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Sci. 90: 295302. There has recently been a significant increase in the amount of scientific research relating dairy cattle
behaviour to nutritional management, particularly with respect to the interaction with health. Much of this research has
been focused on using information on the natural behavioural patterns of dairy cattle to improve management practices.
As with calves and mature dairy cattle, the behaviour of growing replacement dairy heifers interacts with various
nutritional management practices. For replacement dairy heifers, much of the research in this area has been focused on
how these interactions relate to different feed delivery methods, the amount of feed provided, and stocking density at the
feed bunk. This review describes empirical work focused on understanding how nutritional management practices affect
heifer behaviour, and how these practices may affect the health, productivity, and welfare of these animals. In particular,
evidence will be provided for those practices (such as total mixed ration feeding, feeding for ad libitum intake, and reducing
feed bunk competition) that promote feeding behaviour patterns related to more consistent nutrient intake and improved
rumen health. Even though the behavioural effects of these nutritional practices are known, additional research is still
needed to further demonstrate the possible physiological consequences (e.g., rumen pH, growth rate) of these changes in
behaviour. This information is needed so that further recommendations can be made on these practices so these animals
may be raised for maximum production potential, while maintaining their behavioural needs, health and overall welfare.
De Vries, T. J. 2010. Le comportement et son rôle dans la gestion de la nutrition des génisses laitières en croissance. Can. J.
Anim. Sci. 90: 295302. On a assiste depuis peu à une intensification importante des recherches scientifiques sur les liens
entre le comportement des bovins laitiers et la prise en charge de leur nutrition, surtout sur le plan de la santé. Une bonne
partie de ces travaux font appel à l’information sur le comportement naturel des animaux pour améliorer les pratiques
d’élevage. Comme c’est le cas avec les veaux et les bovins laitiers adultes, le comportement des génisses de remplacement en
croissance influe sur diverses pratiques en matière de nutrition. La recherche correspondante sur les génisses laitières de
remplacement se concentre dans une large mesure sur les interactions du comportement avec les méthodes de distribution
des aliments, la quantité d’aliments fournie et la densité de peuplement aux mangeoires. Cet article décrit les travaux
empiriques visant à élucider la manière dont les techniques de prise en charge de l’alimentation influent sur le comportement
des génisses, et sur la façon dont pareilles pratiques influent sur la santé, le rendement et le bien-être des animaux. Plus
précisément, l’auteur présente des preuves sur les pratiques (distribution d’une ration complète mélangée, alimentation à
satiété et réduction de la compétition aux mangeoires) qui encouragent des habitudes alimentaires associées à une ingestion
plus uniforme d’aliments et à une meilleure santé du rumen. Bien qu’on connaisse l’incidence de ces pratiques
nutritionnelles, il faudrait entreprendre d’autres recherches pour mieux illustrer les conséquences physiologiques éventuelles
(à savoir, pH du rumen, taux de croissance) de tels changements de comportement. On a besoin de ce genre d’information
pour formuler d’autres recommandations sur ces pratiques, afin que les animaux puissent être élevés jusqu’à leur potentiel
de production maximal, sans que leurs besoins comportementaux, leur santé et leur bien-être général en souffrent.
Dairy producers replace 25 to 35% of their herd each heifer rearing program in which sound feeding, breeding,
year with 2-yr-old heifers (Lehenbauer 1998). Overall and management decisions are made. Feeding programs
herd improvement is only possible if these replacements must be designed to allow replacement heifers to be bred
are healthy, productive, and genetically superior to those efficiently, calve out by 24 mo of age, and produce to
being culled. This is achieved through a replacement their maximum potential. To achieve this, heifers must
be fed to grow at a high rate, while not becoming over-
conditioned, as well as to remain healthy. The costs
Presented at the Joint Annual Meeting of the Canadian
Society of Animal Science, the American Dairy
Science Association, and the American Society of Abbreviations: DMI, dry matter intake; TMR,
Animal Science, held in Montreal in July 2009. total mixed ration
295
296 CANADIAN JOURNAL OF ANIMAL SCIENCE
associated with replacement dairy heifer rearing are different ways, typically separate from the roughage, on
second only to the feed costs of lactating cows, repre- top of the roughage (‘‘top dressing’’), or mixed in with
senting 15 to 20% of total farm expenses (Heinrichs the roughage.
1993). Of these costs, feed represents the single greatest DeVries and von Keyserlingk (2009a) recently demon-
expense in heifer rearing costs (Gabler et al. 2000). strated, in a short-term behavioural study, that replace-
It follows, therefore, that the primary nutritional goals of ment dairy heifers, when given a choice of ration
a successful dairy replacement rearing program are to components, or when provided grain concentrate top
feed heifers for maximal production potential, at a low dressed on hay, exhibit a greater peak in dry matter
economic and environmental cost, without compromis- intake (DMI) following feed delivery than when fed a
ing health or welfare. TMR (Fig. 1). This greater peak in DMI comes as a
We have seen much advancement, particularly in the result of the consumption of the grain concentrate
past 10 years, in replacement heifer nutritional research portion of the ration in one or two large meals prior
Can. J. Anim. Sci. Downloaded from www.nrcresearchpress.com by 105.104.230.235 on 05/17/20
directed to these goals. Concurrently, we have also seen to consuming the hay portion of the ration (Fig. 2). In a
a significant rise in the amount of scientific research follow-up, longer-term study (over a period of 13 wk),
relating behaviour to the management of dairy cattle, Greter et al. (2010a) found similar feeding patterns in
particularly with respect to the interaction with health replacement dairy heifers provided with concentrate
(Weary et al. 2009). Much of this research has been top-dressed on haylage. Rapid consumption of the
focused on using information on the natural beha-
concentrate portion of the ration in few, very large
vioural patterns of dairy cattle to make management
meals, as observed in these studies, may cause large
decisions that are beneficial for the health, productivity,
post-prandial drops in rumen pH, as demonstrated in
and welfare of these animals (von Keyserlingk et al.
2009). Although most of the research in this area has young (Quigley et al. 1992; González et al. 2008) and
focused on calves and mature dairy cattle, there is a mature (Østergaard and Grohn 2000; Beauchemin et al.
growing body of literature in which the interactions of 2002) dairy cattle. Even though Greter et al. (2010a) did
replacement dairy heifer behaviour and nutritional not measure rumen pH, they did observe more liquid
management are considered. This review describes those fecal consistency in heifers fed the top-dressed ration
For personal use only.
primary areas where researchers have considered the suggesting that these heifers experienced some level of
interaction of behaviour and nutritional management in sub-acute ruminal acidosis (Plaizier et al. 2008). In both
replacement dairy heifer rearing. Specifically, I will previous studies on feeding methods for heifers (DeVries
review the work on different feed delivery methods, and von Keyserlingk 2009a; Greter et al. 2010a), con-
controlling the amount of feed provided, and varying centrate was provided only one time per day. It could be
stocking density at the feed bunk. As the amount of hypothesized that more frequent provision of concen-
literature in this area is limited, this review also identifies trate within a day would, as in mature dairy cattle
aspects of these, and other, nutritional management (French and Kennelly 1990), prevent large post-prandial
practices that require further research. drops in rumen pH; more research is needed to test
this hypothesis. Interestingly, feeding concentrate, in a
FEED DELIVERY METHODS
Total mixed rations (TMR) are designed as a homo- 3.00
genous mixture with the goal to help minimize the Choice
2.50
selective consumption of individual feed components Top dressed
and ingredients by dairy cattle, promote a steady-state 2.00
TMR
DMI (kg)
Grass hay
1.50
to a novel TMR (for a period of 7 wk). Moreover, the
Feed
delivery increased competition for feed observed in heifers fed
1.00 a top-dressed ration, compared with those fed a TMR,
0.50
was also maintained after they were all switched to the
novel TMR. These heifers previously fed the top dressed
0.00 treatment also maintained looser fecal consistency.
0:00 3:00 6:00 9:00 12:00 15:00 18:00 21:00 Interestingly, the feeding and competitive behaviour
Time (h) patterns persisted throughout the 7 wk of that experi-
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nutrient dense diet fed in a limited amount (Hoffman in the diet while feeding time and meal duration
et al. 2007; Zanton and Heinrichs 2007). Limit feeding increased (Table 1). This suggests that the addition of
allows for the effective control of average daily gain in straw might help meet the natural foraging behaviour
replacement heifers, and also effectively decreases fecal patterns of these heifers. Interestingly, DMI also de-
excretion, increases feed efficiency, and reduces feed creased with the addition of the straw to the ration.
costs in some cases. Unfortunately, limit feeding does Based on these intakes, these researchers found that all
pose behavioural concerns for dairy heifers. Hoffman requirements for maintenance and growth of 1.0 kg d1
et al. (2007) found that limit feeding reduced eating and could be sufficiently met with 10% straw in the diet.
lying time, resulting in animals spending more time Further, they found that sufficient nutrients were
standing while not eating, a known risk factor for consumed with 20% straw in the diet to meet the
lameness (Greenough and Vermunt 1991). These re- requirements to achieve a 0.9 kg d1 growth rate.
searchers also found that limit feeding increased voca- Overall, these findings suggest that, if a ration is
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lization levels in heifers (Hoffman et al. 2007), as well as balanced properly, the addition of an inexpensive, low-
increased the amount of aggressive ‘reaching’ to acquire nutritive feedstuff may help reduce DMI and enable
feed (Hoffman 2007). Limit feeding dairy cattle has also producers to target caloric intake for desirable weight
been associated with increased levels of oral stereo- gain and development while allowing heifers to engage
typies, including tongue rolling, constant head nodding, in more natural foraging behaviour. As the Greter et al.
and bar biting/licking (Redbo et al. 1996; Redbo and (2008) study was not designed to evaluate the long-term
Nordblad 1997; Lindström and Redbo 2000). The effects on growth rate, it is recommended that further
changes in behaviour associated with limit feeding may research be conducted to validate these potential effects.
be attributed to hunger and frustration as a result of This research should also consider whether the cost of
lack of satiety (Watts and Stookey 2000; Valizadeh et al. gain changes with such a feeding strategy. Further
2008). The lack of satiety observed in limit-fed heifers research is also needed to determine if the increased
results from not only having feed available in a limited feed efficiency and reduced feed cost gained from a
amount, but also for a very short duration. The 1 to limit-fed diet could be maintained if such a diet were
2 h of feeding duration observed in limit-fed heifers supplemented with a low-nutritive feedstuff to help
For personal use only.
(Hoffman et al. 2007) is a stark contrast from the 49 h satisfy the natural foraging needs of replacement dairy
that dairy cattle, under natural grazing conditions, heifers.
would engage in foraging behaviour throughout the Another possible way to reduce the behavioural effects
day (Hafez and Bouissou 1975). Moreover, there is of limit feeding would be to feed smaller portions of the
potential for some of these behavioural effects to result
ration more frequently throughout the day. Providing
in negative health implications. For example, increased
fresh feed more frequently has been shown to cause
time spent standing, particularly on hard flooring
mature dairy cattle to distribute feeding behaviour more
surfaces, may increase the risk of hoof pathologies
(Cook et al. 2004; Vanegas et al. 2006). Further, the
consumption of a highly fermentable ration, when eaten Table 1. Dry matter intake, energy intake and measures of feeding
rapidly and ruminated less, may cause greater within- behaviour of growing dairy heifers fed a diet with added straw. [Adapted
and modified from Greter et al. (2008). Reproduced with the permission
day bouts of sub-acute ruminal acidosis (Moody et al. of Journal of Dairy Science]
2007), which are known to be detrimental to health and
feed efficiency. Experimental dietsz Effects
An alternative to limiting the amount of feed pro-
vided would be to limit the nutrient density of a feed 10% 20%
Item Control straw straw SE Linear Quadratic
offered ad libitum (Hoffman et al. 1996). One possible
1
method is to add a low-nutritive, low-value feedstuff to DMI (kg d ) 8.1 7.1 6.2 0.28 B0.001 NSy
the diet that would satisfy the natural feeding behaviour ME 21.0 17.2 14.6 0.86 B0.001 NS
(Mcal d 1)
patterns of limit-fed animals, as feeding duration would Feeding time 180 193 199 13 0.05 NSy
be increased. In addition to achieving satiety, this (min d 1)
feeding method may also reduce the risk of sub-acute Feeding rate 0.06 0.04 0.04 0.004 0.002 0.12
ruminal acidosis. A low-nutritive feedstuff will decrease (kg min1)
Meal frequency 9.6 8.4 8.7 0.92 0.03 0.03
passage rate and increase rumination time, thus increase (no. d 1)
saliva production and rumen buffering. Dietary dilution Meal duration 38.5 42.0 43.4 3.45 0.03 NS
will also reduce volatile fatty acid production through (min meal 1)
reduced concentration of rapidly fermentable carbohy- Meal size 0.9 0.9 0.7 0.09 0.04 NS
drates in the ration. In a recent study by Greter et al. (kg meal1)
(2008), straw was added (by 10 and 20% of DM) to a z
Control17.0% corn silage, 52.1% grass silage, 30.9% concentrate;
TMR fed ad libitum to replacement dairy heifers. These 10% strawcontrol diet with 10% rye straw; 20% strawcontrol diet
researchers found that daily DMI, feeding rate, meal with 20% rye straw.
y
size, and meal frequency decreased with increased straw NSP0.15.
DE VRIES * DAIRY HEIFER NUTRITIONAL BEHAVIOUR 299
equally throughout the day (DeVries et al. 2005). down to 15 cm per heifer), resulted in heifers spending
Spreading feed intake throughout the day through 25 to 50% less time eating. González et al. (2008) also
multiple feedings may not only help satisfy the natural demonstrated that high levels of feed bunk competition
feeding behaviour patterns heifers need during their day, (when comparing two, four, or eight heifers per feeding
thus diminishing hunger-related behaviour, but also have bin) reduced feeding time for heifers that were fed a high
positive effects on feed efficiency (Rakes et al. 1957; concentrate diet ad libitum. Interestingly, despite the
Nocek and Braund 1985; Shabi et al. 1999). Further reduction in feeding time, researchers have not reported
research is encouraged to address this hypothesis. any effect of competition for feed on the DMI of
replacement dairy heifers fed ad libitum (González
STOCKING DENSITY AT THE FEED BUNK et al. 2008; DeVries and von Keyserlingk 2009b) or,
The intensification of the dairy industry, along with less surprisingly, those that are limit fed (Keys et al.
rapid growth in herd sizes, has resulted in housing dairy 1978; Longenbach et al. 1999).
Can. J. Anim. Sci. Downloaded from www.nrcresearchpress.com by 105.104.230.235 on 05/17/20
cattle, including young stock, at higher densities (i.e., To maintain similar DMI in situations of high
over stocking). This management practice is often competition, heifers compensate by eating faster
justified, as it can be argued that heifers spend only a throughout the day, particularly during periods of peak
small fraction of their day consuming feed (3 to 3.5 feeding activity, and by shifting their intake patterns
h d1; Greter et al. 2008) and, thus, it could be assumed such that a greater proportion of their DMI occurs in the
that the provision of a feeding place for each individual later hours after feed delivery (Fig. 3; González et al.
heifer within a pen is not necessary. However, cattle tend 2008; DeVries and von Keyserlingk 2009b). Competition
to synchronize their behavior, that is, many animals in
for feed access also appears to change the meal pattern-
the group will feed, ruminate, and rest at the same times
ing of heifers, resulting in consumption of fewer meals
(Miller and Wood-Gush 1991; Rook and Huckle 1995).
per day, which are larger and longer in duration (DeVries
When cattle are fed in groups, the initiation to feed by
one animal will often stimulate the other animals, and von Keyserlingk 2009b). Given that within-day
regardless of whether they show signs of hunger (Curtis rumen pH declines increase with meal size (Allen 1997),
and Houpt 1983). Further, synchronized peaks in large and long meals may have significant impact on
rumen fermentation. Results from González et al. (2008)
For personal use only.
Feed Feed
activity. Alternatively, the synchrony between animals 0.90
delivery delivery
has the potential to break down in such situations of 0.60
high competition, and may lead to animals feeding at
different times to avoid excessive aggression (Miller and 0.30
Wood-Gush 1991). These effects of feed competition, 0.00
with their associated affects on behaviour, growth, and 0:00 3:00 6:00 9:00 12:00 15:00 18:00 21:00
health of replacement dairy heifers fed under various
feeding strategies, are documented in a growing body of (b) 0.08
Feeding rate (kg/min)
literature.
0.06
Competition for feed appears to have similar effects,
across feeding strategies, on the feeding behaviour of 0.04
replacement dairy heifers. DeVries and von Keyserlingk
(2009b) recently found that ad libitum fed heifers 0.02
exposed to high feed bunk competition (two heifers
per feed bin) tended to have 10% shorter daily feeding 0.00
times (192 vs. 213 min d1) than heifers with no feed 0:00 3:00 6:00 9:00 12:00 15:00 18:00 21:00
bunk competition (one heifer per feed bin). Similarly, Time (h)
decreases in feeding time have also been reported in
studies on limit-fed heifers. Keys et al. (1978) found that Fig. 3. Hourly averages for (a) DMI (kg) and (b) feeding
increasing pen stocking density and, thus, decreasing rate (kg min 1) for growing dairy heifers (8 mo old, weighing
235 kg) fed a TMR ad libitum noncompetitively (one heifer
feed bunk space (from 0.81 to 0.20 m per heifer) for per feed bin; n 12) or competitively (two heifers per feed bin;
yearling heifers linearly decreased their time spent eating n 12). Daily DMI averaged 6.2 and 6.3 kg d 1 (SE0.3)
by 26%. In a series of experiments on feed bunk space, and feed rate averaged 0.038 and 0.047 kg min 1 (SE0.004)
Longenbach et al. (1999) reported that heifers (ranging for the noncompetitive and competitive treatments, respec-
from 4 to 21 mo of age) subjected to increasing com- tively. [Modified from DeVries and von Keyserlingk (2009b).
petition (by reducing feed bunk space from up to 47 Reproduced with the permission of Journal of Dairy Science.]
300 CANADIAN JOURNAL OF ANIMAL SCIENCE
greater feed bunk competition for concentrate in heifers, of sub-acute ruminal acidosis. Future research should
fed roughage ad libitum separately, resulted in lower address how moderate overstocking and social status
rumen pH, greater lactate concentration, increased interact to affect feed access and maintenance of
serum haptoglobin levels, and increased proportion of nutrient intake in ad libitum and group-fed replacement
abscessed livers. For heifers fed a higher forage ration ad dairy heifers.
libitum, these rumen effects are likely far less extreme
with the exception of situations where excessive sorting CONCLUSIONS
of the ration is observed. Sorting against long fiber There is a growing body of literature on dairy cattle
particles by heifers (Hoffman et al. 2006, Greter et al. behaviour and how this field of science can aid in
2008; DeVries and von Keyserlingk 2009a, b), could making science-based recommendations for best man-
exacerbate any reductions in rumen pH caused by agement practices. For replacement dairy heifers, much
changes in meal pattern. Alternatively, increased daily of this research has been focused on the interactions
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time spent eating through reduced competition for feed between nutritional management and behaviour, parti-
may increase total salivary secretion (Beauchemin et al. cularly how these relate to different feed delivery
2008) and thus be beneficial for rumen fermentation and methods, the amount of feed provided, and stocking
fibre degradation (Owens et al. 1998). Unfortunately, density at the feed bunk. This research has provided us
there are no data to date to support these hypotheses on with a basic understanding of how nutritional manage-
the effects of feed competition on rumen fermentation ment practices may affect heifer behaviour, and how
for replacement heifers that are fed higher forage rations these practices may affect the health, productivity, and
ad libitum or limit fed lower forage rations, and thus welfare of these animals. Future research is needed to
further research in this area is required. further demonstrate the physiological consequences of
Another concern when feed bunk competition is high changes in behaviour that may result from these
for replacement dairy heifers is the variable effect it nutritional management practices. This information is
has on individual animals kept in groups. Recent work needed so that further recommendations can be made
has demonstrated that competition for feed in dairy on these practices so these animals may be raised for
heifers increases the day-to-day variation in meal maximum production potential, while maintaining their
For personal use only.
duration, feeding time and meal size (DeVries and von behavioural needs, health and overall welfare.
Keyserlingk 2009b), resulting in inconsistent feeding
behaviour patterns. González et al. (2008) also found
that the variability in feeding time and body weight ACKNOWLEDGEMENTS
tended to increase in groups of heifers fed competitively, The paper reviews concepts and data largely generated
suggesting disparity between group members in their in collaboration with colleagues, particularly Marina
ability to access feed and maintain DMI. Similarly, von Keyserlingk, at the University of British Columbia
Longenbach et al. (1999) reported that competition Animal Welfare Program. The research was funded, in
increased the variability in live weight gain within pens part, by a Natural Sciences and Engineering Research
of replacement dairy heifers. It has been demonstrated Council of Canada (NSERC, Ottawa, ON, Canada)
in heifers that competition for feed is most intense Discovery Grant awarded to T. J. DeVries and through
during the first hour after feed delivery, with these a NSERC Collaborative Research Development Grant
effects greatest in situations where competition for feed with the Dairy Farmers of Canada (Ottawa, ON,
bunk access is increased as the number of animals able Canada).
to feed simultaneously decreases (Keys et al. 1978). The
variability in feeding behaviour and weight gain ob- Allen, M. S. 1997. Relationship between fermentation acid
production in the rumen and the requirement for physically
served in these studies may be explained, in part, by
effective fiber. J. Dairy Sci. 80: 14471462.
certain animals dominating the feed bunk after feed Beauchemin, K. A., Maekawa, M. and Christensen, D. A. 2002.
delivery and consuming excess DM. At the same time, Effect of diet and parity on meal patterns of lactating dairy
this likely increases the risk that subordinate animals cows. Can. J. Anim. Sci. 82: 215223.
show reduced DMI; further research is warranted to Beauchemin, K. A., Eriksen, L., Norgaard, P. and Rode, L. M.
substantiate this explanation. Further, sorting of an ad 2008. Short communication: Salivary secretion during meals in
libitum fed ration could also contribute to this varia- lactating dairy cattle. J. Dairy Sci. 91: 20772081.
bility. Excessive sorting by dominant animals can reduce Borland, K. and Kesler, E. M. 1979. Complete rations for
the nutritive quality of the feed that is available to those Holstein calves 8 to 18 weeks of age. J. Dairy Sci. 62: 304309.
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Environmental influences on claw horn lesions associated
(DeVries et al. 2005; Hosseinkhani et al. 2008). As a
with laminitis and subacute ruminal acidosis in dairy cows.
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maintain adequate nutrient intake to meet their energy Coppock, C. E. 1977. Feeding methods and grouping systems.
and essential nutrient requirements, while the dominant J. Dairy Sci. 60: 13271336.
heifers that preferentially consume the concentrate Coppock, C. E., Bath, D. L. and Harris, B. 1981. From feeding
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K. A. 2003. Short communication: Diurnal feeding pattern of R., Veira, D. M. and von Keyserlingk, M. A. G. 2008. The
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