing continuous and restricted access (2 h/d) to food. During times of restricted access, pigs visited the feeder less frequently but for a longer period during each visit. However, unlike the present study in which calves were able to maintain weight gains, the restricted-access pigs failed to maintain their daily food intake and weight gains, perhaps due to limitation of gut fill. Similarly, Martinsson (1992) and Martinsson and Burstedt (1990) reported that cows given only 8 h of access to silage-based diets per day decreased their daily DMI by 6% compared with cows provided unlimited access. The ability of the young calf to compensate for restricted access to milk in the current study may in part be due to the liquid diet being consumed. Because food intake is controlled in part by gastric emptying (Rayner and Gregory, 1989), the low DM content of the milk diet should allow for a very fast rate of passage, allowing for increased rate of intake during the period that milk was available.

A number of lines of evidence suggest that calves on the 4-h treatments were not unduly affected by limiting the time of milk availability. First, the overall time on the teat and pattern of visits over 24 h was fairly similar, regardless of treatment. In a previous study, calves responded to restricted milk intake by doubling the number of visits to the feeder, likely reflecting increased feeding attempts by the hungry calves (Jensen and Holm, 2003). In contrast, the calves in the 4-h treatment groups almost never visited the teat during the 20 h/d that milk was not available. In addition, the daily patterns of lying and time spent at the water bowl were very similar in the 3 treatment groups. Indeed, lying times and patterns were well within the ranges previously reported for milk-fed calves (Hänninen et al., 2005). In combination, these results suggest that calves fed 4 h/d were not unduly disturbed by the lack of milk availability, although more research is required to document the behavioral signs of "hunger" in milk-fed calves. It would also be interesting to investigate how calves respond to more frequent delivery of fresh milk throughout the day.

Even calves with continuous access to milk consume their largest meals immediately after the delivery of fresh milk (Appleby et al., 2001). This preference for fresh milk is likely associated with milk temperature. In the current study, and in our previous work with this type of teat-feeding system, milk is delivered to calves immediately after milking and is thus close to body temperature when first delivered. When using computer-controlled calf feeders that deliver milk at a more constant temperature, visits to the nipple appear to be randomly distributed throughout the day (Hammell et al., 1988).

Although the benefits of providing ad libitum access to water are well known, a large proportion of producers fail to provide an adequate supply of water to young calves (Heinrichs et al., 1994). One advantage to intermittent milk feeding is that the same apparatus can also be used to provide water to calves when milk is not available. In the current study all calves had ad libitum access to water from a bowl drinker, but providing additional water through the nipple feeding system had no detrimental effects on production or behavior.

In conclusion, calves fed unlimited quantities of milk through a teat-feeding system can adjust their milk feeding behavior to restricted periods of milk availability. Calves spend slightly less time drinking and tend to consume less milk, but are able to maintain similar weight gains to calves provided continuous access to milk.


   ACKNOWLEDGEMENTS
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 ACKNOWLEDGEMENTS
REFERENCES
 
We thank the staff and students at the University of British Columbia’s Dairy Education and Research Center and Animal Welfare Program. We are grateful to Ina Genshein for help in running this experiment, and Cassandra Tucker, Jeff Rushen, and Anne Marie de Passillé for help planning this experiment. Funding was provided by the NSERC Research Partnership Support Program made possible by contributions from the Dairy Farmers of Canada.

Received for publication August 28, 2005. Accepted for publication January 19, 2006.


   REFERENCES
TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 ACKNOWLEDGEMENTS
 REFERENCES
 


Appleby, M. C., D. M. Weary, and B. Chua. 2001. Performance and feeding behaviour of calves on ad libitum milk from artificial teats. Appl. Anim. Behav. Sci. 74:191–201.

Bøe, K., and O. Havrevoll. 1993. Cold housing and computer-controlled milk-feeding for dairy calves: Behaviour and performance. Anim. Prod. 57:183–191.

Bornett, H. L. I., C. A. Morgan, A. B. Lawrence, and J. Mann. 2000. The flexibility of feeding patterns in individually housed pigs. Anim. Sci. 70:457–469.

Canadian Council on Animal Care. 1993. Guide to the care and use of experimental animals. Vol. 1. E. D. Olfert, B. M. Cross, and A. A. McWilliam, ed. CCAC, Ottawa, Canada.

Chua, B., E. Coenen, J. van Delen, and D. M. Weary. 2002. Effects of pair versus individual housing on the behavior and performance of dairy calves. J. Dairy Sci. 85:360–364.[Abstract]

de Passillé, A. M. B. 2001. Sucking motivation and related problems in calves. Appl. Anim. Behav. Sci. 72:175–186.[Medline]

de Passillé, A. M. B., R. J. Christopherson, and J. Rushen. 1993. Nonnutritive sucking and the postprandial secretion of insulin, CCK and gastrin in the calf. Physiol. Behav. 54:1069–1073.[Medline]

Diaz, M. C., M. E. Van Amburgh, J. M. Smith, J. M. Kelsey, and E. L. Hutlen. 2001. Composition of growth of Holstein calves fed milk replacer from birth to 105-kilogram body weight. J. Dairy Sci. 84:830–842.[Abstract]

Hammell, K. L., J. H. M. Metz, and P. Mekking. 1988. Sucking behavior of dairy calves fed milk ad libitum by bucket or teat. Appl. Anim. Behav. Sci. 20:275–285.

Haley, D. B., J. Rushen, I. J. H. Duncan, T. M. Widowski, and A. M. de Passillé. 1998. Effects of resistance to milk flow and the provision of hay on non-nutritive sucking by dairy calves. J. Dairy Sci. 81:2165–2172.[Abstract]

Hénninen, L., A. M. de Passillé, and J. Rushen. 2005. The effect of flooring type and social grouping on the rest and growth of dairy calves. Appl. Anim. Behav. Sci. 91:193–204.

Heinrichs, A. J., S. J. Wells, H. S. Hurd, G. W. Hill, and D. A. Dargatz. 1994. The national dairy heifer evaluation project – A profile of heifer management practices in the United States. J. Dairy Sci. 77:1548–1555.[Abstract]

Jasper, J., and D. M. Weary. 2002. Effects of ad libitum milk intake on dairy calves. J. Dairy Sci. 85:3054–3058.[Abstract/Free Full Text]

Jensen, M. B. 2004. Computer-controlled milk feeding of dairy calves: The effects of number of calves per feeder and number of milk portions on use of feeder and social behavior. J. Dairy Sci. 87:3428–3438.[Abstract/Free Full Text]

Jensen, M. B., and L. Holm. 2003. The effect of milk flow rate and milk allowance on feeding related behavior in dairy calves fed by computer controlled milk feeders. Appl. Anim. Behav. Sci. 82:87–100.

Larson, L. L., F. G. Owen, J. L. Albright, R. D. Appleman, R. C. Lamb, and L. D. Muller. 1977. Guidelines towards more uniformity in measuring and reporting calf experimental data. J. Dairy Sci. 60:989–991.[Abstract/Free Full Text]

Lupoli, B., B. Johansson, K. Uvnäs-Moberg, and K. Svennersten-Sjaunja. 2001. Effect of suckling on the release of oxytocin, prolactin, cortisol, gastrin, cholecystokinin, somatostatin, and insulin in dairy cows and their calves. J. Dairy Res. 68:175–187.[Medline]

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