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NUTRITION, FEEDING, AND CALVES

Effects of Resistance to Milk Flow and the Provision of Hay


on Nonnutritive Sucking by Dairy Calves1
D. B. HALEY,*,†,2 J. RUSHEN,* I.J.H. DUNCAN,†
T. M. WIDOWSKI,† and A. M. DE PASSILLÉ*
*Dairy and Swine Research and Development Centre,
Agriculture and Agri-Food Canada, Lennoxville, PQ, Canada J1M 1Z3
†Department of Animal and Poultry Science,
University of Guelph, Guelph, ON, Canada N1G 2W1

ABSTRACT INTRODUCTION
This study examined effects of resistance to milk During the milk feeding period, most veal calves
flow and the provision of hay on the duration of and many dairy replacement heifers are fed from a
nutritive sucking and subsequent nonnutritive suck- bucket, are kept individually in stalls or hutches, and
ing by dairy calves. In a series of four experiments, 12 are often tethered (9, 30). Much concern has been
male Holstein calves were individually fed milk from raised about the effect of social and behavioral restric-
an artificial teat. Resistance to milk flow was varied tion on the well-being of calves raised under such
by adjusting the orifice size within the milk supply conditions [review; (22)]. Increased interest has
tube. Using a Latin square design, each calf was fed arisen in the development of group housing systems
the same quantity of milk using four orifice sizes (one for raising veal calves and replacement heifers. One
per day for 4 consecutive d). The duration of nutritive behavioral problem that often occurs in group housing
sucking (time required to finish the milk meal) was systems is nonnutritive sucking (i.e., calves suck on
longer when calves were fed from the smallest orifice pen fixtures or on other calves without receiving any
size (0.16-cm diameter) than when calves were fed nutritive reinforcement) (19, 26, 35). The sucking of
from the largest orifice size (0.55-cm diameter). other calves (cross-sucking) is directed at the ears,
Calves compensated for resistance to milk flow in an muzzle, tail, scrotum, navel, prepuce, and the udder
attempt to maintain milk intake; however, changes in area (8, 12, 24) and is generally thought to be
sucking rate alone were probably not responsible for detrimental to calf health (28, 32). Cross-sucking can
observed differences in the rate of milk consumption result in injury to the area sucked (24); can lead to
from different orifice sizes. The duration of nonnutri- the consumption of urine ( 8 ) ; and, in extreme cases,
tive sucking (sucking of the teat following the milk may reduce BW gain (12). For these reasons, there
meal) was significantly reduced when calves were fed has been an interest in the understanding of the
from the smallest orifice compared with that when sucking behavior of calves (21).
Nonnutritive sucking of pen fixtures and of other
calves were fed from the largest orifice. Nonnutritive
calves has not been reported for calves suckling a cow.
sucking was reduced but not eliminated even with the
Thus, it has been hypothesized that nonnutritive
longest meal duration. The provision of hay to calves
sucking is done to compensate for the deprivation of
following the meal also significantly reduced the
nutritive sucking (sucking for milk), which would
amount of nonnutritive sucking. Reduction in flow
normally be performed during nursing (12, 16, 31).
rate when calves drink milk through a teat and the
In support of this hypothesis, calves fed from a bucket
provision of hay after the meal can reduce the inci-
spend more time sucking a dry teat than do calves fed
dence of nonnutritive sucking following the meal and
from an artificial teat (2, 10, 14, 17, 35), and cross-
may help to limit cross-sucking in group housing
sucking occurs less frequently when calves are fed
systems.
from an artificial teat than when calves are fed from a
( Key words: behavior, calves, nonnutritive sucking,
bucket (11, 15, 27).
cross-sucking) If nonnutritive sucking is performed to compensate
for a lack of nutritive sucking, then nonnutritive
sucking may be reduced by increasing the amount of
Received December 30, 1997. nutritive sucking that occurs. Metz ( 2 0 ) used two
Accepted April 3, 1998.
1Lennoxville Contribution Number 572. sizes of teat orifices to affect milk flow rate and found
2Corresponding author (Lennoxville address). that nonnutritive sucking tended to be reduced with a

1998 J Dairy Sci 81:2165–2172 2165


2166 HALEY ET AL.

smaller teat orifice. However, the results were not calves received 100 to 400 g/d of long alfalfa hay at
statistically analyzed. 1600 h. Water was available for ad libitum intake via
Cross-sucking and other forms of nonnutritive a self-filling float-activated water bowl, which was
sucking are most common immediately after a milk cleaned once each week.
meal (7, 19). Kilgour and Dalton ( 1 8 ) have sug-
gested that access to dry feed is a possible solution to Apparatus
sucking problems, and Hoyer and Larkin ( 1 5 ) ob-
served that sucking among calves fed from a bucket A milk delivery system was developed that allowed
was limited when calves were encouraged to eat hay calves to suck milk from an artificial teat (13). A
or chaff immediately after they finished their milk rubber teat (20 × 100 mm; wall thickness, 4 mm) was
meal; however, sucking behavior was not directly linked to a plastic container by 70 cm of clear tubing
tested. (1.6-cm diameter). The delivery tube was attached to
The objective of this research was to compare the the milk source by a connector that was equipped
effects of resistance to milk flow on meal duration and with a shut-off valve that was closed at the end of the
subsequent nonnutritive sucking. We hypothesized meal. Resistance to milk flow was altered by changing
that an increase in the resistance to milk flow and an the orifice size of a washer inserted within the milk
increase in the amount of nutritive sucking would supply tube. The teat was affixed to the pen 70 cm
reduce or eliminate nonnutritive sucking after the from the floor, and the milk source was positioned at
meal. We also examined whether the provision of hay the same height at an angle to facilitate complete
to calves would distract calves from nonnutritive drainage of the container. At this height, the milk
sucking after the meal. flow under the force of gravity through each of orifice
sizes tested was 0.55-cm diameter, 1.40 L/min;
0.39-cm diameter, 0.70 L/min; 0.27-cm diameter, 0.30
MATERIALS AND METHODS L/min; and 0.16-cm diameter, 0.13 L/min. Milk flow
under the force of gravity was approximately doubled
Calves and Housing for each successive diameter of the orifice. Regular
meals were fed using the 0.39-cm orifice, which cor-
Calves were housed and fed according to Recom- responded with the diameter of the tubing supplied
mended Codes of Practice for the Care and Handling by a commercially available teat feeding system
of Special-fed Veal Calves ( 1 ) , and all experimental (Group C.A.L.F. Supersuckler; Milk Specialties Co.,
procedures were approved by an experimental ethics Dundee, IL). For regular feedings, teats were re-
committee at the Dairy and Swine Research and De- moved 10 to 15 min after the calf finished the milk.
velopment Centre in compliance with the guidelines
of the Canadian Council on Animal Care. Twelve
male Holstein calves separated from their dam within Behavioral Observations
24 h of birth were housed individually in pens (2.1 × Video cameras (Panasonic WV-BL200; Panasonic
1.85 m ) that permitted visual, auditory, olfactory, Canada Inc., Mississauga, ON, Canada) were posi-
and limited physical contact with at least one adja- tioned to record a close, side view of each calf as it
cent neighbor. Calves were kept on a Pasture Mat sucked the teat. Output from the 12 cameras was
flooring (Promat Ltd., Seaforth, ON, Canada) and taped simultaneously, each at a rate of 15 frames/s
were provided with fresh wood shavings for bedding using three time-lapse video cassette recorders
each day. All pens were in the same artificially venti- (Panasonic AG-6730; Panasonic Canada Inc.) and
lated room with a preset minimum temperature of three, four-way video splitters (Uniplex Sprite Video
20°C and a maximum allowable variation of 8°C per Multiplexer; Dedicated Microcomputers Ltd., Reston,
24-h period. Artificial lighting was provided on a VA). Videotape recording began just before feeding
12-h photoperiod from 0600 to 1800 h. and ended 25 min after the calf finished the milk. All
tests were performed during the morning feeding
Feeding period. As each calf finished the milk, the attending
technician made a signal in view of the camera to
Calves were individually fed whole milk via a rub- indicate the end of the meal. The shut-off valve was
ber teat. The daily quantity was equal to 10% of the subsequently closed to prevent calves from sucking
mean BW of the group and was divided into two equal air or tasting any droplets of milk left in the supply
portions fed at 0745 and 1545 h at a mean tempera- tube. For experimental meals, the teats were left in
ture of 30°C (27 to 33°C). Depending on their age, place for 25 min after the milk was consumed.

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NONNUTRITIVE SUCKING BY DAIRY CALVES 2167

Videotapes were analyzed by continuous observa-


tion. The behavior of each calf was observed from the
beginning of the meal until 20 min after the meal
ended. Most nonnutritive sucking occurs during the
10 min following a milk meal ( 5 ) .
The following behavior patterns were recorded.
Nutritive sucking. Nutritive sucking was
recorded during the meal when the calf was sucking
on the teat and ingesting the milk. Nutritive sucking
was operationally defined as the rhythmic opening
and closing movements of the mouth; the tongue was
cupped around the bottom portion of the teat. The
teat had to be straight and oriented along its length
within the mouth of the calf (i.e., not folded or bent
over). Sucking had to last at least 2 s to be recorded.
Nonnutritive sucking. Nonnutritive sucking was
based on the same criteria as nutritive sucking but Figure 1. Mean flow rate of milk during sucking by calves ( ⁄)
compared with the flow rate through the same orifice size at
was scored after the meal ended, when calves were no gravity ( ◊) . Flow rate during sucking by calves is also expressed as
longer sucking for milk. a percentage of gravity flow rate (13). Dimensions of orifice sizes:
Manipulate. Any other oral behavior directed at large, 0.55-cm diameter; medium, 0.39-cm diameter; small,
0.27-cm diameter; and very small, 0.16-cm diameter.
the teat (e.g., chewing or licking the teat) that was
not classified as sucking behavior was categorized as
manipulative behavior. Experiment 3: Effects of Resistance to Milk Flow
Recumbent. Recumbent behavior was recorded on the Sucking Rate of Calves
when a calf was in any lying position.
Jaw movements. For the purpose of examining Results from preliminary studies ( 1 3 ) showed
sucking rate, jaw movements were recorded. A com- that, to some degree, calves could adjust their rate of
plete open and close cycle of the mouth of the calf was milk intake to compensate for resistance to milk flow
scored as one jaw movement. (Figure 1). The objective of this experiment was to
determine whether differences in the rate of milk
intake with two different orifice sizes were due to
Experiments 1 and 2: Meal Duration changes in the sucking rate of the calves as measured
and Nonnutritive Sucking by the frequency of jaw movements.
For this experiment, the standard feeding system
Two experiments were designed to test the effects was modified by replacing the simple connector piece
of resistance to milk flow on the duration of nutritive at the base of the milk container with a Y-shaped
and nonnutritive sucking. connector. Tubes exiting the Y-shaped connector
In Experiment 1, 12 calves were tested at a mean merged into one common tube to which the teat was
age of 8 ± 0.4 wk. Calves were blocked by weight and attached (Figure 2). Washers with different orifice
were fed 3.4 L of whole milk from rubber teats with a sizes were inserted on either side of the Y-shaped
0.16-cm or 0.55-cm diameter. Using a crossover de- connector. Resistance to milk flow was increased or
sign, calves randomly received milk from both decreased by directing the flow of milk during the
orifices, one per day on 2 consecutive d. meal through one side of the Y connector and then the
In Experiment 2, we tested four orifice sizes to other. The total length of tubing remained 70 cm.
examine the effects of very slight differences in milk Ten calves were blocked by weight and tested at a
mean age of 15 wk. At 30-s intervals during the meal,
flow resistance. The same 12 calves from Experiment
milk flow was alternated back and forth through
1 were tested at a mean age of 13 wk and were fed 5.0
orifices with 0.55 and 0.39-cm diameter, which con-
L of milk from four orifice sizes: 0.55, 0.39, 0.27, and stituted a 30% change in orifice size. Resistance to
0.16 cm. A repeated Latin square design was used, milk flow was manipulated during the first 5 min of
and calves randomly received milk from each orifice the meal, and calves always began the meal by suck-
size, a different size each morning over 4 consecutive ing milk through the 0.55-cm orifice. Sucking rates
d. Calves were blocked by BW, generating three 4 × 4 were calculated by counting the number of jaw move-
Latin squares. ments during each 30-s interval. Sucking rates during

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2168 HALEY ET AL.

Figure 2. Diagram showing the modified feeding system used to manipulate milk flow rate during the meal.

periods of different resistance were compared. For of the data was examined by the univariate proce-
practical purposes, jaw movements were only counted dure, and data were log-transformed. Untransformed
during periods of continuous sucking for 10 s or data are presented in the results section and in the
longer. To maximize the quality of the videotaped figures as means ( ±SE). Effects of milk flow
image, each calf was recorded at 60 frames/s. Be- resistance and the provision of hay on the duration of
havior following the meal was not examined in this various behavior patterns were analyzed by ANOVA
experiment. using the general linear models procedure with the
following factors: treatment, block, day, calf within
Experiment 4: Effects of the Provision of Hay block, and the interaction of treatment and block.
on Nonnutritive Sucking Following the meal, more than 90% of the nonnutri-
tive sucking occurred during the first half of the
The objective of this experiment was to test 20-min observation period. Statistical results were
whether the provision of hay after the milk meal the same for all behavior patterns considered,
affected the amount of nonnutritive sucking. whether 10 or 20 min were analyzed; therefore, we
The mean age of the 12 calves during this experi- have only reported results from the initial 10 min
ment was 14.5 wk. On 2 consecutive d, calves sucked following the meal.
6.2 L of milk (11% of group mean BW) through the The mean sucking rate of calves during consecutive
0.39-cm orifice. Treatments were administered as 30-s periods was analyzed by ANOVA using a
calves finished their milk. Treatments were 1 ) ap- repeated measures design with the following factors:
proximately 50 g of hay placed inside the feeder and flow rate, block, calf within block, and the interaction
immediately removed (sham fed) or 2 ) approxi-
of flow rate and block.
mately 50 g of hay placed in the feeder and left for the
Multiple mean comparisons were made using
calf to consume (hay fed). Using a crossover design
Tukey’s test.
and blocking by BW, all calves randomly received
both treatments, one each day on consecutive days.
All calves were sham fed the evening before the test RESULTS
to provide some experience with the general ex-
perimental procedure. Experiments 1 and 2: Meal Duration
and Nonnutritive Sucking
Statistical Analysis
All sucking was directed at the teat, and little, if
Statistical analyses were performed using the SAS/ any, obvious sucking of objects or other calves oc-
STAT for Windows software package (25). Normality curred. In Experiment 1, there was an overall effect of

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NONNUTRITIVE SUCKING BY DAIRY CALVES 2169

orifice size on the duration of both nutritive sucking


for milk ( P < 0.001) and nonnutritive sucking of the
teat after the meal ( P < 0.01). Compared with the
time necessary to finish the milk when the
0.55-cm orifice was used (3.23 ± 0.28 min), calves fed
from the 0.16-cm orifice took longer to finish the milk
(9.84 ± 0.89 min), and the duration of nonnutritive
sucking from the 0.16-cm orifice (3.44 ± 0.40 min)
was shorter than that from the 0.55-cm orifice (5.08 ±
0.49 min). When calves sucked from the 0.16-cm
orifice, the duration of sucking to finish the meal
(13.28 min) surpassed the combined total sucking
time (nutritive plus nonnutritive) when calves
sucked from the 0.55-cm orifice (8.31 min). Orifice
size did not affect the amount of time calves spent
manipulating the teat following the meal (0.55-cm
Figure 4. Mean duration of nonnutritive sucking (i.e., sucking of
orifice = 2.52 ± 0.53 min; 0.16-cm orifice = 2.58 ± 0.44 the teat following the meal when no milk was available) by calves
min; P > 0.10). fed from the large (0.55-cm diameter), medium (0.39-cm di-
In Experiment 2, there was an overall effect of milk ameter), small (0.27-cm diameter), and very small (0.16-cm di-
ameter) orifice sizes. Bars with different superscripts (a, b ) differ
flow resistance on meal duration ( P < 0.0001). Dura- ( P < 0.05) according to Tukey’s test.
tion of nutritive sucking was significantly longer
when calves were fed from the 0.16-cm orifice than
when calves were fed from the 0.27-, 0.39-, and
0.55-cm orifices (Figure 3). Milk flow resistance had cant differences in nonnutritive sucking among the
an overall effect on nonnutritive sucking of the teat orifice sizes were detected. As in Experiment 1, when
after the meal ( P < 0.0001). Less nonnutritive suck- calves were fed from the 0.16-cm orifice, the duration
ing was recorded following meals fed from the of nutritive sucking (14.45 min) surpassed the total
0.16-cm orifice than that following meals fed from the sucking time (nutritive plus nonnutritive) recorded
other three orifice sizes (Figure 4). No other signifi- for calves fed from each of the other orifice sizes
(10.12, 9.90, and 10.44 min for the 0.55-, 0.39-, and
0.27-cm orifices, respectively). Orifice size had no
effect on the amount of time calves spent manipulat-
ing the teat following the meal (0.55-cm diameter =
2.71 ± 0.64 min, 0.39-cm diameter = 2.56 ± 0.67 min,
0.27-cm diameter = 2.81 ± 0.54 min, and 0.16-cm
diameter = 2.18 ± 0.48 min; P > 0.10). Calves spent
more time ( P < 0.05) recumbent following meals fed
from the 0.16-cm orifice (1.27 ± 0.39 min) compared
with the time spent lying following meals fed from the
0.55-cm orifice ( 0 min). Size of the orifice used for
feeding had no effect on other behavior patterns.

Experiment 3: Effects of Resistance


to Milk Flow on Sucking Rate
A significant overall difference was detected in the
sucking rate of calves when milk was fed through two
different orifice sizes ( P < 0.01). When there was less
Figure 3. Mean ( ±SE) duration of nutritive sucking of milk by resistance to milk flow with the 0.55-cm orifice, the
calves fed from the large (0.55-cm diameter), medium (0.39-cm sucking rate of the calves was higher (2.30 ± 0.03 jaw
diameter), small (0.27-cm diameter), and very small (0.16-cm
diameter) orifice sizes. Bars with different superscripts (a, b ) movements/s) than that when milk was fed through
differ ( P < 0.05) according to Tukey’s test. the 0.37-cm orifice (2.27 ± 0.03 jaw movements/s).

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2170 HALEY ET AL.

Experiment 4: Effects of the Provision of Hay showed a consistent reduction; oral manipulation of
on Nonnutritive Sucking the teat following the meal was the same for different
meal durations. Nonetheless, calves spent signifi-
Provision of hay to calves following the meal had
cantly more time in a recumbent position when fed
an overall effect on nonnutritive sucking ( P < 0.01);
from the 0.16-cm orifice, and the same muscles may
nonnutritive sucking was reduced when hay was fed
not be used for sucking and manipulation of the teat.
(0.51 ± 0.15 min) compared with that when hay was
Calves were sensitive to changes in milk flow rate
sham fed (1.88 ± 0.04 min). No differences in the
and were able, in part, to compensate for increased
amount of time that calves spent performing other
resistance to milk flow to maintain similar rates of
behavior patterns were observed ( P > 0.10).
milk intake. During preliminary studies (13), the
shortest meal duration was achieved with the
DISCUSSION 0.39-cm orifice, and a further increase in orifice size
A longer duration of sucking during the meal did not reduce meal duration. When a very large
resulted in a reduction in subsequent nonnutritive orifice was used (1.02-cm diameter), calves sucked
sucking, but nonnutritive sucking was not eliminated. the milk at a speed equal to only 29% of the natural
Calves compensated for resistance to milk flow, in flow rate through the same orifice at gravity. The
part, by changing their sucking rate. In addition, calves were clearly slowing down the speed of milk
calves performed less nonnutritive sucking when hay flow. Rate of milk intake by the calves appeared to
was fed following the meal. reach a plateau at approximately 1 L/min, which may
Reduced nonnutritive sucking following longer reflect a limit on the amount of milk that calves can
periods of nutritive sucking provided evidence in sup- swallow. When further resistance to milk flow was
port of the hypothesis that nonnutritive sucking is applied using an orifice size of 0.08 cm, calves some-
performed to compensate for a lack of nutritive suck- times stopped sucking completely although milk was
ing (7, 16, 31, 33). Rushen and De Passillé ( 2 3 ) have still available.
found that prior nonnutritive sucking reduced subse- Wolff ( 3 4 ) found that calves had the same sucking
quent nonnutritive sucking more than the ingestion rate on an artificial teat whether or not they were
of milk. receiving milk (2.1 to 2.5 sucks/s). We found similar
The reduction in nonnutritive sucking may be due sucking rates, and, although analysis showed a
in part to the physiological consequences of sucking higher sucking rate with the 0.55-cm orifice, such a
behavior. Wise and LaMaster ( 3 ) suggested that small difference may not be of much biological sig-
sucking enhances some physiological processes as- nificance. The small difference in sucking rate ob-
sociated with the digestion of milk. Calves allowed to served in the present study ( 1 % ) probably cannot
suck a dry teat after drinking their milk from a alone explain the difference in the rate of milk con-
bucket have higher concentrations of insulin and sumption (20%) between the two orifice sizes. Two
cholecystokinin in their blood ( 5 ) . These metabolic other factors that might affect the rate of milk intake
hormones are thought to be important in satiety in include the degree of pressure within the oral cavity
addition to other factors such as stomach distension and the position or movement of the tongue. With
or absorption of digested nutrients (23). Prolonging greater resistance to milk flow, calves may reduce
the period of nutritive sucking may allow these hor- their sucking rate, applying more force in the same
mones to reach higher concentrations in the blood amount of time to overcome resistance to milk flow.
before the calf stops ingesting milk. Digestion of the Calves might have been sucking harder and perhaps
milk may reduce sucking motivation by acting as using different muscles in the mouth.
negative feedback via stomach distension or circulat- Although nonnutritive sucking after the meal was
ing hormone concentrations, or both. If the amount of reduced, it was not eliminated. When calves were fed
time for the digestion of milk or the concentration of from the 0.16-cm orifice, the duration of nutritive
blood hormones is important in regulating nonnutri- sucking surpassed the total sucking duration (nutri-
tive sucking, an increase in the duration of the meal, tive plus nonnutritive) for calves fed from each of the
regardless of feeding method, may be useful in reduc- other three orifice sizes. These results suggest that
ing nonnutritive sucking. calves may be motivated to perform some nonnutri-
It could be argued that, following meals fed from tive sucking at the end of a milk meal, regardless of
the 0.16-cm orifice, the sucking muscles of the calves meal duration. De Passillé et al. ( 6 ) found that the
were too fatigued to allow for much nonnutritive ingestion of milk stimulates sucking by calves. The
sucking. However, only actual sucking of the teat ingestion of milk during the meal might have continu-

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NONNUTRITIVE SUCKING BY DAIRY CALVES 2171

ally stimulated the calves to suck even at the end of Thibeault, Richard Lanctôt, and other staff at the
the longest nutritive period. Most milk-fed calves are Dairy Centre for their cooperation and for looking
likely motivated to perform some nonnutritive suck- after the calves. Essential technical assistance was
ing, including calves raised with their dam. However, provided by Dominique Lévesque, Sylvie Provencher,
performance of nonnutritive sucking for calves housed and Marjolaine St-Louis.
with the dam may be directed at the teats instead of
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