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Since survival of the young mammals depends on sucking success, it is assumed that

sucking motivation must be strong and that sucking deprivation would result in
frustration which could have a negative impact on their welfare. This concern as well as
that regarding cross-sucking between calves and intersucking between older animals
has stimulated research on the motivation of calves for non-nutritive sucking. Nonnutritive sucking is elicited by the ingestion of milk, and the lactose concentration in
milk, rather than that of fat or protein, is the main factor stimulating non-nutritive
sucking. Every time a calf drinks milk, it is stimulated to suck and deprivation of sucking
may interfere with digestive processes or satiety. To understand the behaviour of the
calf during nutritive sucking, we examined the effects of milk flow on calves sucking
and butting an artificial feeding system. Slowing and stopping the flow rate stimulates
butting and can lengthen the duration of sucking for the meal. It also stimulates the calf
to switch teats when a second teat is available. These findings are well correlated to
our observations of the calf suckling a cow in experimental manipulations. The duration
of the meal is not correlated with milk intake, but it is influenced by milk availability and
how hungry the calf is. Furthermore, calves will butt more often when there is less milk
available in the dams udder, presumably because milk flow is slower. Consequently, it
is the occurrence of butting rather than sucking duration that would be a good indicator
of milk intake when the calf is suckling the dam. Finally, we recommend a combination
of slower milk flow and hay feeding and the provision of a non-nutritive artificial teat to
reduce the occurrence of cross-sucking following a milk meal.

1. Introduction
Because of the importance of sucking behaviour for young calves reared for
meat production or milk production, it is necessary to understand the underlying
motivation and consequences of this behaviour. Understanding sucking
behaviour should help us control cross-sucking (sucking of ears, tails, prepuce
and other body parts) and inter-sucking (milk stealing and udder sucking in
older animals), develop calf milk-feeding systems that are better adapted to the
calf, and develop management systems that ensure good productivity in
cow/calf rearing systems.
The issue of behavioural deprivation must also be addressed for artificially
reared calves. The issue of behavioural deprivation is central to animal welfare
(Hughes and Duncan, 1988), since in most modern farms, the animals cannot
perform many of the behaviours regularly seen in less restrictive environments.
Animal welfare groups see such behavioural deprivation as a major problem
with intensive animal husbandry, but in order to know whether behavioural
deprivation reduces welfare, we must understand the nature of the motivation
underlying each particular behaviour (Hughes and Duncan, 1988). Calves
reared separately from their mother are often fed by bucket and can only suck
at objects in the pen or at pen mates to satisfy their sucking motivation. When
calves can suckle their dam, they are rarely reported to suck on other calves
(Krohn et al., 1999 and Schuch et al., 1999), but cross-sucking and intersucking
can occur after weaning off the cow (Keil et al., 2000). In order to determine

how much sucking deprivation affects the calves welfare, we need to


understand more about the causes and consequences of sucking.
In this paper, we summarise the research we have done in four areas: (1)
identifying the factors that motivate calves to perform non-nutritive sucking; (2)
identifying the factors that motivate calves to perform sucking and non-nutritive
sucking on a teat feeding system and using this information to understand the
consequences of preventing calves from carrying out normal sucking behaviour
and to control cross-sucking in calves kept in groups; (3) documenting the
effects of the calf suckling (nursing) the dam on the dams physiology as well as
on the calfs growth; (4) examining the motivation for sucking and butting when
the calf is suckling the cow under different conditions of milk availability and calf
hunger level.

2. Non-nutritive sucking by calves


Individual housing for veal calves has been criticised by animal welfare groups.
However, the research on veal calf housing shows that adoption of alternative
housing systems (especially those where the animals are kept in groups) may
not automatically improve animal welfare (Rushen, 1994). Different types of
housing systems tend to produce different welfare problems and changing
housing systems often results in switching from one set of welfare problems to
another. For example, although group housing systems for calves enable the
calves to show a wide range of behaviours, the incidence of disease and
mortality rates are often higher than for individual housing (Rushen, 1994). One
solution is to ask how each sort of housing system can be improved. One factor
that might increase disease transmission in grouped calves is the cross-sucking
that occurs between calves. Dairy producers in particular are hesitant to keep
heifers in groups because of this cross-sucking, which many believe also leads
to inter-sucking and milk stealing (Spinka, 1992 and Keil et al., 2000).
Intersucking is reported to occur in 111% of heifers and cows (Wood et al.,
1967, Keil et al., 2000 and Spinka, 1992). When these animals are identified or
suspected, they are usually fitted with a nose-ring to control this behaviour,
because it is believed to cause udder malformations and mastitis (Keil et al.,
2000). However, this technique has been criticised from an animal welfare point
of view (Keil et al., 2000). In order to improve group-housing systems to control
these abnormal/undesirable behaviours, we need to understand the factors that
cause calves to suck.
Sucking bouts on the dam occur 410 times a day depending on the age of the
calf and last on average 710 min (Hafez and Lineweaver, 1968, Ewbank,
1969, Odde et al., 1985, Day et al., 1987, Lidfors and Jensen, 1988 and Spinka

and Illmann, 1992). The main functional goal of sucking behaviour seems
obvious: to obtain milk. However, when young ruminants are raised separately
from their mothers, they suck each other and at parts of their pens despite
apparently adequate nutrition. It has often been suggested that this non-nutritive
sucking behaviour is due to an increased motivation for sucking behaviour
resulting from the lack of opportunities to suck the mother (Sambraus, 1985).
Does the calf need to suck and if so when?
In a series of studies (reviewed in de Passill and Rushen, 1997), we examined
the factors that can cause or inhibit non-nutritive sucking by calves. To test
experimentally the calves motivation to suck, we developed a model consisting
of an artificial, dry rubber teat. The calves never received milk through the teat,
which allowed us to separate the effects of sucking behaviour itself from the
effects of milk ingestion. Non-nutritive sucking at such artificial teats can be
readily elicited in young calves (e.g. Hammell et al., 1988 and de Passille et al.,
1992) and is performed in the typical sucking positions. As well as sucking, the
calves will often butt at the teat, which is a normal part of their suckling
behaviour (Hafez and Lineweaver, 1968 and Lidfors et al., 1994). Most of the
suckling and butting occurs during the first 10 min following a milk meal (Fig. 1).
In contrast to what one would expect if non-nutritive sucking was stimulated by
hunger, non-nutritive sucking is far more common immediately after the meal
than before (de Passill et al., 1992) and disappears after weaning calves off
milk (Lidfors, 1993). Non-nutritive sucking is slightly higher in calves receiving a
lower ration of milk (Rushen and de Passill, 1995) indicating that non-nutritive
sucking is not completely independent of food intake in the longer term. It is
important to note that we are looking at the well fed calfs motivation for nonnutritive sucking. This does not mean that a very hungry calf would not be
motivated to perform sucking on all objects and other calves if it was not able to
obtain milk. Nevertheless, halving the amount of milk the calves drink during a
meal (Rushen and de Passill, 1995) does not increase the amount of nonnutritive sucking that occurs after the meal, which suggests that neither the
amount of milk in the stomach nor the oral sensations from ingesting milk inhibit
non-nutritive sucking.
In fact, our results suggest that non-nutritive sucking is elicited, rather than
reduced by the ingestion of milk: de Passill et al. (1992) found that some nonnutritive sucking occurred when calves did not drink milk at meal time or drank
water, but this was considerably less than when the calves drank milk. Simply
injecting small quantities of milk into the mouth of the calf is sufficient to

stimulate considerable sucking (Rushen and de Passill, 1995). Much less


sucking is found when the calves taste either water or a suspension of grain
and the amount of non-nutritive sucking increases as the concentration of milk
replacer increases (de Passill et al., 1997a) (Fig. 2). This suggests that it is
specifically the taste of milk that is important in eliciting sucking. In a number of
experiments, we have altered the concentration of each of the main ingredients
of reconstituted milk. Changes in the concentrations of butter fat, casein and
lactoserum proteins have little effect on the amount of non-nutritive sucking that
occurs. However, increases in lactose concentrations increase non-nutritive
sucking, while reductions in lactose concentrations have the opposite effect.
Converting lactose to glucose and galactose has little effect suggesting that it is
the concentration of sugars rather than lactose per se which is the main factor
(de Passill and Rushen, 1998).
If the ingestion of milk does not reduce sucking motivation, what does? Rushen
and de Passill (1995) examined whether the performance of sucking behaviour
itself reduces the motivation to suck. We gave calves a small portion of milk in a
bucket and gave half of the calves a dry teat to suck. We then gave all the
calves another small portion of milk and gave them all teats. When the calves
sucked the teat after drinking the first portion of milk, the duration of sucking
following the second portion was lower than that following the first portion. This
did not occur if the calves did not receive the teat after the first portion of milk.
This suggests that the performance of sucking behaviour itself is more effective
in reducing the underlying motivation than is the actual ingestion of milk.
In order to examine the duration of this inhibitory effect, we gave the teat to only
half the calves immediately after a meal. Forty minutes later, we gave all of the
calves a teat, but little sucking occurred and this was not higher in the calves
that could not suck immediately after the meal (Rushen and de Passill,
1995). de Passill et al. (1992)found that the sucking motivation that was
elicited by the ingestion of the milk waned during the 10 min following the meal.
These results suggest that the elicited motivation to suck wanes fairly quickly
even in the absence of opportunities to suck. This supports the observations
that tying up a calf for 10 min after the milk meal reduces the occurrence of
cross-sucking in group-reared milk-fed calves (Graf et al., 1989).
Sucking by calves is often claimed as a behavioural need (Sambraus,
1985 and de Wilt, 1985). However, the impression often given in discussions of
behavioural needs is of animals being continuously in a state of high motivation
(and hence, frustration) unless they are able to perform the behaviour. Our
results provide little support that this view can apply to sucking behaviour. It

seems likely that until the calves drink milk, they are not motivated to suck
regardless of sucking deprivation. Moreover, the sucking motivation elicited by
ingesting milk soon decays regardless of whether any sucking occurs.
Nevertheless, the performance of this non-nutritive sucking following the milk
meal may have a physiological consequence that may be of value to the animal
and in this respect, sucking may be important for calves.
2.1. Physiological consequences of non-nutritive sucking
Our results show that non-nutritive sucking by young calves reduces sucking
motivation. There is some evidence that performance of normal feeding
behaviour in a number of species contributes to satiety, even if it does not result
in food ingestion (e.g. Toates, 1986). It is not yet known whether performance of
such behaviour achieves this effect by influencing the underlying physiological
processes involved in satiety.
de Passill et al. (1993) found that insulin and cholecystokinin (CCK) in the
hepatic portal vein after the meal were higher when the calves sucked a dry teat
after the meal. The increase in CCK and insulin concentrations was positively
correlated with the duration of sucking, but not with the duration of other oral
behaviour directed at the teat. This suggests that actual sucking behaviour is
important rather than just the extra sensory stimulation from having the teat in
the mouth. The physiological mechanism underlying the effect is unclear,
although increased vagal stimulation may be involved (Uvns-Moberg, 1983).
The possible satiety effects and the widespread metabolic effects of insulin and
CCK mean that deprivation of sucking behaviour cannot be assumed to be
inconsequential for animal well-being and growth even if this does not affect
nutrient intake.
2.2. Can these results help us control cross-sucking?
From these results we would predict that cross-sucking in calves would tend to
occur immediately after meals of milk rather than at other times, and that it
would be reduced if the calves could suck a teat immediately after the meal.
Most cross-sucking between calves has been reported to occur following
feeding (Lidfors, 1993, de Passille and Caza, 1997 and Veissier et al., 1998)
and very little occurs after calves are weaned from milk (Lidfors, 1993). de
Passill and Caza (1997) examined milk fed veal calves that were housed in
groups and drank milk from a bucket. Some of the groups were allowed to suck
a dry rubber teat after each milk meal. They found that most cross-sucking did
occur immediately after each milk meal, as predicted (Fig. 3). Furthermore, the
calves that were offered rubber teats after each meal preferred sucking the teat

to sucking the other calves in the pen, so the teat had a marked effect on crosssucking between calves by reducing its occurrence by more than 75% (Fig.
3). Jung and Lidfors (1999) report similar results. These results support the
findings that calves are motivated to do non-nutritive sucking after the milk meal
and that, in a production setting, satisfaction of this motivation will reduce the
occurrence of cross-sucking.
Little is known about the ontogeny of intersucking. It is thought that intersucking
evolves from cross-sucking (Spinka, 1992). If cross-sucking was controlled we
would expect no intersucking. Sucking in itself or via the hormones secreted
with sucking, CCK for example, has been shown to be important in the
development of bonding between the newborn and its dam (Nowak et al.,
1997 and Nowak et al., 1999). When this bonding does not occur with the cow,
perhaps because the calf did not suckle the dam for long enough, the calf may
bond to something else and this may lead to intersucking. In support of this
idea, Dobrecni and Juhs (1999) report a lower percent of intersuckers in
herds where calves are allowed to suckle their dam for 5 days or more rather
than one day or less. Perhaps, five days of suckling permitted a bond to form
between the calf and the dam and the dam and the abrupt separation from the
cow at that time caused the bond to be broken in an acute way. Spinka
(1992) has suggested that to avoid or stop intersucking, the heifer doing the
intersucking must be rejected as is done by the dam in a natural weaning
situation or by the heifer being intersucked when the intersucking heifer is fitted
with a nose-ring. It would be interesting to further examine the role of bonding of
the calf on the cow, animal or object being sucked and the effects of braking this
bond on the occurrence of intersucking.

3. Motivation to suck and butt when on a teat feeding system


If non-nutritive sucking is performed in order to compensate for the inability to
perform normal nutritive sucking, could we reduce its occurrence by increasing
the opportunity to suck for milk? Cross-sucking occurs less frequently when
calves are fed from an artificial teat than when calves are fed from a bucket
(Graf et al., 1989, Hoyer and Larkin, 1954 and Szucs et al., 1983). Calves fed
from a bucket have been reported to spend more time sucking a dry teat than
do calves fed from an artificial teat (Hammell et al., 1988). When calves take
longer to suck their milk from a teat bucket, they do less non-nutritive sucking
following the meal (Hepola et al., 1999). Furthermore, Metz (1984)reported
reduced non-nutritive sucking when calves obtained milk from an artificial teat
that had a smaller orifice so that the milk flow rate was reduced, and hence,

drinking time was longer. We examined how nutritive sucking could reduce
post-prandial non-nutritive sucking.
To do this, we devised a nutritive teat feeding system (Haley et al.,
1998a and Haley et al., 1998b) where we could control the flow rate of the milk
delivery such that we could lengthen the duration of the meal, and
consequently, the length of time the calves sucked to obtain their milk. We used
washers with different size orifices so that we could control the flow rate of the
milk. The first thing we found was that the flow rate when the calves sucked was
not related linearly to the size of the orifice. At each step, we doubled the size of
the orifice and effectively doubled the flow rate when milk was flowing freely
under gravity. However, when the calves were sucking, the flow rate did not
increase linearly. Calves appeared to be able to adjust their rate of sucking so
that when the size of the orifice was large, the calves slowed down the flow rate
compared to what would have been found had the milk simply flowed under the
influence of gravity. In contrast, when the size of the orifice was reduced, calves
appeared to be able to increase the rate of consumption so that the flow rate
during sucking was much greater than occurred under gravity (Fig. 4). For most
orifice sizes, the calves sucked at about 1 l/min, which may reflect the optimum
flow rate for calves; only when the size of the orifice was very small did the flow
rate during sucking drop much below this. Hence, for the three largest orifices,
we were not able to substantially increase the duration of non-nutritive sucking,
and consequently, the duration of non-nutritive sucking was not greatly reduced
(Fig. 5). However, when the size of the orifice was very small, the flow rate
during sucking was very low, the duration of nutritive sucking more than doubled
and non-nutritive sucking was reduced by half (Haley et al., 1998a). However,
with the very small orifice we found that the duration of nutritive sucking alone
greatly exceeded the combined duration of nutritive and non-nutritive sucking
for the other sizes of the orifice. Despite this, some non-nutritive sucking still
occurred. Therefore, increasing the duration of nutritive sucking is an effective
way of reducing non-nutritive sucking following a milk meal. However, making
the meal longer may in itself reduce the duration of non-nutritive sucking but
may not eliminate it entirely (Haley et al., 1998a). hrberg and Lidfors
(1999) have reported that cross-sucking was reduced as effectively by a slow
flow meal in a bucket (10 min) as by access to a floating nipple during the meal
and for 15 min post milk ingestion. But once again, the authors did not report a
complete elimination of cross-sucking with these treatments. These results
suggest that the non-nutritive sucking after a meal is not done purely to
compensate for a reduced duration of nutritive sucking. The ingestion of milk

during the meal may have continually stimulated the calves to suck even at the
end of the longest milk meal.
When calves are allowed non-nutritive sucking at the milk-feeding station, time
in the station doubles to 12 min on average and cross-sucking is reduced about
10-fold (Schuch et al., 1999). All these results demonstrate that the calfs
motivation to suck after a meal is very high and that providing the opportunity
for non-nutritive sucking in an experimental or in a production setting will satisfy
the calfs sucking motivation and reduce the occurrence of cross-sucking. It is
clear that the sucking motivation following a milk meal is strong and consistent.
We have also examined some other ways of reducing non-nutritive sucking
after a meal. We have demonstrated that the motivation to suck wanes within
1015 min post-meal as satiety mechanisms set in (de Passill et al.,
1992). Graf et al. (1989) have reported that 10 min of tying up the calves after
their milk meal was sufficient to control the occurrence of cross-sucking.
However, this last solution may be less appropriate as it will deprive the calf of
the physiological effects of non-nutritive sucking (de Passill et al., 1993).
Providing water through the teat at the end of the meal also reduced nonnutritive sucking (Gaboury and de Passill, 1997). We also found that providing
hay at the end of milk meal stimulated the calf to eat hay rather than perform
non-nutritive sucking and this also lead to a reduction in the performance of
non-nutritive sucking at the end of the milk meal (Haley et al., 1998a). However,
neither of these treatments eliminated the non-nutritive sucking. These results
indicate that a combination of slower milk flow, water feeding through the teat at
the end of the milk meal, hay feeding at the end of the milk meal and the
provision of a non-nutritive artificial teat will reduce the motivation for nonnutritive sucking at the end of a meal and, consequently, would be ways of
reducing the occurrence of cross-sucking following milk meals.
Finally, we also examined the calves behaviour on the teat feeding system to
determine factors that stimulate the calves to butt and change teats so that we
may be able to interpret the calf behaviour while nursing a cow. A stoppage or a
drop in milk flow and a slow flow rate stimulate butting (Haley et al., 1998b) and
teat switching (de Passill and Gaboury, 2000) as well as some pacing by the
calf. Calves are sensitive to milk flow rate and can adjust their sucking
behaviour to changes in flow rate. Therefore, an increase in butting and teat
changes can reflect a problem in milk availability when a calf is sucking the dam
or a milk feeding system.

4. When the dam is suckled by her calf


The early separation of the calf from the cow is seen as a keystone of the
modern dairy industry, and as essential to maximum production. However,
some segments of the population are disquieted by this, considering it
unnatural. Moreover, minimising disease when calves are raised separately
from their mothers remains difficult (Rushen, 1994). In addition, there have
been few studies of how early weaning of replacement heifers might affect their
later productivity. The commercial success of ecological milk in certain
countries (where the calves must be kept for a period of time with their dam)
has renewed interest in exploring any possible benefits of cow/calf rearing
systems.
Uncontrolled access by the calf to the mother can reduce milk yield at milking
by 40%, although there are no obvious long term changes in milk production.
However, calf growth rates are far higher, and calving-conception intervals may
be reduced (Metz, 1987). Furthermore, the negative effect on milk yield at
milking may be reduced if the number of nursings are controlled. Krohn et al.
(1999) studied cows who were allowed to nurse their calves (replacement
heifers) after milking for 8 weeks. The total amount of milk produced by the
cows was increased, so that although milk yields at milking were reduced, this
decrease was much less than the quantity drunk by the calves, suggesting an
overall economic advantage. Incidence of mastitis were nearly halved. Growth
rates of heifers nursing their dams was much greater than for weaned heifers.
At Lennoxville, we examined milk production of Holstein cows that were milked
twice daily, and also nursed their calves twice a day, 2 h after milking. Average
milk yields at milking during 9 weeks of nursing were 20% lower than controls.
When the amount drunk by the calves was added to the amount of milk
obtained at milking, nursed cows did not differ from control cows in total milk
produced. Residual milk following oxytocin injections was higher in nursed cows
than in control cows, and during milking, plasma oxytocin concentrations were
higher for control cows than for nursed cows, suggesting a lack of milk ejection
(de Passill et al., 1997b). In contrast to the previous studies, these results
show no obvious advantage for milk production by keeping the calf with the
cow. The amount of milk obtained at milking was reduced by the quantity drunk
by the calf. This difference from Krohn et al.s (1999) results may be due to the
time of nursing being too close to the previous milking. However, in keeping with
Krohn et al.s results, the nursing calves, during the nursing period, were

healthy and had growth rates double those of control calves that received the
standard ration of the Centre.

5. Motivation to suck and butt when suckling the cow


We also examined the behaviour of the calf when suckling the cow under
controlled experimental conditions where we modified milk availability as well as
calf level of hunger. We altered the availability of milk in the cows udder by
partly milking the cow immediately prior to a nursing. We also altered the level
of hunger of the calf, either by depriving it of one meal or by tube feeding. When
the cow had been fully milked, the calf sucked for twice as long and butted three
or four times more frequently than when the cow had been partly milked but
ingested the same amount of milk in both cases. When the calf was made more
hungry it sucked for twice as long, butted at a similar rate, but drank more milk
than at a normal nursing. When the calf was satiated via tube feeding 4 l before
the nursing, it sucked for a shorter time, butted at a similar rate and drank less
milk than when the calf was tube fed only 0.4 l (de Passill et al., 1996).
Consequently, the duration of the calfs sucking behaviour is not a very good
predictor of milk intake as has previously been suggested (Lidfors et al.,
1994, Mendl and Paul, 1989 and Cameron, 1998). However, a very high rate of
butting may indicate a problem of milk availability.
Lidfors et al. (1994) found that butting by calves during nursing occurred at the
beginning of the meal, but also toward the end of a period of sustained sucking.
They suggested that the end of nutritive sucking and the beginning of nonnutritive sucking during a nursing bout could be seen by the marked increase in
the amount of butting towards the end of the meal. Haley et al.s (1998b) results
using an artificial teat feeding system support this point. They found the highest
frequency of butting at the beginning of the milk meal and also once the milk
flow ended (Fig. 6). By turning the milk flow on and off during a milk meal, they
found that calves increase butting dramatically when the milk is turned off. This
is further supported by Mayntz and Costas (1998) results showing that
following an oxytocin surge, and consequently, liberal milk availability, nursing
bouts uninterrupted by butting, teat release or teat changes are longer than
when there is no oxytocin surge. Consequently, the rate of butting by the calf
may give some indication of milk transfer.

6. Conclusion
In conclusion, sucking is an important behaviour for the calf because it may
affect metabolic hormone secretion and performing sucking reduces sucking
motivation. Sucking is stimulated by the intake of milk and when the calf has not
learnt that the mother is the proper sucking stimulus (Krohn et al., 1999), we
must provide appropriate stimuli to satisfy the calfs motivation to suck when it is
fed milk. We recommend a combination of slower milk flow and the provision of
a non-nutritive artificial teat and hay feeding at the end of the milk meal to
reduce the occurrence of cross-sucking following a milk meal.

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