Invited review: Development and expression of

dairy calf feeding behaviour

E. K. Miller-Cushon1 and T. J. DeVries2,3
1
Department of Animal Sciences, University of Florida, 2250 Shealy Drive, Gainesville,
Florida, 32601, USA; and 2Department of Animal and Poultry Science, University of Guelph,
Kemptville Campus, 830 Prescott Street, Kemptville, Ontario, Canada K0G 1J0.
Received 23 October 2014, accepted 14 April 2015. Published on the web 11 May 2015.
Miller-Cushon, E. K. and DeVries, T. J. 2015. Invited review: Development and expression of dairy calf feeding behaviour.
Can. J. Anim. Sci. 95: 341350. Feeding and housing practices for dairy calves impact performance and growth early in life,
Can. J. Anim. Sci. Downloaded from cdnsciencepub.com by 177.182.139.121 on 09/05/23

as well as feeding behaviour. There is also increasing evidence that early exposure to different feeding and housing strategies
influence the development of feeding behaviour. The dairy calf needs access to milk in sufficient quantities to maintain
health and high-levels of growth. In addition, intake of solid feed, such as grain concentrate, is necessary early in life to
establish fermentation and develop the rumen. In dairy calves, feeding behaviour patterns can have direct impact on nutrient
consumption and, consequently, growth. Further, recent research has provided evidence that behaviour patterns that
develop in response to management factors early in life may persist once learned, potentially having longer-term implications
for health and welfare. Thus, there is potential for various nutritional, housing, and management factors to impact the
learning of feeding behaviour early in the life of dairy calves. This review provides several examples of early management
factors that influence feeding behaviour of dairy calves and, in turn, impact calf growth and welfare. For example, providing
continuous, ad libitum access to milk results not only in greater growth, but also in meal patterns that more closely resemble
the natural behaviour of a calf suckling its dam. Housing management also has the potential to influence when, and how
much, a calf eats. Whereas social facilitation promotes intake, particularly at weaning, competition for feed in group-housed
calves restricts feeding patterns and degree of feeding synchrony. With regards to provision of solid feed, recent data suggest
that providing physically effective hay, in addition to grain concentrate, may be beneficial for the rumen environment
and consequent growth and efficiency. However, the physical form and presentation of forage may also influence feeding
behaviour of calves, in particular feed sorting, which can impact both immediate nutrient intakes as well as the development
and persistence of that behaviour. For example, calves provided hay and concentrate as a mixture begin to sort their feed
early in life, with the pattern of sorting depending on hay particle size. Feed sorting in young calves may reflect a motivation
to consume a proportion of hay in the diet. In general, it appears to be beneficial for early growth and welfare to support milk
and solid feed intake through unrestricted allowances and less-competitive feeding environments. Continued research in this
area is needed to assess the longevity of learned behaviour patterns, and what factors may influence their persistence.

Key words: Behaviour, feeding, dairy calf

Miller-Cushon, E. K. et DeVries, T. J. 2015. Article de synthèse sollicité: Développement et expression du comportement

alimentaire des veaux laitiers. Can. J. Anim. Sci. 95: 341350. Les pratiques alimentaires et de logements des veaux laitiers ont
un impact sur la performance et la croissance tôt dans leur vie, ainsi que sur le comportement alimentaire. Il y a aussi de plus
en plus de preuves que l’exposition précoce aux différentes stratégies d’alimentation et de logement a une influence sur le
développement du comportement alimentaire. Le veau laitier doit avoir accès au lait en quantités suffisantes pour maintenir
la santé et de forts taux de croissance. De plus, la consommation d’aliments solides, tels que le concentré de grains, est
nécessaire tôt dans la vie pour établir la fermentation et pour le développement du rumen. Chez les veaux laitiers, les profils
de comportement alimentaire peuvent avoir un impact direct sur la consommation des éléments nutritifs et, par conséquent,
sur la croissance. De plus, des études récentes indiquent que les comportements qui se développent en réponse aux facteurs de
gestion tôt dans la vie peuvent persister une fois appris, et ainsi avoir des implications à long terme sur la santé et le bien-être.
Donc, il y a un potentiel pour que les différents facteurs nutritionnels, de logement et de gestion aient un impact sur
l’apprentissage des comportements alimentaires dans la vie précoce des veaux laitiers. Cet article de synthèse offre plusieurs
exemples des facteurs de gestion précoce qui ont une influence sur le comportement alimentaire des veaux laitiers et qui,
à leurs tours, ont un impact sur la croissance et le bien-être des veaux. Par exemple, offrir l’accès ad libitum en continu au lait
se solde par une plus grande croissance, mais aussi par des profils de repas qui ressemblent plus étroitement au comportement
naturel d’un veau qui allaite auprès de sa mère. La gestion du logement a aussi le potentiel d’avoir une influence sur quand
et combien le veau se nourrit. Tandis que la facilitation sociale favorise la consommation, particulièrement au sevrage,
la compétition pour les aliments chez les veaux logés en groupes restreint les profils d’alimentation et le degré de
synchronisation de l’alimentation. Par rapport à fournir la nourriture solide, des données récentes suggèrent qu’offrir le
foin matériellement efficace en plus du concentré de grains peut avoir un effet bénéfique à l’environnement du rumen et la
croissance et l’efficacité qui en découle. Par contre, la forme physique et la présentation des fourrages peuvent aussi

3
Corresponding author (e-mail: tdevries@uoguelph.ca).
Can. J. Anim. Sci. (2015) 95: 341350 doi:10.4141/CJAS-2014-163 341
 342 CANADIAN JOURNAL OF ANIMAL SCIENCE

influencer le comportement alimentaire des veaux, particulièrement au niveau du tri alimentaire, ce qui peut avoir une
influence immédiate sur la consommation des éléments nutritifs ainsi que le développement et la persistance de tels com-
portements. Par exemple, les veaux à qui l’on donne le foin et le concentré comme mélange commencent a trier leur nourriture
de façon précoce avec le profil de tri qui dépend de la taille de particule du foin. Le tri alimentaire chez les jeunes veaux peut
refléter la motivation à consommer une portion de foin dans la diète. De façon générale, il semble que ce soit avantageux pour
la croissance et le bien-être précoce de soutenir la consommation de lait et d’aliments solides sans restrictions et dans des
environnements d’alimentation ayant peu de compétition. Davantage de recherche dans le domaine est nécessaire pour
évaluer la longévité des profils de comportement acquis ainsi que pour trouver les facteurs qui peuvent influencer leur
persistance.

Mots clés: Comportement, alimentation, veau laitier

Modern practices for managing dairy calves greatly with different approaches for milk feeding level, method
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influence the development and expression of feeding
behaviour. The early separation of the calf from the cow
has been described as the keystone of the modern dairy
industry (de Passillé 2001) and, as a result, all aspects
of calf feeding are controlled by management practices,
providing much opportunity to influence when and how
the calf consumes feed. A diversity of management prac-
tices for dairy calves are currently employed on farms,
with varying approaches to both milk feeding manage-
ment and strategies for introducing solid feed, such as
concentrate and hay. Understanding the role of manage-
ment factors in performance of feeding behaviour can
provide insight into feeding patterns which may, im-
mediately or in the future, negatively impact health and
welfare. Further, recent research into the role of learning
in feeding behaviour of dairy calves suggests that early
management strategies may influence behaviour after
weaning and, potentially, later in life.
FEEDING BEHAVIOUR UNDER NATURAL
CONDITIONS
Suckling behaviour of the calf with the dam has been
observed in natural settings in beef cattle and semi-
wild herds of cattle. Suckling patterns follow a circadian
rhythm, with increased activity at dawn and dusk (Odde
et al. 1985). In general, calves perform 4 to 10 suckling
bouts per day, which last an average of 7 to 10 min per
bout (de Passillé 2001). Suckling bout frequency and total
time spent nursing decrease as lactation progresses, in
relation with milk production decline (Day et al. 1987).
In the presence of their dam, calves begin to graze
early in life. Extensively reared beef calves have been
reported to graze slowly (‘‘picking’’ the pasture) for short
periods of time (10  15 min) in the first weeks of life,
with grazing time increasing linearly from 20 min at
10 d to 360 min at 100 d of age, 70% of grazing time of
adult cow (Nicol and Sharafeldin 1975). Diurnal grazing
patterns of calves mirrored those of the adult cows, with
similar peaks in feeding activity, but reduced activity at
off-peak times compared with adult cattle (Nicol and
Sharafeldin 1975).
Modern dairy production systems impose a number of
restrictions on feeding behaviour of the milk-fed calf, as
calves are fed artificially apart from the dam. Practices
for raising dairy calves vary considerably among farms,
of milk delivery (e.g., bucket vs. teat; Hammell et al.
1988), social grouping (e.g., group or pair vs. individual;
Chua et al. 2002), and type of solid feed provided (e.g.,
provision of forage vs. highly digestible grain; Castells
et al. 2012). Management practices dictate when and
what calves can eat and, consequently, influence devel-
opment of feeding behaviour and are closely linked with
performance and welfare.
ROLE OF LEARNING IN FEEDING BEHAVIOUR
The development of feeding behaviour relies on a variety
of learning mechanisms (Day et al. 1998; Galef and
Giraldeau 2001). Dietary preference and selection develop
in response to associations between sensory properties of
foods and post-ingestive consequences (Provenza et al.
1992), with social environment influencing the likelihood
of encountering particular stimuli through social facil-
itation (Galef 1995). Feeding patterns are also subject to
learning, as the start of a meal is influenced by condi-
tioned responses to external stimuli (Weingarten 1983;
Reppucci and Petrovich 2012). In cattle, operant con-
ditioning can be used to train cattle to approach the feed
bunk in response to an auditory cue (Wredle et al. 2004),
and there is anecdotal evidence to suggest that cattle
learn to approach the feed bunk in response to activity
associated with feed delivery. When animals are housed in
groups, feeding patterns are also subject to social factors.
Social facilitation is the process by which the behaviour
of a conspecific, such as another animal in the pen,
releases the performance of the same behaviour in the
subject (Nicol 1995); for example, if a single cow begins
to feed, others will soon follow suit (Albright 1993).
Learning of feeding behaviour is of interest with
regard to dairy calves, as there is evidence of critical
periods early in life where learning occurs rapidly and,
thus, early experiences may influence longer-term beha-
viour patterns. Weaning from milk is a sensitive period
of learning for young animals, during which feed intake
is increasing and learning about appropriate feed types
must occur rapidly (Provenza and Balph 1987). Squibb
et al. (1990) reported that lambs exposed to a particular
type of shrub at the period of transition from non-
ruminant to ruminant (4 to 8 wk of age) demonstrated a
more persistent preference for that food, compared with
lambs exposed earlier or later in life. Similarly, Arnold

and Maller (1977) determined that exposure to foraging
environments (differing in variety of plants growing in
different regions of Australia) had longer-term effects
on dietary preference when exposure occurred earlier
(in the first 6 mo), rather than later in life. In addition to
influencing development of feed preferences, early feed
exposure can influence the development of motor skills
used for foraging. Foraging motor skills are influenced
by prior exposure to plant types, with animals foraging
more efficiently when they have previous experience
with similar plant types (Arnold and Maller 1977). Also,
exposure to particular plant types appears to affect for-
aging style and success. For example, lambs exposed to
either grass or a shrub developed different prehension
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patterns, described as jerking or chewing, and had dif-
ferent foraging success, as indicated by bite size, bite rate,
and rate of rate of intake, when offered the unfamiliar
feed type (Flores et al. 1989).
In dairy calves, aspects of the environment, such as
milk feeding method, social environment, and strategies
for introducing solid feed, greatly influence the expres-
sion of feeding behaviour and have potential to impact
the learning of behaviour patterns.
FEEDING OF MILK AND MILK REPLACER
Feeding behaviour of dairy calves depends greatly on
milk feeding management. For much of the past few
decades, dairy calves have conventionally been fed 10%
of their birth body weight in milk, an amount that
translates to between 4 and 5 kg d1 of liquid milk, and is
less than 50% of reported ad libitum intakes of suckling
calves (Appleby 2001; Tedeschi and Fox 2009). Calves
fed restricted quantities of milk consume the entire milk
allotment rapidly at the time of delivery, and have fre-
quent unrewarded visits to the milk feeder (Jensen 2004;
De Paula Vieira et al. 2008), suggesting that they are
Time spent engaged in activity (min/h)
14
12
10
0
0:00
Fig. 1. Diurnal feeding activity of calves provided milk at two feeding levels: ad libitum and restricted at 5 L d 1 (2.5 L provided at
0800 and 1600). Time spent engaged in non-nutritive sucking is shown for calves provided restricted amounts of milk. [Adapted
from Miller-Cushon et al. (2013a); reproduced with permission from the Journal of Dairy Science.]
MILLER-CUSHON AND DEVRIES * DAIRY CALF FEEDING BEHAVIOUR 343
hungry and, thus, experiencing poor welfare (De Paula
Vieira et al. 2008). Further, when fed via an artificial teat,
restricted-fed calves spend considerable amounts of time
engaged in non-nutritive sucking throughout the day
(Fig. 1) and have non-nutritive sucking bouts at a fre-
quency similar to milk meal frequency in calves fed
milk ad libitum (Miller-Cushon et al. 2013a). Sucking has
functional consequences for the calf, causing the release
of hormones involved in postprandial satiety, such as
insulin, cholecystokinin, and gastrin (de Passillé et al. 1993)
and it is a behavioural need in itself, as the motivation
to suck is reduced to a greater extent by sucking time
than by milk ingestion (Rushen and de Passillé 1995;
de Passillé 2001).
In contrast to conventional restricted milk feeding, en-
hanced milk feeding programs provide elevated quanti-
ties of milk or ad libitum access to milk, allowing diurnal
milk meal patterns closer in resemblance to those of a
suckling calf, with feeding activity distributed over the
day (Fig. 1; Miller-Cushon et al. 2013a). Meal frequen-
cies similar to suckling calves have been reported in ad
libitum milk feeding systems, with reports of average
meal frequency ranging from 4 to 10 meals per day
(Appleby 2001; De Paula Vieira et al. 2008; Miller-
Cushon et al. 2013a). Calves have peaks in feeding
activity corresponding to fresh milk delivery, as well as
increased activity at sunrise and sunset (Senn et al. 2000;
Appleby 2001; Miller-Cushon et al. 2013a). In general,
providing greater quantities of milk appears to be bene-
ficial in allowing preferred feeding patterns and support-
ing increased rate of weight gain and greater structural
growth [e.g., approximately 1.0 kg d1 on enhanced
milk feeding programs vs. 0.45 kg d1 on conventional
restricted milk feeding programs; as reviewed by Khan
et al. (2011)]. However, enhanced milk feeding programs
may present challenges at time of weaning, if the calf has
ad libitum milk feeding
milk feeding time - 5 L/d
non-nutritive sucking
2:00 4:00 6:00 8:00 10:00 12:00 14:00 16:00 18:00 20:00 22:00 0:00
Time of day
 344 CANADIAN JOURNAL OF ANIMAL SCIENCE
not begun consuming sufficient amounts of solid feed
prior to removal of milk. When provided greater quanti-
ties of milk, calves have less frequent and smaller meals of
concentrate (Miller-Cushon et al. 2013a), which delays
rumen development (Terré et al. 2007; Hill et al. 2008). In
some cases, poor weight gain through weaning negated
any body weight advantage arising from an enhanced
milk feeding program (Borderas et al. 2009; DePassillé
et al. 2011). Consistent weight gain through weaning is
improved with gradual weaning programs that encou-
rage solid feed intake (Khan et al. 2007).
Milk feeding level has potential to influence the calf
beyond the milk feeding stage, as there is evidence that
early rate of growth is associated with increased milk
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production into the first lactation (Soberon and Van
Amburgh 2013). In addition, milk feeding level and meal
patterns may have long-term metabolic consequences for
the calf, as large, infrequent milk meals (3.5 L feedings
twice daily) have been found to increase serum insulin to
glucose ratio (0.19 vs. 0.12 pg mmol 1; Terré et al. 2009),
suggesting that this pattern of feeding may elicit insulin
resistance (Bach 2012). From a behavioural standpoint,
there is some evidence to suggest that milk feeding level
may influence patterns of solid feed intake for a short
period of time after weaning. Compared with calves
provided milk ad libitum, calves fed restricted amounts
of milk have greater rates of intake while feeding initially
post-weaning, and feed more continuously in response
to feed delivery, taking fewer pauses during their first
meal (Miller-Cushon et al. 2013a). These differences in
feeding patterns were reported only initially after wean-
ing, suggesting no longer-term impact of milk feeding
level on feeding behaviour of calves housed individually
(Miller-Cushon et al. 2013a). However, it is possible
that differences in feeding patterns resulting from early
experience are more likely to persist in situations where
there is an interaction with social factors, such as com-
petitive pressure at the feed bunk.
SOCIAL ENVIRONMENT
Social environment is an influential factor in the devel-
opment of feeding behaviour, as feeding activity is socially
facilitated. Compared with calves housed individually,
calves reared in groups of two or more begin consuming
solid food earlier in life and consume more solid food
(Babu et al. 2004; Hepola et al. 2006; De Paula Vieira
et al. 2010), in more frequent daily meals (Miller-Cushon
et al., in preparation). Thus, social facilitation encourages
early intake of feed and is beneficial in supporting a
successful transition at weaning (Chua et al. 2002;
Miller-Cushon et al. 2014a).
When calves are housed in groups, competition be-
comes a factor that may restrict access to milk early in
life. Frequencies of competitive displacements at the milk
feeder or teat increase with decreasing teat availability
(von Keyserlingk et al. 2004; Miller-Cushon et al. 2014b)
and increasing group size (Jensen 2004; Jensen and
Budde 2006). Increased competition for access to milk
can reduce meal frequency and milk intake during short-
term experiments (von Keyserlingk et al. 2004) and early
in the milk-feeding stage (Miller-Cushon et al. 2014b).
However, pair-housed calves have been found to com-
pensate for restricted teat access over the duration of the
milk-feeding stage by increasing meal frequency, total
feeding time, and therefore milk intake (Miller-Cushon
et al. 2014b). Competition for access to teats also reduces
the degree of synchrony between milk and solid feed
meals (Miller-Cushon et al. 2014b). Further research is
encouraged to assess implications of competition for teat
access on the development of feeding patterns and milk
intake when calves are housed in larger, more competi-
tive groups (e.g., groups of 1015 calves housed with
computer-controlled milk feeders with access to only one
or two teats; Weber and Wechsler 2001; Jensen and
Holm 2003).
Early social environment influences the development
of social behaviour, impacting social hierarchy later in
life (Warnick et al. 1977) and response to competition
(Duve et al. 2012) or a novel environment (De Paula
Vieira et al. 2012). For example, Duve et al. (2012) reported
that calves reared with social contact (pair-housing)
accessed feed more successfully in a competitive chal-
lenge than calves reared in individual pens. There is
also evidence that differences in meal patterns developed
in response to early environment may persist when
mediated by different social factors, such as degree of
competition. For example, differences in meal frequency
resulting from early social housing have been found to
persist when calves are housed similarly, in social groups,
after weaning (Miller-Cushon et al., in preparation).
Further, dairy calves raised with restricted access to
milk feeding stations were found to consume their feed
more rapidly in shorter meals post-weaning, compared
with calves raised with unrestricted teat access, despite
being fed non-competitively in the post-weaning period
(Miller-Cushon et al. 2014b). As rate of feed intake can
be considered an indicator of social constraint (Nielsen
1999), it is likely that these post-weaning differences in
feeding behaviour were mediated by learned competitive
behaviour: calves reared with restricted teat access con-
tinued to displace each other more frequently and had
reduced synchrony in their feeding compared with calves
reared in less-competitive pens (Miller-Cushon et al.
2014b). Similarly, Greter et al. (2010) reported that dairy
heifers with previous exposure to different feed delivery
methods (a total mixed ration compared with forage top-
dressed with gain concentrate) had persistent differences
in competitive behaviour: heifers displaced each other
more frequently and spent more time feeding in response
to feed delivery when they had experience competing for
access for concentrate at feed delivery when provided a
top-dressed ration.
Feeding patterns developed in response to pre-
vious experience with competition may persist, having
a longer-term impact on health. Differences in post-
weaning feeding patterns resulting from early competitive
 MILLER-CUSHON AND DEVRIES * DAIRY CALF FEEDING BEHAVIOUR
environment, as observed by Miller-Cushon et al. (2014b),
are consistent with differences in feeding patterns result-
ing from competition for access to feed in older animals:
in heifers and adult cows, reducing available feeding sta-
tions or bunk space increases intake rates (Hosseinkhani
et al. 2008; Proudfoot et al. 2009; DeVries and von
Keyserlingk 2009) and competitive interactions around
feed (DeVries et al. 2004; Huzzey et al. 2006), while
decreasing meal frequencies (Hosseinkhani et al. 2008;
DeVries and von Keyserlingk 2009). The effect of a
competitive environment on feeding patterns may have
implications for long-term health of dairy cattle. Meal
size and frequency can impact the rumen environment, as
ruminal pH declines following meals, and the rate of
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decline is proportional to meal size (Allen 1997). Further,
reduced feeding time may decrease salivary secretion,
which acts as a buffer in the rumen (Beauchemin et al.
2008). Thus, it is possible that feeding patterns char-
acterized by greater rates of intake and larger, infrequent
meals may lead to larger post-prandial drops in rumen
pH, potentially increasing the risk of subacute ruminal
acidosis (Krause and Oetzel 2006). In addition, increased
competitive behaviour for access to feed may result in
greater within-pen variability in feed access and weight
gain (González et al. 2008) and day-to-day variability
in feeding time and meal characteristics (DeVries and
von Keyserlingk 2009), negatively impacting the perfor-
mance and welfare of smaller, less-dominant animals.
Provision of Solid Feed
In the rearing of dairy calves, a primary goal is to
encourage solid feed intake to support rapid rumen
development and facilitate weaning from milk or milk
replacer. Learning about novel feeds in ruminants occurs
through a process of sampling available food types and
developing associations between sensory characteristics
of a feed and post-ingestive feedback (Provenza and
Balph 1987). Ruminants develop preferences for feeds
associated with positive post-ingestive feedback, such as
foods paired with nutrients, such as glucose (Burritt and
Provenza 1992) and aversions for feeds associated with
negative post-ingestive feedback resulting from toxins,
such as lithium chloride (Burritt and Provenza 1989), or
excess amounts of a nutrient, such as urea (Kertz et al.
1982). The integration of sensory properties of a food
with its post-ingestive feedback influences the perceived
palatability of the feed type, which, in turn, shapes
dietary selection patterns as animals choose feeds to
select and avoid (Provenza et al. 1992). In a number
of cases, young ruminants have demonstrated an ability
to employ this process of dietary learning to select a
balanced diet from a variety of feed types; for example,
lambs offered a low protein food and a high protein food
were able to choose a mixture which provided optimal
amounts of protein for growth (Kyriazakis and Oldham
1993; Bach et al. 2012). This dietary learning may be due
to association of particular foods with negative post-
ingestive feedback resulting from microbial production
345
of excess amounts of nitrogen and resultant production
of high amounts of ammonia (Kertz et al. 1982; Provenza
1995).
Dairy calves are typically provided with highly diges-
tible grain concentrate, as rumen papillae development
occurs in response to butyrate produced in the rumen
through fermentation of carbohydrates (Warner et al.
1956; Sander et al. 1959). Forage provision remains con-
troversial due to the concern that it may displace con-
centrate intake and, consequently, impair rumen papillae
development (Hill et al. 2008; Kertz et al. 1979). How-
ever, there is evidence to suggest that forage provi-
sion does not need to reduce concentrate intake (Khan
et al. 2011; Castells et al. 2012) and, further, may posi-
tively impact rumen environment, increasing rumen pH
(Suárez et al. 2007; Khan et al. 2011) and feed efficiency
(Coverdale et al. 2004). Provision of hay to dairy calves
also reduces the occurrence of non-nutritive oral beha-
viours (Castells et al. 2012), suggesting it satisfies a need
to exercise foraging behaviour. Diurnal feeding patterns
of hay and concentrate indicate that calves spend the
majority of time consuming hay during peak feeding
periods, when fresh feed is delivered, and more time
consuming concentrate than hay throughout the rest of
the day (Miller-Cushon et al. 2013c). It is interesting to
note when calves are offered a choice of hay and con-
centrate, across multiple studies they have been shown to
select a proportion of hay ranging between 5 and 30% of
total dry matter intake (Khan et al. 2011; Castells et al.
2012; Miller-Cushon et al. 2013b). Dietary selection of
forage is likely to depend on characteristics of the forage.
For example, Castells et al. (2012) reported that provi-
sion of different types of hay, straw, and silages resulted
in similar or increased grain concentrate compared with
calves without access to forage, except for provision
of alfalfa hay, which reduced concentrate intake. These
authors suggested that the contrasting response to alfalfa
hay may be due to differences between legumes and
grasses in terms of composition and flow rate from the
rumen (Castells et al. 2012). Dietary selection of hay and
concentrate may also depend in part upon milk allow-
ance, as calves increase concentrate intake, relative to hay
intake, after weaning (Fig. 2; Miller-Cushon et al. 2013b),
at which point they have greater reliance on solid food
for energy.
Changes in dietary selection in relation to milk
allowance could suggest that selection of concentrate
depends on energy content of the feed, which could
cause positive post-ingestive feedback. Feedback related
to rumen environment may also play a role, as adult
cattle have been found to attenuate effects of low rumen
pH by increasing sorting in favour of longer particles
(DeVries et al. 2008, 2014 a, b) and increasing preference
for hay (Keunen et al. 2002). However, evidence to
date suggests that when provided a variety of feed types,
the ability of dairy calves to select a nutritionally
balanced diet prior to weaning is limited. Montoro
and Bach (2012) reported that dairy calves offered six
 346 CANADIAN JOURNAL OF ANIMAL SCIENCE
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Fig. 2. Total feed intake and dietary selection of hay and concentrate (percent of total intake) over time by calves offered these feed
types separately. Calves were weaned during week 7 of age and no milk was provided during week 8. [Adapted from Miller-Cushon
et al. (2013b); reproduced with permission from the Journal of Dairy Science.]
separate ingredients over-consumed protein, and Miller-
Cushon et al. (2014c) reported that protein intake by
dairy calves offered a choice of two pelleted diets was
influenced by source of the protein. In these studies,
dietary selection was apparently driven by palatability
factors (in both cases, a preference for soybean meal;
Miller-Cushon et al. 2014d) to a greater extent than
nutrient requirements. Consequently, sensory properties
of feed, rather than post-ingestive consequences, may
be most influential in the feed preferences and dietary
selection of young or pre-ruminant calves, with innate
preferences dictating feed choices. It is possible that the
ability of young calves to associate individual types of
solid feed with related post-ingestive consequences is
impeded when they are also ingesting milk, as dietary
learning is limited when animals are provided multiple
feed choices (Duncan and Young 2002). It follows that
the composition of particular feeds offered may affect
feeding behaviour and patterns of dietary selection.
Further research is encouraged to assess factors driving
dietary selection in dairy calves, and investigate the moti-
vation of calves to consume forage prior to weaning.
Researchers have recently investigated the extent to
which dairy calves select within a mixed diet, providing
evidence that calves are adept at sorting a ration of con-
centrate and hay early in life (e.g., prior to weaning at
week 6 of age; Miller-Cushon et al. 2013b). Calves have
been found to select in favour of hay prior to weaning
when provided a mixed diet consisting of chopped hay
and concentrate (diet composed of 7090% concentrate;
Montoro et al. 2013; Miller-Cushon et al. 2013b), with
selection in favour of concentrate increasing after wean-
ing off of milk (Miller-Cushon et al. 2013b, c). Feed
sorting is clearly subject to characteristics of the ration;
for example, when provided a mixed diet containing hay
that was finely ground rather than coarsely chopped,
Montoro et al. (2013) found that calves selected in favour
of the concentrate component prior to weaning, suggest-
ing a preference for physically effective ration particles.
The development of feed sorting in dairy calves
appears to be subject to early experience, with the extent
of feed sorting after weaning dependent upon feeding
strategies prior to weaning. Calves exposed to a mixed
diet (of forage and concentrate) prior to weaning have
been found to sort to a greater extent up to 6 wk after
weaning, compared with calves exposed to separate feed
components prior to weaning (Fig. 3; Miller-Cushon
et al. 2013b). Further, differences in the extent of feed
sorting persisted through a transition to a novel ration,
suggesting that expression of feed sorting may depend on
the acquisition of feeding motor skills (such as physical
discrimination between particles) required for sorting
(Miller-Cushon et al. 2013b). Similarly, pre-weaning ex-
posure to rations differing in physical form of forage
resulted in the development of different patterns of
feed sorting (Montoro et al. 2013), which persisted after
transition to a common ration post-weaning (Miller-
Cushon et al. 2013d). Whereas the degree of feed sorting
has been found to vary among calves, dependent upon
prior experience, the pattern of feed sorting appears to
be less flexible. Post-weaning, dairy calves that sort their
feed have consistently been observed selecting in favour
of the higher-energy grain component (Miller-Cushon
and DeVries 2011; Miller-Cushon et al. 2013b, d),
similar to patterns of feed sorting observed in adult
cattle (Leonardi and Armentano 2003; Hosseinkhani
et al. 2008; Miller-Cushon and DeVries 2009). Pre-
weaning environment has been found to have only short-
lived effects on the pattern, or direction, of feed sorting
post-weaning; calves exposed to either concentrate or
hay prior to weaning sorted a mixed diet in favour of
the familiar feed component initially after weaning,
 MILLER-CUSHON AND DEVRIES * DAIRY CALF FEEDING BEHAVIOUR 347
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Fig. 3. Effects of previous exposure to hay and concentrate presented either as a mixture or separately on sorting of dairy calves
offered a mixed ration after weaning. Sorting is shown as intake of NDF (reflecting hay selection) or NFC (reflecting concentrate
selection) as a percent of predicted intake, where predicted intake was based on the composition of the mixed diet (70% concentrate
and 30% hay). Values100% indicate sorting for hay or concentrate, and values B100% indicate sorting against hay or con-
centrate. *PB0.05. [Adapted from Miller-Cushon et al. (2013b); reproduced with permission from the Journal of Dairy Science.]

but differences in sorting patterns did not persist, with all
calves beginning to sort in favour of concentrate and
against hay (Miller-Cushon and DeVries 2011). Thus,
results of studies to date generally suggest that the emer-
gence and persistence, but not the pattern, of feed sort-
ing may depend on the opportunity and motivation to
exercise feed sorting early in life. The emergence of feed
sorting at this point in life may be influenced by the ease
with which the ration is physically separated and sorted
by the calves, as well as how well the ration meets the
changing nutrient requirements of the calf.
The potential to influence longer-term feed sorting
through early management has implications for cattle
welfare. In adult cattle, it is well understood that feed
sorting results in an unbalanced intake of nutrients that
has potential to impair rumen health and negatively im-
pact performance, both at the cow (DeVries et al. 2008)
and herd (Sova et al. 2013) level. No work to date has
investigated the effects of feed sorting in dairy calves on
their rumen development or aspects of rumen environ-
ment, such as pH. However, Greter et al. (2010) reported
an association in dairy heifers between feed sorting and
looser stools, indicating reduced rumen pH (Krause and
Oetzel 2006). The extent of feed sorting is highly variable
between adult cattle (Leonardi and Armentano 2003)
despite efforts to thwart feed sorting at the herd level,
such as increasing feeding frequency (DeVries et al. 2005)
and altering ration dry matter content (Leonardi et al.
2005). This variability must reflect individual differ-
ences in factors controlling the expression of the beha-
viour, which could include early experience. Thus, early
management should aim to minimize the expression
and development of this behaviour. Further work should
assess the longevity of differences in feed sorting resulting
from early experience, as well as the potential to generalize
recent findings across a variety of diets.
CONCLUSION
Feeding behaviour of dairy calves is highly subject to
early management, providing opportunity to influence
the development of feeding behaviour patterns. Feeding
behaviour has a direct effect on performance and health
of the dairy calf and learned behaviour patterns may
persist and impact welfare over a longer period of time.
For example, enhanced milk feeding level benefits growth
and, potentially, longer-term performance, as well as
feeding patterns. Social housing influences feed intake
and feeding patterns and may have persistent effects on
response to competition for access to feed, suggesting
that competition should be minimized in the pre-weaning
housing environment. The approach to presenting solid
feed to calves influences both feeding patterns and feed
sorting, and post-weaning expression of feed sorting may
be subject to previous experience. In summary, dairy
calves should be fed and housed to support the expression
and development of feeding patterns conducive to good
health and welfare.
ACKNOWLEDGEMENTS
Much of the research reviewed in this paper was funded,
in part, by a Natural Sciences and Engineering Research
Council of Canada (NSERC; Ottawa, ON, Canada) Dis-
covery Grant awarded to T. J. DeVries and an Ontario
Ministry of Agriculture Food and Rural Affairs (OMA-
FRA)/University of Guelph Production Systems research
grant.
 348 CANADIAN JOURNAL OF ANIMAL SCIENCE
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