Review: Feeding behaviour of dairy cattle: Meaures

and applications
M. A. G. von Keyserlingk and D. M. Weary
Animal Welfare Program, University of British Columbia, 2357 Mall, Vancouver, British Columbia, Canada V6T 1Z4
(e-mail: marina.vonkeyserlingk@ubc.ca). Received 22 December 2009, accepted 10 June 2010.
von Keyserlingk, M. A. G. and Weary, D. M. 2010. Feeding behaviour of dairy cattle: Meaures and applications. Can. J.
Anim. Sci. 90: 303
309. There is growing scientific interest in feeding behaviour of dairy cattle, in part because dairy
nutritionists are now becoming interested in how changes in feed intake are mediated by changes in behaviour and, in part,
Can. J. Anim. Sci. Downloaded from www.nrcresearchpress.com by 105.104.84.101 on 12/04/19

because changes in feeding behaviour are increasingly recognized as useful indicator of cow health. In this review we
describe key methodological approaches to the study of feeding behaviour in dairy cattle. We also review empirical work
addressing how changes in management and housing can affect this behaviour. We show how cows divide their daily intake
into several discrete feeding events made up of a number of visits or ‘‘meals’’ that are separated by longer periods with little
feeding activity. Feeding behaviour can be described using several measures, including the number and duration of meals,
as well as intake and feeding rate. Feeding behaviour within a group of intensively managed cows is often highly
synchronized, similar to that seen in extensively housed cattle, with delivery of fresh feed appearing to be the primary
factor stimulating feeding by housed dairy cows. Competition at the feed bunk can affect feeding behaviour, increasing the
feeding rate and reducing intake, especially for subordinate animals. We also review empirical work showing that feed
intake, feeding times, and feeding rate are altered when cows are ill. Feeding behaviour changes in the days before calving,
and these changes are greatest among cows at greatest risk of succumbing to disease in the early post partum period. These
results suggest that monitoring changes in feeding behaviour may be useful in early detection and prevention of disease in
transition cows.

Key words: Dairy cattle, feeding behaviour, management, health

For personal use only.

von Keyserlingk, M. A. G. et Weary, D. M. 2010. Le comportement alimentaire des bovins laitiers: quantification et
applications. Can. J. Anim. Sci. 90: 303
309. Les chercheurs s’intéressent de plus en plus aux habitudes alimentaires des
bovins laitiers, en partie parce que les nutritionnistes souhaitent voir de quelle manière les changements de comportement
affectent l’ingestion des aliments et en partie parce qu’on voit de plus en plus dans la modification des habitudes
alimentaires un indicateur utile de la santé de l’animal. Dans cet article, les auteurs décrivent les principales approches
méthodologiques à l’étude du comportement des bovins laitiers sur le plan de l’alimentation. Ils passent également en revue
les travaux empiriques indiquant de quelle manière les modifications au niveau des pratiques d’élevage et du logement des
animaux affectent ce comportement. Les auteurs montrent comment les vaches divisent leur alimentation journalière en
segments composés de visites ou « repas » que séparent de plus longues périodes d’inactivité. On peut décrire les habitudes
alimentaires en recourant à diverses mesures, notamment le nombre et la durée des repas ainsi que le taux d’ingestion et
d’alimentation. Dans les groupes de vaches laitières étroitement élevées, le comportement alimentaire s’avère souvent
synchronisé, un peu comme cela se produit chez les bovins en claustration, l’arrivée de nourriture fraı̂che semblant être le
principal facteur qui stimule la prise d’aliments. La compétition à l’auge modifie parfois le comportement pour accroı̂tre le
taux d’alimentation tout en réduisant la quantité d’aliments ingérée chez les animaux subordonnés. Les auteurs ont aussi
examiné des travaux empiriques indiquant qu’il y a modification de l’ingestion des aliments, des périodes d’alimentation et
du taux d’alimentation quand les vaches sont malades. Les habitudes alimentaires changent peu avant le vêlage, et les
modifications les plus importantes surviennent chez les animaux qui courent le plus de risques de succomber à la maladie
durant la période suivant immédiatement la mise bas. Ces résultats laissent croire qu’en surveillant les changements au
comportement alimentaire, on dépisterait et préviendrait plus facilement la maladie chez les vaches en transition.

Mots clés: Bovins laitiers, comportement alimentaire, zootechnie, santé

Much of the recent research on dairy cattle nutrition Andersen (2000)]. Identifying these physiological factors
has focused on metabolic and nutritional factors that has proven helpful in formulating rations, including
contribute to the physiological regulation of feed in- those for early lactation cows whose energy demands are
take [see reviews by Allen (2000) and Ingvartsen and difficult to meet (Ingvartsen and Andersen 2000).

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; NDF, neutral detergent fibre
303
 304 CANADIAN JOURNAL OF ANIMAL SCIENCE

However, feed intake is mediated by the cow’s beha-

viour, including where, when and how she eats the diet
provided (Grant and Albright 1995). Surprisingly little
research to date has focussed on feeding behaviour,
including the effects of housing, management, and
environment on motivation and ability of dairy cattle
to access feed.
In this paper we describe methodological approaches
to the study of feeding behaviour in dairy cattle and
review recent empirical work addressing how changes in
management and housing affect this behaviour. One
practical goal of this research is identifying factors that Fig. 1. Cumulative daily dry-matter intake by one mid-
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can reduce a cow’s ability to access feed. A second area
of application is to use changes in feeding behaviour as
an early indicator of disease. Meeting these applied
goals requires an understanding of how feeding beha-
viour can be measured, so our review begins with this
topic. Drawing mostly upon examples from our own
work, we describe the components of feeding behaviour
in dairy cattle, review studies on how feeder design and
management affect behaviour, and how changes in the
behaviour can be used in detecting disease. For a more
general introduction to conceptual issues in the study of
feeding behaviour we refer readers to Nielsen (1999).
MEASURING FEEDING BEHAVIOUR
For personal use only.
lactation Holstein dairy cow.
total daily dry matter intake (DMI) can be read as the
maximum on the y axis (27.7 kg); total time spent
feeding can be calculated by summing the duration of
the nine meals (249 min). These values can then be used
to calculate derived measures of feeding behaviour, such
as feeding rate calculated on a per meal or daily basis.
Over 24 h the patterning of feeding events into meals
is clear, but over shorter periods it can be more difficult
to clearly assign individual feeding events into meals.
Cows may withdraw their head from the feed bunk
while chewing, and move along the feed bunk before

The feeding behavior of most animals can be recorded taking their next bite. Cows are sometimes displaced
as events that include bites or visits to a feeder (Mayes from their position at the feed bunk by another cow,
and Duncan 1986; Nielsen 1999). Visits to a feed source forcing them to wait or to move to a new location. Cows
or feeder are usually interspaced by numerous short may also leave the bunk for several minutes, perhaps to
intervals that in turn are interspaced by long intervals. engage in social interactions with pen mates or to visit
The availability of technologies such as the electro- the water trough. Cows may also leave the feeder to lie
nic feed recording system (INSENTEC, Marknesse, down in a stall, in which case they are likely to be away
Holland) now allow researchers to continuously monitor from the feeder for a longer period. If visits to the feeder
these events at the feeder as well as how much each are separated by a variety of non-feeding intervals, how
individual cow consumes at each visit. With the Insentec can we objectively define when a meal begins and ends?
system, each cow is fitted with a unique passive One approach to this problem is to plot the frequency
transponder attached to her ear tag. When a cow distribution of intervals between visits to the feeder and
approaches the feed bin an antenna detects her trans- evaluate any discontinuities (see Tolkamp et al. 2000).
ponder and the head gate opens allowing her access to Figure 2 illustrates the frequency distribution of inter-
feed and the electronic system records the time and vals expressed on a log scale. This distribution is clearly
the initial weight in the bin. When a cow exits the bin, the bimodal, with the intersection between the two peaks
head gate closes and the system again records the
time and bin weight. These values are then used to
calculate the duration of each visit and the amount of
feed consumed.
A high-producing Holstein can eat more than 25 kg of
dry matter in a day. Figure 1 illustrates how one cow
patterned this intake over a 24-h period. Intake was
divided into several discrete feeding events or ‘‘meals’’
that were separated by longer periods when the cow
showed little feeding activity. The cumulative intake plot
can be used to visualize a variety of important measures
of feeding behaviour. For example, the number of
distinct meals can be seen as the number of steps on
the graph (nine meals), and the duration and intake Fig. 2. Frequency distribution of the intervals (n 5911)
during each of these meals can be calculated from the between the feeding visits to the feeder for 20 early-lactation
displacement on the x and y axes, respectively. Similarly, Holstein cows recorded continuously for 10 d.
 VON KEYSERLINGK AND WEARY * DAIRY CATTLE FEEDING BEHAVIOUR
occurring at about 25 min. This break point can be used
to define within-meal intervals (represented by the peak
on the left with a maximum at about 0.5 min), and
between meal intervals (the peak on the right of the
graph with a maximum at about 180 min). According to
this approach an interval of less than the break point
(called ‘‘meal criterion’’) can be defined as within a
meal. For example, two feeding events separated by a
break of only 10 min would be considered by this
definition as being part of the same meal, but if the cow
returned to the feeder after a break of 50 min we would
consider this event to be part of a new meal. The meal
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criterion can also be identified mathematically by
modelling the two (or more) distributions and calculat-
ing where these distributions intersect (e.g., DeVries
et al. 2003a, 2009).
TEMPORAL AND SPATIAL VARIATION
With the measures outlined above we can begin to better
describe how feeding behaviour varies in time. Cattle
often show a crepuscular pattern in feeding activity,
with clear peaks around dawn and dusk (Shabi et al.
2005). Figure 3 provides clear illustration of diurnal
variation in feed bunk attendance for a group of
intensively managed lactating cattle, showing two clear
For personal use only.
peaks in feeding activity. Cows spent very little time
feeding late at night and early in the morning. Activity
peaked when fresh feed was first provided at about 0600
and again when fresh feed was provided at 1500.
In addition to the temporal variation described above,
spatial variation in feeding behaviour may also be
important. Spatial variation may be driven by social
factors. For example, Manson and Appleby (1990)
measured the spacing between adjacent cows at the
feed bunk and found that this spacing was non-random,
at least when cow numbers were low and cows had some
opportunity to choose where to feed. Cows tended to
clump together at the feeder, but avoided positions
immediately adjacent to another cows. Cows very
different in social rank showed the greatest avoidance,
Fig. 3. The diurnal pattern of feeder activity for four groups
of 12 min-lactation Holstein dairy cows. Each group was
observed during a 10-d period with once a day feeding (at
0600) and 10 d with twice a day feeding (at 0600 and 1500)
using a cross-over design (see DeVries et al. 2005).
305
presumably because low ranked cows avoided feeding
beside high ranked individuals.
Unfortunately, little is known about other factors
affecting spatial variation in feeding. In our own
research we have noted that cows sometimes appear to
systematically move between feeding positions, appear-
ing to ‘‘graze’’ the feed bunk, a behaviour that seems
non-functional given that the same diet to provided at
all locations. Cows may be responding to subtle varia-
tion in feed quality associated with sorting or poor feed
mixing, but to our knowledge no work on cattle has
addressed this behaviour.
Feeding Management
Figure 3 also illustrates the effects of feeding manage-
ment of the diurnal feeding pattern: when cows were
provided fresh feed twice a day (at 0600 and 1500) there
were two clear peaks in bunk attendance, showing that
cows respond to availability of fresh feed at the feed
bunk. Work by our group has shown that a primary
factor stimulating housed dairy cattle to feed is fresh
feed delivery (DeVries and von Keyserlingk 2005). There
is also a peak in feeding after cows return from the
milking parlour, but this effect is minor compared to
the delivery of fresh feed.
One common management practice believed to sti-
mulate feeding activity is feed push-up. When fed a total
mixed ration (TMR), dairy cows have a natural
tendency to sort through the feed and toss it forward
where it is no longer within reach. Producers commonly
push the feed closer to the cows in between feedings to
ensure that cows have continuous feed access. In an
observational study, Menzi and Chase (1994) noted that
the number of cows feeding increased after feed push up
and concluded that feed push-ups had ‘‘minor and brief
effects’’ on the feed bunk attendance. In a more recent
controlled study, we tested the stimulatory affect of feed
push-up by increasing the number of push-ups during
the late evening and early morning (DeVries et al.
2003b). If some feed was already available at the feed
bunk, the act of pushing up extra feed had little or no
affect on the cow behaviour.
Physical Design of the Feed Barrier
There are several aspects of the feeding environment
that affect the cow’s ability to access feed, including the
amount of available feed bunk space per animal and
the physical design of the feeding area. When cows are
forced to compete for access to food, socially dominant
cows spend more time eating than do herd-mates lower
in social rank (Manson and Appleby 1990). Reducing
the space available for cows to eat increases competition
(Mentink and Cook 2006). For example, DeVries et al.
(2004) showed that doubling feeding space from 0.5 m to
1.0 m per cow reduced by half the number of aggressive
interactions while feeding. This reduction in aggressive
behaviour allowed cows to increase feeding activity by
 306 CANADIAN JOURNAL OF ANIMAL SCIENCE
24% at peak feeding times, an effect that was strongest
for subordinate animals.
The physical design of the feeding area can also
influence feeding behaviour. One of the most obvious
features of the feeding area is the physical barrier that
separates the cow and the feed, and research shows how
some designs can reduce aggressive interactions at the
feed bunk. Endres et al. (2005) compared the effects of a
post-and-rail versus a headlock feed line barrier on the
feeding and social behaviour of dairy cows and found
that during periods of peak feeding activity (90 min after
fresh feed delivery) subordinate cows had lower feeding
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times when using the post-and-rail barrier. This differ-
ence was likely due to positive effects of the headlock
barriers in reducing competitive interactions; there were
21% fewer displacements at the feed bunk when cows
accessed feed by the headlock barrier compared with the
post-and-rail barrier.
Huzzey et al. (2006) retested the effects of these two
types of feed bunk barriers over a range of stocking
densities: 0.81, 0.61, 0.41 and 0.21 m/cow (correspond-
ing to 1.33, 1.00, 0.67 and 0.33 headlocks/cow). Daily
feeding times were higher and the duration of inactive
standing in the feeding area (waiting to gain access to
the feed bunk) was lower when using a post-and-
rail compared with a headlock feed barrier. As well,
For personal use only.
regardless of barrier type, feeding time decreased and
inactive standing increased as stocking density at the
feed bunk increased. Cows were displaced more often
from the feeding area when the stocking density was
increased, and this effect was greater for cows using the
post-and-rail feed barrier. This effect was again greatest
for the subordinate cow, particularly at high stocking
densities.
More recent work has also investigated how competi-
tion affects feeding behaviour. Hosseinkhani et al.
(2008) followed the feeding behaviour of non-lactating
Holstein cows (n 36) eating TMR from either their bin
(one cow/bin) or a shared bin (two cows/bin). Cows that
had to compete for feed access increased their feeding
rate throughout the day. These cows also had fewer
meals, which tended to be larger and longer than cows
with access to their own bin. Competition had little
effect on daily DMI, but changed the daily distribution
intake; higher intakes occurring during the hours after
feed delivery after much of the feed sorting had already
occurred. Proudfoot et al. (2009) investigated the effects
of competition using a similar design and reported that
competitively fed multiparous cows increased their
feeding rate and were more likely to be displaced from
the feed bunk compared with non-competitively fed
cows. Thus, overstocking the feed bunk increases social
competition, including displacements from the feeder,
resulting in poor access to feed.
The research reviewed above shows that headlock
feed barriers can reduce competitive displacements by
cows, even though this hardware was not designed for
this effect. Some work has now begun to address feeder
design changes specifically intended to reduce competi-
tion among cows. Providing partitions between feeding
stations (‘‘feed stalls’’; Fig. 4) provides additional
protection to cows while feeding (DeVries and von
Keyserlingk 2006). Feed stalls reduce aggression and
competitive displacements, effects that again are greatest
for subordinate cows. This reduction in aggression
allows cows to increase daily feeding time and reduce
the time they spent standing in the feeding area waiting
to access the feeder. Thus the provision of more bunk
space, particularly when combined with feed stalls,
improves access to the feed bunk and reduces competi-
tion at the feed bunk, especially for subordinate cows.
Feed stalls may also help reduce the between-cow
variation in the composition of the ration consumed
by preventing subordinate cows from being forced to
access the bunk only after dominant cows have sorted
the feed (DeVries et al. 2005). A less aggressive feeding
environment may also have longer-term health benefits;
cows engaged in aggressive interactions at the feed bunk
are likely at higher risk for hoof health problems
(Leonard et al. 1998). A final advantage of feed stalls
is that they can facilitate the use of alternative flooring
surfaces in the feeding area; access to a soft, dry
standing area at the feeder can improve hoof health
(Manske et al. 2002).
Sorting
Cows sort mixed rations, typically preferring the grain
component and discriminating against longer forage
(Leonardi and Armentano 2003). One effect of this
sorting behaviour is that cows with early access to fresh
feed may consume a much richer diet than was intended
(Stone 2004). Sorting against long fibre particles is also
associated with lower rumen pH (DeVries et al. 2008).
This effect may be particularly troublesome for early
lactation cows, where greater sorting of a higher
concentrate, lower fibre diet (DeVries et al. 2007,
2008), coupled with rapidly increasing DMI (Kertz
et al. 1991), can increase the risk of ruminal acidosis
Fig. 4. Feed stall installed on a post and rail feed barrier
system.
 VON KEYSERLINGK AND WEARY * DAIRY CATTLE FEEDING BEHAVIOUR
(DeVries et al. 2008). This in turn may result in
inconsistent DMI, poor feed efficiency, reduced feed
digestibility and protein synthesis, and increased risk of
disease.
Sorting can also reduce the nutritive value of the diet
remaining in the feed bunk, predominantly in the hours
after delivery of fresh feed (DeVries et al. 2005); for
example, neutral detergent fibre (NDF) increases
throughout the day as a result of feed sorting. Sorting
changes the forage to concentrate ratio of the remaining
feed, affecting the nutrient intake of animals that access
the feeder latter in the day (see Fig. 5). Increasing the
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frequency of feed delivery from once daily to twice daily
reduced the amount of sorting, but feeding four times
per day had no additional effect (DeVries et al. 2005).
Cows are able to sort rations even when eating at a high
rate (DeVries et al. 2007), perhaps because cows select
for particles that can be consumed more rapidly as well
as those that are more palatable (Pyke et al. 1977).
These results, coupled with the finding that increasing
the frequency of feed provision increases access to the
feed bunk, suggests that a higher frequency of feed
delivery reduces variation among cows in the quality of
food consumed. Subordinate cows are displaced from
the feed bunk less frequently when the group is fed more
often, indicating that these cows have greater access to
For personal use only.
feed, particularly fresh feed, when the frequency of feed
delivery is high.
Sorting is a part of feeding behaviour, but research on
sorting to date has focused on the consequences of the
behaviour (i.e., temporal variation in feed remaining in
the feeder) rather than the behaviour per se. We see this
as one important shortcoming in the literature, and
encourage new work focusing on the behaviours cow use
to physically sort the diets they are provided. We believe
that this more basic research may shed light on how diet
forage concentrate
100
75
% of TMR
50
25
0
Fresh 1x 2x 2x
Feed Orts
Fig. 5. Forage to concentrate ratio TMR and orts estimated
from the initial NDF content values for the TMR and the final
NDF content of the orts of feed delivered once per day (1),
twice per day (2 ) or four times per day (4 ) [adapted from
DeVries et al. (2005)].
307
could be better formulated to minimize the negative
effects of sorting, or perhaps even formulated to allow
cows the ability to express this behaviour without harm
to cow health.
Changes in Feeding Behaviour During Illness
Changes in feeding behaviour have long been used to
help identify when animals become ill [reviewed by
Weary et al. (2009)]. Sowell et al. (1998) reported some
of the first work on cattle showing that healthy feedlot
steers spent 30% more time at the feed bunk than
morbid steers, and a greater percentage of healthy steers
visited the feed bunk immediately following feed deliv-
ery. Recent research has shown that these changes can
also be useful in detecting illness in dairy cattle,
especially during the transition period when cows are
most vulnerable to metabolic and infectious diseases.
Figure 6 illustrates how patterns of feed intake differ for
healthy cows and cows diagnosed with metritis. The
most dramatic differences in the diurnal feeding pattern
occur during times of highest bunk attendance between
0600 and 1800. A number of recent papers have shown
that feeding behaviour in the days before calving may
play an important role in identifying cows at risk of
disease in the early post-partum period. For example,
Urton et al. (2005) showed that cows diagnosed with
acute metritis after calving spent less time feeding during
the prepartum period (day
12 to day
2 prior to
calving), perhaps because cows with lower intakes
have poorer immune function (Hammon et al. 2006).
Huzzey et al. (2007) found that cows diagnosed with
severe metritis 7 to 9 d post partum consumed less feed
and spent less time at the feed bunk during the 2-wk
period before calving, nearly 3 wk before the first
clinical signs of disease. The odds of have severe metritis
increased by 1.72 for every 10 min decrease in feeding
time during the week before calving, and the odds
increased nearly threefold for every 1 kg decline in DMI.
More recent work has also shown that similar declines
4x
Orts
Fig. 6. Diurnal feed intake of cows that remain healthy and
cows diagnosed with clinical metritis after calving (Healthy
n 45, Metritis n22) from 5 to 10 d after calving [adapted in
part from Huzzey et al. (2007)].
 308 CANADIAN JOURNAL OF ANIMAL SCIENCE
in feeding behaviour are observed in cows at risk of
sub-clinical and clinical ketosis (Gonzalez et al. 2008;
Goldhawk et al. 2009).
In the studies reviewed above it is not clear if the
changes in feeding behaviour increase the risks of illness
or if illness cause changes in behaviour; experimental
research is now required to separate cause and effect.
Such research may also help explain the correlation
reported in some previous research between measures
of feeding activity and milk production. Indeed, Shabi
et al. (2005) report that the correlation between feeding
behaviour and milk production is stronger than that
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between intake and production. We hypothesise that
this correlation is driven by ill animals having both low
production and depressed intakes.
CONCLUSIONS
Research to date on dairy cattle nutrition has focused
almost exclusively on the effects of the nutrients
provided. In this review we have shown that sophisti-
cated methods of describing feeding behaviour are now
available, and we have argued that knowledge of feeding
behaviour can provide insights that improve our ability
to feed intensively managed dairy cows. The question is
no longer just what do they eat, but how do they eat it?
For personal use only.
An improved understanding of feeding behaviour is
helping to avoid practical problems such as competition
at the feed bunk and excessive sorting. This research is
also showing how changes in management can have
facilitate feed access, particularly for the subordinate
cow, and how changes in feeding behaviour can be used
to evaluate cow health. More generally, we argue that
changes in feeding behaviour can provide insight into
how well a cow is able to function within the physical
and social environment she is provided.
ACKNOWLEDGEMENTS
We thank our past and current students in the Uni-
versity of British Columbia’s Animal Welfare Program,
in particular Katy Proudfoot, Julie Huzzey and Trevor
DeVries, for their hard work in undertaking much of the
research cited in this review. Funding for the University
of British Columbia’s Animal Welfare Program is
provided by Canada’s Natural Sciences and Engineering
Research Council (Ottawa, ON, Canada) Discovery
Grant Program and the Industrial Research Chair
Program with industry contributions from the Dairy
Farmers of Canada (Ottawa, ON, Canada), Westgen
Endowment Fund (Milner, BC, Canada), Pfizer Animal
Health (Kirkland, QC, Canada), BC Cattle Industry
Development Fund (Kamloops, BC, Canada), the
BC Milk Producers (Burnaby, BC, Canada), BC
Dairy Foundation (Burnaby, BC, Canada), BC Dairy
Education and Research Association (Abbotsford, BC,
Canada), and Alberta Milk (Edmonton, AB, Canada).
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