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Animal (2015), 9:6, pp 10381044 The Animal Consortium 2015

doi:10.1017/S1751731115000063
animal

Relationship between locomotor play of dairy calves and their


weight gains and energy intakes around weaning
G. G. Miguel-Pacheco1a A. Vaughan1,2, A. M. de Passill2 and J. Rushen2
1
Applied Animal Behaviour and Animal Welfare Programme, The University of Edinburgh, Easter Bush Veterinary Centre, Roslin EH25 9RG, UK; 2University of British
Columbia, PO Box 1000, 6947 Highway 7, Agassiz, BC, Canada V0M 1A0

(Received 24 September 2013; Accepted 11 December 2014; First published online 12 February 2015)

Play behaviour has been proposed as a measure of good welfare in growing animals and locomotor play by calves is often reduced
after weaning off milk. Adjusting weaning age according to individual calves abilities to eat solid feed maintains energy intake
and weight gain during weaning. We investigated the effects of this method of weaning on locomotor play of calves and the
relationship between locomotor play and energy intake and weight gains. We measured the running behaviour of 56 Holstein
heifer calves before and after weaning. Calves were housed in groups of eight, fed milk, grain starter and hay from automated
feeders. Weaning began when their voluntary intake of grain starter reached either 200 or 400 g/day, and weaning was completed
when starter intake reached either 800 or 1600 g/day. Before weaning, older calves ran less than young ones; and the duration of
running correlated with weight gains and digestible energy intake. Immediately after weaning, digestible energy intake and
locomotor play decreased but no correlation was observed between these variables. One week after weaning, the duration of
running was correlated with both energy intake and weight gain. Digestible energy intake increased but locomotor play continued
to decrease. The amount of running a calf does after weaning partly reects energy intake and weight gain, supporting
suggestions that locomotor play is good indicator of welfare and tness of growing animals. However, the decline in locomotor
play following weaning is not solely due to decreased energy intake.

Keywords: play, dairy calf, weaning, animal welfare

Implications jumping and bucking) is common in young calves (Reinhardt,


1980; Jensen et al., 1998) and is susceptible to many welfare
Play behaviour is common in young calves and is linked
relevant factors such as food availability (Krachun et al.,
to energy intake. Dairy calves fed milk ad libitum grow
2010), space allowance (Jensen and Kyhn, 2000; Mintline
considerably faster and display more play than those fed
et al., 2012), social contact (Duve and Jensen, 2011) and
1015% BW of milk per day. In current weaning procedures,
pain (Rushen and de Passill, 2012; Mintline et al., 2013). For
calves reduce their play. Adjusting weaning age according to
young and growing animals, play behaviour has been
an individuals ability to eat solid feed maintains energy
proposed as an indicator of good management practices
intake and weight gain during weaning. This may have
(Held and pinka, 2011).
positive implications on the welfare of calves, and play
For dairy calves, weaning off milk at the ages typical in dairy
behaviour may be useful as a welfare assessment tool for
production can be stressful (Weary et al., 2008) and can result
growing animals.
in reduced energy intake and poor growth (e.g. Sweeney et al.,
2010; Rushen and de Passill, 2012). Locomotor play is also
Introduction reduced at weaning but the correlation between energy intake
and locomotor play is variable (Krachun et al., 2010; Sweeney
Positive welfare indicators, such as play behaviour, highlight et al., 2010; Rushen and de Passill, 2012) and the relationship
the presence of good welfare (Boissy et al., 2007; Yeates and with growth rate is unknown. Adjusting the weaning age of
Main, 2008). Locomotor play (consisting mainly of running, calves according to their individual intakes of solid feed can
a
Present address: School of Veterinary Medicine and Science, University of
prevent the drop in energy intake and growth after weaning,
Nottingham, Sutton Bonington Campus, Leicestershire, LE12 5RG, UK. but results in greater differences between calves in weight

E-mail: svxgm@nottingham.ac.uk gains (WGs) during and immediately after weaning, and in the

1038
Locomotor play of dairy calves around weaning

age and duration of weaning (Roth et al., 2009; de Passill and and ADF = 33.6%) and water troughs (Insentec, Marknesse,
Rushen, 2012). However, it is not known how this method of the Netherlands). Weigh scales (Smart1, Westernscale Inc.,
weaning affects locomotor play. Port Coquitlam, BC, Canada) were located at the milk feeder
The aim of the study was to examine the effects on loco- and water trough. Automatic feeders were equipped with
motor play of weaning calves off milk according to their sensors that recognized each calfs ear tag, allowing feeders
intakes of solid feed. We examined whether locomotor play to record individuals intake and weight at each visit.
was maintained after weaning, and whether locomotor play
was related to (a) digestible energy (DE) intake, (b) pre- and Treatments and experimental design
post- weaning growth and (c) age and duration of weaning. Once calves were moved to group pens, animals were assigned
to one of the four weaning treatments balancing birth weight
and age across treatments (de Passill and Rushen, 2012);
Material and methods resulting in seven experimental pens with eight calves in each
(two calves per treatment per pen). Treatment groups were
The study was carried out in compliance with the Canadian
based on starter intake target at the beginning and end of
Council on Animal Care Guidelines and approved by the
weaning following a 2 2 design (Low-High, Low-Low, High-
Institutional Animal Care Committee.
Low and High-High). For two groups, weaning started when
average grain consumption over 3 consecutive days was equal
Animals, housing and diet to or greater than 200 g/day (Low start). The remaining two
We used 56 Holstein heifer calves (mean birth weight 42.7 kg; groups started weaning when average grain consumption
range 30.0 to 56.1 kg) that were separated from their dams reached 400 g/day over 3 consecutive days (High start). Milk
within 6 h of birth. Each calf was housed in individual pens was reduced accordingly to grain consumption. Weaning was
(1.22 2.44 m) on wood shaving. They were bottle-fed completed when average grain consumption over 3 con-
ad libitum with whole milk twice daily and had constant secutive days was 800 g/day (low end) or 1600 g/day (High
access to water in buckets. At 6 1 days of age, the calves end). If calves had not begun weaning by 74 days of age due to
were moved to group pens (7.08 4.74 m) (Figure 1). Pens had insufcient grain consumption, the milk feeder automatically
slatted metallic oors (2.48 4.60 m) and a sawdust bedded weaned them over a 7-day period. This new group was named
area (4.6 4.74 m). Calves had access to automated feeders computer weaned.
(CF 1000 CS Combi, DeLaval Inc., Tumba, Sweden) that pro-
vided whole milk (12 l/day at 40C; herd averages of 3.97%
fat, 4.10% protein and 3.30% lactose from bulk tank samples)
Measurements
and grain starter (ad libitum; CP = 17.9% on DM basis; main
Behavioural observations. Locomotor play was assessed using
continuous observation between 0800 and 2300 h for a total
ingredients: 16% wheat, 14% barley, 13% canola meal,
duration of 15 h per day. No observations were made between
13% oats, 10% soya, 105 corn and 4% molasses, Unifeed
2300 and 0800 h as locomotor play rarely occurs during this
Ltd., Chilliwack, BC, Canada). They also had ad libitum
period (Krachun et al., 2010). We recorded the start and end
access to hay (DM = 90.8%, CP = 15.1%, NDF = 51.1%
time of each bout of running, enabling measurement of total
duration and frequency of running bouts. Calves were observed
for 2 days from each of three periods related to weaning (as
dened in Table 1), totaling 6 days of behavioural observa-
tions. Five of the 6 observed days were chosen randomly within
each period, the remaining observed day (immediately post
weaning period: 1st day of no milk) was xed for all the calves.
We dened locomotor play as running: that is a rapid forward
movement that may include jumping, bucking and/or
kicking with one or two legs (Jensen et al., 1998; Jensen and
Kyhn, 2000), lasting a minimum of 3 s (Krachun et al., 2010).

Table 1 Observation periods


Period Description

PreW Before weaning 5 days before the beginning


of weaning
PW1 to 3 Immediately post weaning First 3 days of no milk
(weaned)
PW7 to 9 Later post weaning Between the 7th to 9th day
Figure 1 Layout of experimental group pen. after weaning complete

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Miguel-Pacheco, Vaughan, de Passill and Rushen

Bucking and jumping that occurred independently of running respectively (Bodycote Testing Group Inc. Lethbridge, AB,
were not recorded, since the duration of these behaviours was Canada). BW and daily WG were calculated per day and
very short. If a calf did two running bouts that were <3 s apart, averaged per observation period. Age of the calves for the
they were recorded as part of the same bout. before weaning period were based on the age at the beginning
Three video cameras (Panasonic WV-BP 334; Panasonic, of this period (i.e. on the 1st day of the 5 days previous to the
Osaka, Japan) connected to digital video recorders were rst reduction of milk). Data for the duration of weaning, age
located 6 m vertically above each group pen to record calves at which weaning began and at which was completed were
behaviour. Two red lamps tted above the pens enabled used as in de Passill and Rushen (2012). Weaning duration
recording throughout the night. Two cameras were located was dened as total number of days taken for a calf to com-
over the feeding area and one over the bedding area. We plete weaning, from the rst decrease in milk intake to 0 milk.
identied the individual calves using pictures of each animal As the distributions of measures of running were highly
taken from different angles (front, top, back, left and right). positively skewed (Figure 2), non-parametric tests were used
Videos from the different cameras were viewed simulta-
neously using Ommicast software (Genetech Inc., Saint-
Laurent, Quebec) and EZViewLog500 software (Geovision
Japan Inc., Tokyo, Japan) at 4 normal speed. There were
four observers, with each observer watching two group pens
and no overlap between the observers.

Reliability. To assess intra- and inter-observer reliability, all


observers watched a sample video twice. In addition, we
assessed the reliability half way through the observations for
each group. Correlations for the intra- and inter-observer
reliability ranged between 0.90 to 0.99 (Median = 0.98) and
0.90 to 0.99 (Median = 0.95), respectively.

Statistical analysis
All statistical analyses were conducted using Stata/SE 12.0.
(Stata Corp 2011, Texas, USA). Data from ve calves were
excluded from the analysis: two calves were weaned too early
due to technical problems with the milk feeder, two calves had
health problems requiring veterinary treatment and one calf
died. Therefore, the analysis included only data from 51
calves; nine calves from the High-High treatment, nine calves
from High-Low, 11 calves from Low-High, nine calves from
Low-Low and 13 calves from computer weaned group.
Weaning according to grain intake affected duration of
weaning and when weaning started and ended (de Passill and
Rushen, 2012). Thus, this paper examines the relationship
between DE intake, growth, age at weaning and weaning
duration and measures of locomotor play. The total daily
duration and the frequency of running bouts were calculated as
the average of the values in the 2 days of each of the three
observation periods (Table 1). The mean bout duration for each
calf on each day was calculated by dividing the total duration
by the frequency of bouts, and then averaged across the
2 observed days for each period. In order to study the rela-
tionship between energy intake and locomotor play, DE intake
was calculated for the present study according to the observed
days. Raw data for DE intake, BW and daily WG per day were
obtained from de Passill and Rushen (2012) and adjusted
according to the observation periods of the present paper. The
mean daily DE intake for each calf was calculated from their
intakes of milk, grain and hay, averaged across the 2 obser-
vation days and divided by the calfs BW (MJ/kg). The DE for Figure 2 Duration (a), frequency (b) and average bout duration (c) of
milk was considered to be 5.59 Mcal/kg (NRC, 2001); analysis running at each observation period: before weaning (PreW), immediately
of grain and hay gave DE of 3.48 and 2.40 Mcal/kg, post weaning (PW1 to 3) and later post weaning (PW7 to 9).

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Locomotor play of dairy calves around weaning

to analyse all data (Martin and Bateson, 2010). Kruskal between the three measures of running, duration of weaning
Wallis test (Petrie and Watson, 2006) was used to compare and age at which weaning began and was completed. To
locomotor play measurements between weaning treatments reduce the effect of differences between groups of calves, we
per period and to compare age at when weaning started and ranked the values for all variables within groups before cal-
ended, and the duration of weaning between weaning culating the correlations. Results are presented as 25th per-
treatments. Wilcoxon test (Petrie and Watson, 2006) was centile Median 75th percentile and a P-value of 0.05
used to test for differences between treatments in the age at was specied to indicate signicant differences.
which weaning started, ended and weaning duration; and to
test for differences between consecutive periods of obser-
Results
vation in the total running duration, frequency of running
bouts, mean bout duration of running, and DE intake. We Effect of weaning treatment groups
used Spearmans correlations (Petrie and Watson, 2006) to No signicant differences between the weaning treatment
examine the relationship between the three measure of groups in each period were observed for total duration, fre-
running WG and DE intake in each period. Before weaning, quency and mean bout duration of running (Table 2). Milk,
Spearmans correlations were also calculated between the grain and hay DE intake per kilogram of BW before weaning
three measures of running, age of the calves at this period are shown in Table 3. Weaning treatments affected the age
and birth weight of the calves. Correlations were also cal- when weaning started (P < 0.001), ended (milk 0, P < 0.001)
culated in the immediately post-weaning period (PW1 to 3) and the duration of weaning (P < 0.001) (Table 3).

Table 2 Median and inter-quartile range (IQR) for duration, frequency and mean bout duration of running before weaning (PreW), immediately post
weaning (PW1 to 3) and late post weaning (PW7 to 9) according to weaning treatment groups
Weaning treatment

Measures of running HH HL LH LL CW P

Running before
Duration (s/day) 49.0 (31.0 to 66.0) 86.5 (12.5 to 93.0) 62.5 (36.5 to 111.5) 78.0 (25.0 to 115.0) 36 (23.5 to 68.0) 0.49
Frequency (bouts/day) 4.5 (3.5 to 13.0) 11.0 (2.5 to 18.0) 12.0 (6.0 to 15.5) 10.5 (3.0 to 14.5) 4.5 (3.5 to 7.5) 0.77
Mean bout duration (s/bout) 7.7 (6.9 to 9.7) 5.1 (4.8 to 6.6) 6.2 (5.2 to 7.7) 7.4 (5.4 to 7.9) 6.1 (5.5 to 7.5) 0.41
Running PW1 to 3
Duration (s/day) 0 (0 to 2.5) 0 (0 to 5.0) 4.5 (0 to 6.5) 0 (0 to 3.0) 0 (0 to 3.0) 0.37
Frequency (bouts/day) 0 (0 to 0.5) 0 (0 to 1.0) 1.5 (0 to 1.5) 0 (0 to 0.5) 0 (0 to 0.5) 0.61
Mean bout duration (s/bout) 0 (0 to 4.0) 0 (0 to 5.0) 3.0 (0 to 4.3) 0 (0 to 6.0) 0 (0 to 4.0) 0.37
Running PW7 to 9
Duration (sec/day) 16.0 (4.0 to 21.0) 26.5 (18.0 to 64.0) 17.5 (11.0 to 67.0) 23.0 (18.5 to 33.5) 16.5 (3.5 to 30.5) 0.26
Frequency (bouts/day) 1.5 (1.0 to 4.0) 4.5 (1.5 to 10.5) 3.5 (2.0 to 10.0) 4.5 (3.5 to 5.0) 3.5 (1.0 to 6.0) 0.98
Mean bout duration (s/bout) 6.0 (4.0 to 6.3) 5.1 (4.8 to 6.1) 5.8 (3.7 to 6.2) 5.7 (4.6 to 6.7) 5 (4.0 to 6.0) 0.86
HH = High-High (weaning started at 400 g/day ended at 1600 g/day); HL = High-Low (weaning started at 400 g/day and ended at 800 g/day; LH = Low-High (weaning
started at 200 g/day and ended at 1600 g/day); LL = Low-Low (weaning started at 200 g/day and ended at 800 g/day); CW = computer weaned.

Table 3 Median and inter-quartile range (IQR) of total digestible energy (DE) intake per kilogram of BW and from each feed (milk, hay and grain)
before weaning; and the median (IQR) of age (days) at weaning (day of rst reduction of milk) and age (days) at 0 milk and duration of weaning
(days) according to treatment groups
Weaning treatment

Variables HH HL LH LL CW

DE intake (MJ/kg of BW) 0.4 (0.3 to 0.4) 0.4 (0.4 to 0.4) 0.4 (0.4 to 0.4) 0.4 (0.3 to 0.4) 0.4 (0.3 to 0.4)
Milk 0.3 (0.3 to 0.4) 0.3 (0.3 to 0.4) 0.4 (0.3 to 0.4) 0.4 (0.3 to 0.4) 0.3 (0.3 to 0.3)
Grain 0.1 (0.04 to 0.1) 0.1 (0.01 to 0.02) 0.03 (0 to 0.1) 0.03 (0.02 to 0.03) 0.01 (0 to 0.1)
Hay 0.01 (0 to 0.02) 0.02 (0.01 to 0.02) 0 (0.04) 0.02 (0.01 to 0.03) 0.03 (0.01 to 0.04)
Age at weaning (days) 43 (36 to 64) 55 (46 to 60) 50 (31 to 60) 57 (48 to 59) 74 (72 to 74)
Age at 0 milk (days) 67 (57 to 72) 61 (51 to 63) 69 (52 to 74) 66 (55 to 68) 79 (78 to 79)
Duration of weaning (days) 11 (8 to 18) 5 (3 to 6) 18 (13 to 26) 7 (7 to 9) 5 (5 to 7)
HH = High-High (weaning started at 400 g/day ended at 1600 g/day); HL = High-Low (weaning started at 400 g/day and ended at 800 g/day; LH = Low-High (weaning
started at 200 g/day and ended at 1600 g/day); LL = Low-Low (weaning started at 200 g/day and ended at 800 g/day); CW = computer weaned.

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Miguel-Pacheco, Vaughan, de Passill and Rushen

Table 4 Medians and 25th and 75th percentile for digestible energy Table 5 Spearmans correlations (and probability P in parentheses)
intake (DE) per kilogram of BW and weight gain (WG) before weaning between the total duration of running, frequency of running bouts and
(PreW), immediately post weaning (PW1 to 3) and later post weaning mean bout duration of running bouts digestible energy intake (DE) per
(PW7 to 9) and age at start of weaning (Start), age at end of weaning kilogram of BW and weight gain (WG) before weaning (PreW), imme-
(End) and duration of weaning in days (Duration) diately post weaning (PW1 to 3) and later post weaning (PW7 to 9)
Variables Period 25th Median 75th Period Measure of running DE intake/kg BW WGs

DE (MJ/kg of BW) PreW 0.34 0.37 0.40 PreW Total duration 0.40 (<0.05) 0.47 (<0.05)
PW1 to 3 0.29 0.33 0.36 Bout frequency 0.42 (<0.05) 0.44 (<0.05)
PW7 to 9 0.31 0.36 0.43 Mean bout duration 0.13 (>0.10) 0.05 (>0.10)
WG (kg/days) PreW 0.83 1.10 1.45 PW1 to 3 Total duration 0.05 (>0.10) 0.04 (0.77)
PW1 to 3 0.70 1.00 1.45 Bout frequency 0.009 (0.95) 0.03 (0.81)
PW7 to 9 1.30 1.50 1.78 Mean bout duration 0.12 (>0.10) 0.07 (0.61)
Weaning age (days) Start 43 59 72 PW7 to 9 Total duration 0.25 (0.08) 0.34 (<0.05)
End 59 69 77 Bout frequency 0.24 (0.09) 0.30 (<0.05)
Duration (days) 5 8 13 Mean bout duration 0.07 (0.63) 0.17 (0.23)

Variability between calves weaning duration correlated with running duration, running
There were large differences between both individual calves frequency or mean bout duration (P > 0.10).
and during the three periods in the amount of running they Table 5 shows the correlations between the three
performed (Figure 2). The total duration of running in the period measures of running, DE intake and WG of the calves in each
immediately following weaning was signicantly lower than period. Before weaning (PreW), DE intake correlated with the
before weaning (W = 11379.63; P < 0.001) and increased in total duration of running and the frequency of running bouts,
the subsequent period (W = 11376.63; P < 0.001) (Figure 2). but not with mean bout duration. Similar ndings were
The daily frequency of running bouts followed a very similar observed between WGs and all three measures of running
pattern, being higher before weaning than immediately after (Table 5). Immediately after weaning (PW1 to 3), neither DE
weaning (W = 11373.75; P < 0.001), and increasing in the intake nor WG correlated with any variable of running. In the
next post-weaning period (W = 11362.88; P < 0.001) subsequent post-weaning period (PW7 to 9), we observed
(Figure 2). Changes in the mean bout duration followed a similar ndings to the before weaning period (PreW): DE
similar pattern: mean bout duration was lower immediately intake correlated with total duration of running and the fre-
after weaning compared with before (W = 11379.75; quency of running bouts, but not with mean bout duration.
P < 0.001) and increased slightly in the second post-weaning WGs positively correlated with both total duration of running
period (W = 11367.13; P < 0.001) (Figure 2). and frequency of running bouts, with a non-signicant trend
Table 4 shows the variation between calves in DE intake and for a correlation with mean bout duration (Table 5).
daily WG for each observation period, as well as the distribu-
tion of weaning ages and duration of weaning. DE intake
decreased signicantly immediately after weaning (PW1 to 3)
Discussion
compared with before weaning (PreW) (W = 11364.63;
P < 0.001), and the variability between calves appeared to Our results demonstrate that the amount of locomotor play
increase slightly (Table 4). DE intake increased during the showed before and after weaning reect both the energy
second post-weaning period (PW7 to 9) in comparison to the intake and growth rates in dairy calves. Locomotor play was
previous period (PW1 to 3) (W = 11367.00; P < 0.001). markedly reduced in the period immediately following
weaning, conrming previous ndings (Krachun et al., 2010).
Relationship between locomotor and other traits This decrease in total duration of running was associated
Before weaning, the age of the calves when the observations with a reduction in both frequency of bouts and mean bout
were taken was negatively correlated with the total duration duration. The reduction in frequency of bouts may reect that
of running (r = 0.36; P = 0.009) and running bout fre- the calf is less likely to start a running event. Additionally, the
quency (r = 0.38; P = 0.006); but no signicant correla- reduction of mean bout duration may indicate that the calf
tion was observed with mean bout duration (P = 0.88). The did not continue running or switch to an alternative beha-
birth weight of the calves was not correlated with the total viour. The decrease after weaning occurred despite the fact
duration of running (r = 0.11; P = 0.43) and the running that the weaning method prevented the large drop in energy
bout frequency (r = 0.03; P = 0.84); but it was positively intake that often occurs during weaning (de Passill and
correlated with mean bout running duration (r = 0.30; Rushen, 2012). This suggests that the decrease in locomotor
P > 0.04). During the immediately post-weaning period play following weaning is not solely due to a decrease in
(PW1 to 3), none of the variables of age related to weaning energy intake, as no correlation was observed immediately
(age when weaning began and age when it ended) and after weaning and may reect the emotional responses of

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Locomotor play of dairy calves around weaning

calves to weaning (Budzynska and Weary, 2008). Alter- weaning of calves according to intake of solid feed does
natively, it could be a specic response to the loss of milk, improve energy intake and WG during and after weaning but
because the improved energy intake after weaning can only does not prevent the marked reduction in locomotor play
result from an increase in the amount of solid feed ingested. that occurs immediately after weaning. This suggests that
We found a relationship between locomotor play and both reduction in energy intake is not the sole reason why loco-
energy intake and WG before and after weaning. Before motor play decreases at weaning. The occurrence of loco-
weaning, calves that ran more had better WGs and tended to motor play during and immediately after weaning off milk
have higher energy intake. Previously, Krachun et al. (2010) may be a behavioural indicator that a calf is adapting to
found a positive correlation between energy intake and loco- weaning and is able to maintain its energy intake and WG.
motor play after weaning, with variable correlations before These results support suggestions that measures of loco-
weaning, but the correlation with WG was not examined. Low motor play in young calves can be used to assess farm
levels of energy intake have been found to reduce play in other management practices associated with weaning, but more
animals such as deer fawn and foals (Mller-Schwarze et al., research is needed to understand the effect of other factors
1982; Pollard and Littlejohn, 2000; Nicol et al., 2005). Imme- on the amount of locomotor play that calves show.
diately after weaning we did not nd similar relationships. The
relationships between locomotor play, WG and energy intake
were also observed 7 to 9 days after weaning was completed. Acknowledgements
The previously unreported link between running and WG and The authors thank J. P. Parent, Ruby Wong, Jason and Gosia
energy intake provides further support for the suggestion that Zdanowicz. They also thank the staff at the Ethology Lab
locomotor play may be a sign of good welfare for growing (Agriculture and Agri Food Canada, Agassiz, BC, Canada) and
animals (Held and pinka, 2011), especially at a time when the dairy staff (University of British Columbia, Agassiz, BC) for
their growth potential is high. their invaluable help. Funding was provided by Agriculture and
These results add to the growing body of research that has Agri-Food Canada and the Natural Sciences and Engineering
observed a link between individual animals behaviour and both Research Council of Canada (both Ottawa, ON, Canada).
energy intake and growth rates (e.g. Stanton, 2012). Dairy
calves differ greatly in their growth responses to weaning: low
energy intake and WGs have been found to be associated with References
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