Response of dairy cows and calves to early separation:

Effect of calf age and visual and auditory

contact after separation

Ilona Stehulova a,*, Lena Lidfors b, Marek Spinka a

a Ethology Group, Research Institute of Animal Production, CZ-104 01 Prague-Uhrıneves, Czech Republicb Department of Animal Environment and Health, Section of Ethology, Swedish University of Agricultural Sciences,

P.O. Box 234, SE-532 23 Skara, Sweden

Accepted 2 March 2007

Available online 4 June 2007

Abstract

We evaluated how the behavioural and heart rate response of dairy cows and calves to mutual separation

was affected by two factors: the age of the calf at separation (1, 4 and 7 days) and the presence or absence of

visual and auditory contact between the mother and her calf after separation. In total, 46 cow–calf pairs were

randomly allocated to one of six treatments according to a 3 � 2 factorial design. The cows’ behavioural

response to separation was recorded at 0–1, 9–10, 25–26 and 50 and 51 h after separation. The calves’

behavioural response was recorded during the 24 h after separation. The heart rate was measured 60 min

before and 60 min after separation. Cows from the later separation treatments spent more time standing

( p < 0.01), placed their heads outside the pen ( p < 0.001), vocalised ( p < 0.05) and sniffed both the air

( p < 0.01) and other animals ( p < 0.001) more frequently compared to cows separated from their calves on

day 1. The behavioural response was also more intense when the cows could see and hear their calves: they

placed their heads outside the pen ( p < 0.001), vocalised ( p < 0.001) and sniffed the air ( p < 0.01) more

often than the no contact cows. Heart rate increased in all cows immediately after separation and was not

influenced by the calf’s age or by contact with the calf. After separation, older calves stood and moved more

( p < 0.001), placed their heads outside the pen more often ( p < 0.01) and showed more explorative

behaviour (i.e. sniffing walls and bedding, p < 0.01; licking walls, p < 0.05) than the calves separated on

day 1. Furthermore, the calves with visual and auditory contact with their mothers tended to place their

heads outside the pen more often ( p < 0.1) than the calves from the no contact treatments. Heart rate

increased rapidly in all calves after separation; however, the increase lasted longer in the later separated

calves ( p < 0.05). At 3 weeks of age, the calves separated on day 7 habituated more quickly to the novel

situation ( p < 0.001). Calves separated on day 4 were more likely to engage in social play than the calves

www.elsevier.com/locate/applanim

Applied Animal Behaviour Science 110 (2008) 144–165

* Corresponding author. Tel.: +420 267 009 590; fax: +420 267 710 779.

E-mail address: stehulova.ilona@vuzv.cz (I. Stehulova).

0168-1591/$ – see front matter # 2007 Published by Elsevier B.V.

doi:10.1016/j.applanim.2007.03.028

separated on days 1 and 7 ( p < 0.05) and the calves denied post-separation contact with their dams also

showed more social play compared to calves without this contact ( p < 0.05). The results indicate that the

response of cows and calves to separation is more intense and lasts longer when separation is delayed and

that this effect is prolonged and further intensified when they are allowed visual and auditory contact after

separation. On the other hand, delayed separation influences later social behaviour of calves in a way that

may enhance their coping abilities.

# 2007 Published by Elsevier B.V.

Keywords: Dairy cattle; Mother–young bond; Separation; Weaning; Behaviour; Heart rate

1. Introduction

On most commercial dairy farms in Europe and North America it is common to separate

calves from their mothers within the first 24 h after birth. With an increasing focus on animal

welfare in today’s society, alternative methods for separating dairy calves from their mothers are

being put into practice. For instance, the regulation in organic farming systems in Sweden is to

keep the calf with its mother during the colostrums feeding period, i.e. the first 4 days post-

partum (KRAV, 2005). However, some organic farmers believe that the separation after a few

days of contact between the cow and her calf causes more distress than separation after 24 h. It is

believed that later separation is, particularly distressing for the cow and may have a negative

impact on her subsequent milk let-down and behaviour in the milking parlour (Hartmann, 1994).

In cattle, the mother–young bond is established during the critical first few hours after birth.

Despite the suggestion by Hudson and Mullord (1977) that the mother–offspring bond can be

established after as little as 5 min of contact immediately after birth, the behavioural response

shown by both the mother and the young has been reported to be mild when separation occurred

within 24 h after birth (Weary and Chua, 2000; Flower and Weary, 2001, 2003). After the bond

has been established, separation will cause a more intense behavioural response. Both cows and

calves show increased vocalisations, activity levels and place their heads outside of the pen more

often (Lidfors, 1996; Weary and Chua, 2000; Flower and Weary, 2001). Furthermore, cows

separated from their calves later after birth call with a higher fundamental frequency (Weary and

Chua, 2000). Later separated cows have also been reported to ruminate less often immediately

after separation occurs (Lidfors, 1996). The fact that the behavioural reaction to separation gets

stronger with age is thus well established; however, other aspects and consequences of delayed

separation are not well documented.

First, separation may be performed in such a way that it prevents any further contact, for

example, by moving the calf to another building. Alternatively, physical separation that allows

for auditory or even visual contact is possible, for instance, by moving of animals to two different

pens which are close to each other. It is not clear which of these two possibilities makes

separation least stressful for the cow and the calf. For instance, the playback study by Marchant-

Forde et al. (2002) suggests that auditory contact increases the stress reactions. Contrary to this,

Price et al. (2003) showed that fenceline contact between cow and calf reduces signs of distress

from weaning in beef calves.

Second, the physiological indicators of stress, such as dynamics of heart rate during and after

separation, have received relatively little attention. If a cow can hear the vocalisations of her calf

after separation at 24 h, her heart rate increases, but the increase is only slightly higher in

response to her own calf compared to the increase in response to the calls of an alien calf

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165 145

(Marchant-Forde et al., 2002), perhaps because separation occurs before the cow is able to learn

her own calf’s calls. In a study by Hopster et al. (1995), cows separated at 2–3 days after

parturition showed only a short and mild increase in heart rate during the 10 min after separation.

To date no research has examined the change in heart rate dynamics in response to separation at

different ages and with/without partial contact between cow and calf after separation.

Third, little is known whether there are potential longer lasting benefits of keeping calves with

their mothers for a few extra days rather than separating them within 24 h after birth. Calves

separated at older ages (10–14 days), showed significantly higher weight gain during these first 2

weeks as well as increased weight gains during the post-separation feeding period (Metz, 1987;

Flower and Weary, 2001). Furthermore, those calves which are separated later tend to have a

lower frequency of diarrhoea (Weary and Chua, 2000). Additional time spent with the dam may

also result in longer lasting effects on the calves’ social behaviour, e.g. higher intensity of licking

and butting unfamiliar calves (Flower and Weary, 2001) which may indicate lower fearfulness of

unknown conspecifics (Krohn et al., 1999). It is not known whether a more subtle difference in

separation age, such as 1 day versus 4 days, would also result in longer lasting differences in

social behaviour.

The main aim of this study was to evaluate how the immediate post-separation behavioural

response and heart rate dynamics of dairy cows and calves were affected by two factors: the age

of the calf at separation (1, 4, 7 days) and presence or absence of visual and auditory contact

between the mother and her calf after separation. In addition, we examined how the reaction of a

calf to a new social situation at 3 weeks of age was affected by its early ontogenetic experience,

i.e. by the age at separation and post-separation contact.

2. Material and methods

2.1. Animals and housing

This study was carried out on a private organic dairy farm in southwestern Sweden, which owned

approximately 300 Swedish Red and White and Swedish Holstein dairy cows. The cows were kept in a

loose-housing barn with 308 lying cubicles bedded with sawdust. The cows were divided into 4 groups of

approximately 80 cows per group according to their stage of lactation. Cows were fed once daily with a total

mixed ration (TMR) consisting of silage, bean and wheat whole crop silage, farm mix (made of wheat, beans

and peas), concentrate and minerals. The cows also had free access to water and minerals. The cows were

milked twice a day in a milking parlour. All of the cows were dried off at least 6 weeks before calving.

Shortly before parturition until separation from their calves, the cows were kept in individual calving

pens (3.3 m � 3 m) with concrete floors bedded with sawdust and cleaned every morning. After calving the

cows had ad lib access to hay, grain mixture and TMR (the same composition as the TMR in dry period prior

to calving) and water. Cows were milked twice a day in the calving pen during the first day after parturition

and thereafter in the milking parlour with all of the other cows at approximately 6:00 and 15:00 h. During all

other hours of the day, the calves could freely suckle from their mothers. Farm workers helped some of the

calves find the dam’s teat or gave the calf the mother’s first colostrum from a teat bottle.

2.2. Experimental design and procedures

In the study, 46 multiparous dairy cows (19 Swedish Red and White cows and 27 Swedish Holstein cows)

between 3 and 9 years old (parity 2–7, median 3.2) were used. The cows were randomly allocated to one of

six treatments according to a block design. The treatment levels were age of calf at separation (days 1, 4, 7)

and presence or absence of visual and auditory contact after separation according to the following

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165146

distribution—(1) day 1: contact, n = 9; (2) day 1: no contact, n = 7; (3) day 4: contact, n = 7; (4) day 4: no

contact, n = 8; (5) day 7: contact, n = 7; (6) day 7: no contact, n = 8. The calves separated from the dam on

day 1 were between 20 and 28 h old, at 4 days of age they were 87–105 h old and at 7 days of age they were

159–177 h old at the time of separation. In all of the six treatments each breed and each sex of the calves

were represented by at least two animals. Cows with visual and auditory contact with their calf after

separation were housed in individual pens. Cows without contact with their calf after separation were

housed in a loose-housing barn (cubicles) with approximately 80 other cows.

The cows and calves were separated between 9:00 and 9:30 h, immediately after the morning milking.

At separation, the cows with post-separation contact with their calves were moved to another building

30 m away. It took about 2 min to move both the animals. In the new building, the cow was placed in an

individual sawdust-bedded pen (2.8 m � 6.7 m) positioned 8 m from the calf’s individual pen for

the following 51 h, so the cow and the calf could see each other (there was free space between the

two pens). The pen with the dam had two neighbouring pens: one calving pen with a highly pregnant

cow and a pen for three calves. The pen for the experimental calf is described below. After the 51 h of

contact the cows were moved into a loose-housing barn. The cows without any post-separation

contact were moved into the loose-housing barn immediately after separation. The distance between

calving pen and loose-housing barn was about 50 m and about 2–3 min were needed to move the cow.

After the cow was moved, the calf was relocated to an individual pen in another building. In the loose-

housing barn, the cows were kept in one large pen with approximately 80 other lactating cows of similar

stage of lactation. In all treatments, cows were given free access to TMR (the same composition as TMR

after calving).

After separation, all calves were placed into individual pens (1 m � 1.2 m) bedded with straw. The pens

had three solid-wood walls (each 1 m high) and a steel-bar door at the front of the pen, which was partly

open to allow the calf to put its head out for exploratory behaviour, drinking milk and water and eating the

farm mixture (grain, peas). The height of the walls also allowed the calf to place its head out of the pen in all

directions. Calves were fed 2 l of milk (or mother’s colostrum in the calves separated at 1 day of age) twice a

day from a teat bottle (the first 2–3 feedings after separation, depending on calf’s ability to learn suck

without assistance) and from teat buckets (until 3 weeks of age) at approximately 6:00 and 14:00 h. Calves

had free access to hay and farm mixture (grain, peas) in a bucket. During the entire rearing period (3 weeks),

each calf had physical contact with 1 calf kept in an adjoining pen and most of the time also had visual and

auditory contact with 1–2 other calves in other calf pens positioned 2 m away. At 3 weeks of age, each

experimental calf was moved into a group pen (4.3 m � 2.5 m) in which three non-experimental calves of

similar age and weight had been housed together for at least 3 days. The pen had a half-slatted, half-solid

floor bedded with sawdust. The calves were fed milk twice a day from open buckets and had free access to

hay in a rack and farm mixture.

The calves were weighed with an electronic scale (TRU-TEST Ltd., Ezi-weigh2, New Zealand) within

24 h after birth and again at 3 weeks of age, just before they were moved into the group pens.

2.3. Behaviour of the cows after separation

The behaviour of each cow was directly observed during four hourly periods between 0 and 1, 9 and 10,

25 and 26 and 50–51 h after separation. The second observation period (9–10 h after separation) started

about half an hour after the afternoon milking was finished. Three different recording methods were used:

(1) instantaneous sampling at 1 min intervals was used to record standing and moving/lying, (2) one-zero

sampling at 1 min intervals was used to record frequency of head out of a pen and (3) continuous sampling

was used to measure total frequency of vocalisation (1 incident of vocalisation = 1 call followed by intake of

breath), sniffing air, sniffing other animals and sniffing walls and bedding (1 incident = 1 occurrence of

behaviour followed by at least 1 s occurrence of a different behaviour) during the observation period. The

recorded behaviours are defined in Table 1. We also recorded the time spent feeding, drinking and

ruminating, but the preliminary analyses did not find any treatments differences for these measures. These

behaviours are not discussed further.

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165 147

2.4. Behaviour of the calves after separation

A black and white video camera (Elbex EP-700, Japan) was positioned 1.5 m above floor level in front of

each calf pen, together with a small spotlight to give enough visibility during the night. A 24 h video

recording of each calf was started immediately before the calf was moved into the individual pen. A time

lapse video recorder (Panasonic AG-6124, set on 24 h mode) with a monitor (Elbex EXM93) were used to

record the calf. The videotapes were analysed for standing and moving/lying using instantaneous sampling

at 1 min intervals. The occurrence of head out of the pen, sniffing walls and bedding and licking walls was

recorded with one-zero sampling at 1 min intervals. The descriptions of the behaviours are given in Table 1.

2.5. Heart rate

The heart rate of the cows and calves were measured by Polar X-trainer PlusTM (Polar Electro Oy,

Finland). The electrodes, transmitter and receiver were attached by a horse belt (Horse ware, Ireland) on

the left side of the body of the cows and calves (according to the user’s guide for the equipment) at least 1 h

before separation. We measured the heart rate in 5 s intervals during the 60 min before and the 60 min after

separation. Since the heart rate could have been influenced by the manipulation, only the rates recorded

during the 30 min before and the 60 min after separation were used for the analysis. The information from

the receiver was downloaded via a POLAR Advantage Interface to the Polar Precision Performance

Software, and then moved to Excel spreadsheets. We removed all the single value peaks if the difference

between the ‘‘peak’’ and both the preceding and following value was higher or lower than 40 bpm (beats

per minute) in cows and 50 bpm in calves. We also removed all of the zero values (indicating an interrupted

contact between the electrodes and body of the cow) and a flat line of values equal to or higher than

250 bpm, as these values indicated that the equipment was defective. When all of the corrections were

made, the mean values of all the valid measurements per minute were calculated for use in the statistical

analysis.

2.6. Behaviour of the calves at 3 weeks of age

Each experimental calf was weighed and moved to a group pen within 15 min before to the start of

observations. Each calf was then observed as a focal animal between 13:00 and 15:00 h (the time of highest

activity before the afternoon feeding) on two consecutive days (4 h in total). Using this direct observation

method, we recorded the total duration of standing and total frequency of head out of the pen, vocalisations,

social behaviour (sniffing animals, licking animals), social play (e.g. butting, running together), individual

play (e.g. running, jumping, head shaking) and explorative behaviour (sniffing walls and bedding, licking

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165148

Table 1

Behaviour of the cows and calves observed immediately after separation

Behaviour Description Recorded in

cows/calves

Standing and moving Animal is standing or walking or is in the process of

standing up or lying down

Cows/calves

Lying Animal is lying with no support of legs Cows/calves

Head out Tip of nose outside of any pen openings or above the railing Cows/calves

Sniffing walls and bedding Muzzle is within 5 cm of walls, steel bars or bedding Cows/calves

Vocalisation Every single call with inhalation between two occurences Cows

Sniffing animals Muzzle is within 5 cm of any body part of other animal Cows

Sniffing air Animal inhalates air with higher frequency than breathing,

head is directed upwards

Cows

Licking walls Tongue is in repeated contact with walls, steel bars, doors Calves

walls). Each occurrence of behaviour followed by at least 1 s occurrence of different behaviour was

recorded as one incidence. The definitions of these behaviours are given in Table 2.

2.7. Statistical analysis

The behavioural reaction of cows to separation was evaluated with a mixed linear model using the PROC

MIXED procedure in SAS (Statistical Analysis System for Windows V8, SAS Institute Inc., Cary, USA).

The fixed effects in the model were the day of separation (day 1, 4 or 7), post-separation visual and auditory

contact (yes/no), the hour of observation (the four hourly periods as a repeated measure, see Section 2.3), the

second-order interactions between the three above factors, and the sex and breed of the calf. We also

included the identity of the cow in the model as a random factor.

The behavioural reaction of the calves to separation was evaluated with a general linear model (PROC

GLM of SAS) was used. The effects were day of separation, post-separation contact, the interactions of the

former two factors, and the sex and breed of the calf.

The changes in heart rate after separation were quantified (for both the cows and calves) using the

following variables: peak heart rate, the latency of peak heart rate after separation, and the average heart rate

during the first, second, third and fourth 15 min periods after separation. These six variables were analysed

using the GLM procedure in SAS, testing for the effects of day of separation, post-separation contact, the

interactions of the former two factors, and the sex and breed of the calf.

The behaviour of calves after their introduction into a group at 3 weeks of age was analysed with a mixed

linear model (PROC MIXED in SAS). The fixed effects in the model were day of separation (day 1, 4 or 7),

post-separation visual and auditory contact (yes/no), the day of observation (the first or the second, see

Section 2.6), the second-order interactions between the above three factors, and the sex and breed of the calf.

We also included the identity of the calf as a random factor.

In all statistical analyses, the pair-wise differences between the 3 day of separation treatments were

tested using t-tests, with the Scheffe adjustment (in SAS) of the significance levels.

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165 149

Table 2

Behaviour of the calves observed at 3 weeks of age

Behaviour Description

Standing and moving Animal is standing or walking or is in the process of standing up or lying down

Lying Animal is lying with no support of legs

Head out Tip of the nose outside of any pen openings

Sniffing walls and bedding Muzzle is within 5 cm of walls, steel bars or bedding

Licking walls Tongue is in contact with walls, steel bars, doors

Sniffing animals Muzzle is within 5 cm of any body part of other animal

Licking animals Tongue is in repeated contact with any body part of other animal

Individual play Run (galloping movement that is not directed towards other calves)

Leap (leaving the ground without horizontal traveling)

Jump (travelling some horizontal distance while off the ground)

Rear (rocking backward into a bipedal position and then falling or jumping forward)

Head shake and jerk (shaking head from side to side or rotating head)

Kick (hind legs rapidly kicked up and back)

Fast turn (rapid direction change while running)

Fast stop (an abrupt halt while running)

Whirl (turning rapidly in a circle while remaining in place)

Social play Butt (head to head or head to body contact)

Kick (striking another animal with one or both hind limbs)

Running together (each movement is closely followed by a response of another calf)

Leaping together (each leap is closely followed by a response of another calf)

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Table 3

Behaviour of the cows during the four hourly periods (0–1, 9–10, 25–26, 50–51 h) after separation as affected by the age of the calves and post-separation contact

Treatment Contact No contact F-value and significance

Day 1 Day 4 Day 7 Day 1 Day 4 Day 7 Day Contact Day � contact

d.f. = 2 d.f. = 1 d.f. = 2

Standing and moving (%) 44.7 � 7.3 63.5 � 7.6 72.7 � 7.6 46.8 � 7.3 63.8 � 7.4 66.8 � 7.5 6.08** 0.04 ns 0.15 ns

Head outside the pen (no.) 7.5 � 2.5 22.9 � 2.7 23.3 � 2.6 4.0 � 2.5 5.6 � 2.6 6.4 � 2.6 8.37*** 35.17*** 4.84**

Vocalisation (no.) 39.0 � 15.5 89.1 � 16.3 96.4 � 16.1 8.9 � 15.5 11.1 � 15.9 33.3�15.9 3.69* 19.29*** 1.25 ns

Sniffing walls and bedding (no.) 4.0 � 1.1 4.5 � 1.1 7.9 � 1.1 3.4 � 1.1 3.2 � 1.1 3.2 � 1.1 1.88 ns 6.21* 1.86 ns

Sniffing animals (no.) 0.5 � 0.8 0.8 � 0.9 2.9 � 0.9 2.5 � 0.8 3.5 � 0.9 6.7 � 0.9 8.69*** 16.4*** 0.56 ns

Sniffing air (no.) 1.8 � 1.1 3.0 � 1.1 7.0 � 1.1 0.9 � 1.1 0.6 � 1.1 2.6 � 1.1 6.03** 8.04** 1.29 ns

Least square means (�S.E.) of number (no.) or percentage (%) of occurrence per 1 h, n = 46 (7–9 per treatment).* p < 0.05.

** p < 0.01*** p < 0.001.

3. Results

3.1. The behavioural response of the cows to separation

The age of the calf at separation had a strong influence on the response of the cow (Table 3 and

Fig. 1). Cows that were separated on days 4 and 7 after birth spent more time standing (day 4,

p < 0.05; day 7, p < 0.01) and placed their heads outside the pen more often (day 4, p < 0.001;

day 7, p < 0.01) than those separated on day 1. Further, the cows separated on day 7 also vocalised

more ( p < 0.01) compared to those separated on day 1. There was no significant difference in

these behaviours between cows separated on days 4 and 7. The cows that were separated on day 7

exhibited a higher frequency of sniffing other cows and/or calves than the cows separated on day 1

( p < 0.001) and day 4 ( p < 0.01). This same trend was found for sniffing air (day 1, p < 0.01; day

4, p < 0.01), however, there was no difference between those separated on days 1 and 4. The

frequency of sniffing walls and bedding was not affected by the day of separation.

Cows housed in individual pens near their calves after separation had visual and auditory

contact with the calves, and these animals showed a much stronger behavioural response to

separation compared to cows housed in a barn without any contact with their calves (Table 3 and

Fig. 2). They placed their head outside of the pen ( p < 0.001), vocalised ( p < 0.001), sniffed the

air ( p < 0.01) and sniffed the walls and bedding ( p < 0.05) more frequently than the cows who

had no contact with their calves. On the other hand, the sniffing of other animals was performed

more often by the cows without contact ( p < 0.001).

We found an interaction between contact and the age of the calf at separation for the head

outside of the pen behaviour (contact � day, F = 4.84, d.f. = 2, p < 0.01). In the contact

treatments, the cows that were separated on days 4 and 7 placed their heads outside the pen more

often compared to those separated on day 1 ( p < 0.01; Fig. 3). The differences between ages

were much lower in cows kept without any contact with their calves after separation (Fig. 3).

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165 151

Fig. 1. Least square means (�S.E.) of four behaviours (% of standing and moving, no. of head outside a pen, sniffing air,

vocalisations) observed in cows after separation from their calves at 1, 4 or 7 days of age of the calves in relation to the

time after separation (0–1, 9–10, 25–26, 50–51 h).

The Swedish Red and White cows seemed to respond more to separation, as they placed their

heads outside the pen more often than the Swedish Holstein cows (15.1 no./h versus 8.2 no./h,

F = 9.21, d.f. = 1, p < 0.01). The Swedish Red and White cows also had a tendency to sniff other

animals (3.5 no./h versus 2.1 no./h, F = 3.65, d.f. = 1, p = 0.06) and sniff the air (3.6 versus 1.8,

F = 3.51, d.f. = 1, p = 0.06) more often than the Swedish Holsteins. No other differences between

the breeds were found for the other behavioural parameters.

‘‘Standing and moving’’ was at its highest level immediately after separation and decreased

with time for all of the treatments (F = 21.96, d.f. = 3, p < 0.001). Placing the head outside of the

pen (F = 26.8, d.f. = 3, p < 0.001), vocalisations (F = 24.6, d.f. = 3, p < 0.001), sniffing the air

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165152

Fig. 2. Least square means (�S.E.) of four behaviours (% of standing and moving, no. of head outside a pen, sniffing air,

vocalisations) observed in cows with or without contact with their calves after separation in relation to the time after

separation (0–1, 9–10, 25–26, 50–51 h).

Fig. 3. Lest square means (�S.E.) of no. of head outside the pen observed in cows after separation from their calves at 1, 4

or 7 days of calf age. Values are presented separately for the cows with or without contact with their calves after separation

(significant differences: **p < 0.01).

(F = 15.14, d.f. = 3, p < 0.001; Figs. 1 and 2), sniffing other animals (F = 25.96, d.f. = 3,

p < 0.001) and sniffing the walls and bedding (F = 8.82, d.f. = 3, p < 0.001) were at the highest

levels immediately after separation, remained high for the 9–10 h observation period, and then

dropped substantially by the next day. The response of the cows separated later (e.g. days 4 and 7)

tended to last longer (Fig. 1), as they tended to increase their frequency of vocalisations between

0 and 1 and 9 and 10 h after separation (day � hour, F = 2.09, d.f. = 6, p < 0.06). There was also

a significant interaction between contact and the time after separation for most of the behaviours

(i.e. head outside the pen: F = 10.45, d.f. = 3, p < 0.001; vocalisations: F = 7.9, d.f. = 3,

p < 0.001; sniffing the air: F = 2.91, d.f. = 3, p < 0.05). The initial response of the cows with

contact was much higher and decreased with time after separation, but the increased activity

lasted for a longer period of time. The frequency of sniffing animals was higher in the no contact

treatments immediately after separation, but decreased in all of the cows between 9 and 10 h after

separation (sniffing animals: F = 15.7, d.f. = 3, p < 0.001; Fig. 2).

3.2. Heart rate of the cows

We were able to use only 41 of the 46 cows for the analysis of heart rate. In 5 cases, more than

75% of the recordings by the POLAR equipment were zero values or a flat line of values higher

than 250 bpm.

Before separation the cows that were separated on day 1 tended to have a higher baseline heart

rate than the cows separated on days 4 and 7 ( p < 0.1, F = 2.58, d.f. = 2). The heart rate of the

cows accelerated immediately after separation and remained higher during the 60 min following

separation compared to their heart rate before separation. However, no significant influence of

age of the calf at separation or visual and auditory contact on heart rate was found (Fig. 4).

3.3. The behavioural response of the calves to separation

The age of the calves clearly influenced their response to separation (Table 4 and Fig. 5). The

calves that were separated on days 4 and 7 spent more time standing and moving (day 4,

p < 0.001; day 7, p < 0.001), they placed their heads outside the pen more often (day 4,

p < 0.05; day 7, p < 0.01) and sniffed the walls and bedding more frequently (day 4, p < 0.01;

day 7, p < 0.05) than those calves that were separated on day 1. There were no differences

between days 4 and 7 for these variables (Fig. 5). A similar trend was found for the frequency of

licking walls, which occurred significantly more often in the calves separated on day 4 compared

to day 1 ( p < 0.05), but there was no difference between the other ages (Fig. 5).

The calves from contact treatments tended to place their heads outside of the pen more often

( p < 0.1). There was no influence of contact with the mother on any other behavioural

parameters (Table 4 and Fig. 6).

Further, we did not find any interactions between the age of the calf at separation and contact

with the dam (Table 4).

The peak of activity for both standing and placing the head outside of the pen was observed

immediately after separation and between 11 and 14 and 20 and 22 h (Figs. 5 and 6). The

frequency of sniffing walls was highest immediately after separation and between 11 and 12 h

after separation. The frequency of licking walls was highest between 9 and 10, 13 and 15 and 20

and 22 h after separation (Figs. 5 and 6).

The bull calves stood and moved more after separation compared to heifer calves (22.3%

versus 16.9%, F = 4.9, d.f. = 1, p < 0.05). Despite this finding, there were no differences

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165 153

between the sexes for any of the other behaviours. Breed did not influence calf behaviour after

separation in any way.

3.4. Heart rate of the calves

Due to technical problems, as previously described, we were only able to use the heart rate

recordings for 39 out of 46 calves for baseline calculations, 38 calves for the interval between 0

and 14 min, 37 calves for the interval between 15 and 29 and 30 and 44 min and 35 for the

interval between 45 and 60 min after separation.

The baseline heart rate was not related to the age of the calves at separation (Fig. 7). Peak and

average heart rate recorded between 0 and 14 min after separation were also similar for

separation at all ages. Differences in heart rate occurred between 15 and 29 min (F = 3.99,

d.f. = 2, p < 0.05) and 30 and 44 min (F = 3.87, d.f. = 2, p < 0.05) after separation; the calves

that were separated on day 7 had a significantly higher heart rate for a longer period of time than

calves that were separated on day 1. Heart rate decreased in all calves between 45 and 60 min

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165154

Fig. 4. Mean heart rate (beats per minute) in cows during the 30 min before and the 60 min after separation (time 0 = time

of separation) when separated from their calves at 1, 4 or 7 days of age of the calves. Values are presented separately based

on the presence (A) or absence (B) of visual and auditory contact with their mothers after separation.

I.S

tehu

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15

5

Table 4

Behaviour of the calves during the 24 h after separation as affected by the age of the calves at separation and post-separation contact

Treatment Contact No contact F-value and significance

Day 1 Day 4 Day 7 Day 1 Day 4 Day 7 Day Contact Day � contact

d.f. = 2 d.f. = 1 d.f. = 2

Standing and moving (%) 12 � 2.8 21.2 � 3.0 26.3 � 2.9 11.3 � 2.8 22.4 � 2.9 24.6 � 2.9 13.47*** 0.03 ns 0.12 ns

Head outside the pen (no.) 2.2 � 1.5 6.2 � 1.5 8.1 � 1.5 1.0 � 1.5 4.3 � 1.5 5.0 � 1.5 6.46** 2.91 ns 0.22 ns

Sniffing walls and bedding (no.) 2.92 � 0.8 6.0 � 1.8 5.3 � 0.8 3.1 � 0.8 4.8 � 0.8 4.2 � 0.8 5.24** 1.26 ns 0.46 ns

Licking walls (no.) 0.2 � 0.2 0.9 � 0.2 0.6 � 0.2 0.2 � 0.3 1.1 � 0.3 0.4 � 0.3 3.46* 0.00 ns 1.08 ns

Number (no.) or percentage (%) of occurrence per 1 h (mean � S.E.), n = 46 (7–9 per treatment).* p < 0.05.

** p < 0.01.*** p < 0.001.

after separation and there was no effect of age on this variable. Further, the heart rate was not

influenced by contact with the dam after separation (Fig. 7).

3.5. Behaviour of the calves at 3 weeks of age—longer lasting effect of early separation

Due to technical reasons, we were only able to observe 37 out of 46 calves at 3 weeks of age

and in each of the treatments 1–2 calves were missing. The behaviour of the calves was

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165156

Fig. 6. Means (�S.E.) of four behaviours (% of standing and moving, no. of head outside a pen, sniffing walls and

bedding, licking walls) observed in calves with or without contact with their mothers after separation in relation to the

time after separation.

Fig. 5. Means (�S.E.) of four behaviours (% of standing and moving, no. of head outside a pen, sniffing walls and

bedding, licking walls) observed in calves after separation at 1, 4 or 7 days of age in relation to the time after separation.

influenced by the age of the calf at separation (Table 5). We found that the age of the calf

influenced the amount of social play, as the calves that were separated on day 4 exhibited a higher

frequency of social play compared to those separated on day 1 ( p < 0.05) and day 7 ( p < 0.05).

A similar trend was found for vocalisations ( p < 0.1), with significant differences between

calves separated on days 1 and 4 ( p < 0.05) and days 4 and 7 ( p < 0.05). The trend was reversed

for the frequency of licking other animals. The earliest separated calves tended to lick the other

calves more often than the calves that were separated on day 7 ( p < 0.1).

The age of the calves at separation also influenced their ability to habituate to a novel

environment, as there were significant interactions between separation age and observation day

for standing and moving (F = 8.61, d.f. = 2, p = 0.001; Fig. 8). The calves that were separated on

day 7 decreased their activity from the first to the second day of observation, as they spent less

time standing and moving. In contrast, the calves that were separated on day 1 stood and moved

more on the second day while those separated on day 4 did not change their activity between the

observation days (Fig. 8). A similar, yet not significant, tendency was found for the frequency of

licking other animals (F = 2.54, d.f. = 2, p < 0.1; Fig. 8) and vocalisations (F = 2.8, d.f. = 2,

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165 157

Fig. 7. Mean heart rate (beats per minute) in calves during the 30 min before and the 60 min after separation (time

0 = time of separation) when separated at 1, 4 or 7 days of age. Values are presented separately based on the presence (A)

or absence (B) of visual and auditory contact with their mothers after separation.

I.S

tehu

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l./Ap

plied

An

ima

lB

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viour

Scien

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10

(20

08

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65

15

8

Table 5

Behaviour of the calves recorded during the two observation days at 3 weeks of age after moving to a group of unfamiliar calves as affected by the age at separation and

post-separation contact

Treatment Contact No contact F-value/significance

Day 1 Day 4 Day 7 Day 1 Day 4 Day 7 Day Contact

d.f. = 2 d.f. = 1

Standing and moving (%) 68.4 � 5.4 73.8 � 5.9 75.2 � 5.1 68.9 � 6 85.5 � 5.2 66.6 � 5.2 2.23 ns 0.07 ns

Head outside the pen (no.) 5.4 � 1.8 3.3 � 1.9 5.4 � 1.6 4.5 � 2.0 8.3 � 1.7 3.0 � 1.7 0.41 ns 0.16 ns

Vocalisation (no.) 10.8 � 12.2 24.5 � 13.3 13.4 � 11.5 12.4 � 13.6 51.0 � 11.8 6.9 � 11.8 3.11 ns 0.5 ns

Sniffing walls and bedding (no.) 18.5 � 4.6 18.3 � 5.0 17.3 � 4.3 18.8 � 5.1 31.5 � 4.4 20.4 � 4.4 1.16 ns 2.09 ns

Licking walls (no.) 2.4 � 0.8 2.6 � 0.8 2.0 � 0.7 1.3 � 0.8 1.4 � 0.7 1.3 � 0.7 0.09 ns 2.54 ns

Sniffing animals (no.) 12.2 � 1.9 11.6 � 2.0 9.6 � 1.8 11.4 � 1.81 14.7 � 1.8 114.7 � 1.8 1.01 ns 0.52 ns

Licking animals (no.) 3.5 � 0.7 3.2 � 0.8 1.9 � 0.7 2.9 � 0.8 2.0 � 0.7 1.1 � 0.7 2.96 ns 2.96 ns

Individual play (no.) 14.9 � 6.2 14.2 � 6.8 11.4 � 6.0 4.8 � 6.9 18.4 � 6.0 20.2 � 6.0 0.64 ns 0.03 ns

Social play (no.) 1. � 1.4 3.7 � 1.6 1.0 � 1.4 2.6 � 1.6 8.5 � 1.4 1.0 � 1.4 4.18* 6.94*

Least square means (�S.E.) of number (no.) or percentage (%) of occurrence per hour (first day, n = 37; second day, n = 36).* p < 0.05.

p < 0.08; Fig. 8). For these variables, the increases in activity between days were apparent for

calves separated from the dam on both days 1 and 4 after birth.

The calves that were kept without any contact with their mothers after separation showed

more social play at 3 weeks of age than those that did have contact with their dams ( p < 0.05).

No other behavioural differences associated with contact with the dam after separation were

found (Table 5).

The behaviour of calves also changed with time after they were moved into a social group at 3

weeks of age (Fig. 9). On the first day of observation (immediately after moving into the group),

the calves showed more explorative behaviour than on the second day (after 24 h), i.e. placed the

heads outside the pen (F = 7.73, d.f. = 1, p < 0.01), sniffed walls and bedding (F = 31.4, d.f. = 1,

p < 0.001) and sniffed other calves (F = 6.44, d.f. = 1, p < 0.05) more often. On the first

observation day, the calves also exhibited a higher frequency of individual play compared to that

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165 159

Fig. 8. Least square means (�S.E.) of the number of recordings. Effect of calf age at separation on the behaviour (% of

standing and moving, vocalisation and licking animals) of the calves during the first and the second day when they were

placed in a group pen at 3 weeks of age.

on the second day (F = 13.60, d.f. = 1, p < 0.001). Conversely, this trend was reversed for the

frequency of licking other animals and licking walls (licking animals: F = 6.09, d.f. = 1,

p < 0.05; licking walls: F = 5.47, d.f. = 1, p < 0.05). The frequency of social play and

vocalisation did not differ between the observation days (Fig. 9).

3.6. Weight gain of the calves

The weight of the bull calves at birth and at 3 weeks of age was significantly higher than the

weight of the heifer calves (birth weight: 48 � 1.8 kg versus 41 � 1.7 kg, p = 0.01, n = 37; 3

weeks: 55 � 1.5 kg versus 49 � 1.4 kg, p < 0.01, n = 35), but weight gain did not differ between

sexes. Further, there was no influence of age at separation or contact with the dam after separation

on weight gain during the first 3 weeks.

4. Discussion

4.1. The behavioural response of the cows to separation

The cows that were separated later showed much stronger and longer lasting response to

separation than those that were separated on the first day after birth. These results agree with the

findings of Lidfors (1996; immediately after birth versus 4 days), Weary and Chua (2000; 1 day

versus 4 days) and Flower and Weary (2001; 1 day versus 14 days). Our results indicate that cows

are less stressed by an earlier separation from the calf. It seems improbable that the increase in the

behavioural reaction was due to a pure calf age effect as Weary and Chua (2000), Flower and

Weary (2001) found that the activity of cows is low when they are kept with their calves and they

do not show any significant differences in activity and vocalisation in relation to calf age. Our

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165160

Fig. 9. Least square means (�S.E.) of the number of recordings. Behaviour of all of the calves during the first and the

second day when they were placed in a group pen at 3 weeks of age (significant differences: *p < 0.05, **p < 0.01,

***p < 0.001).

inability to detect any differences in the majority of the post-separation stress indicators between

the days 4 and 7 separation treatments suggests that the mother–young bond is fully established

before the fourth day after birth. The only exception were those behaviours associated with the

olfactory system (i.e. sniffing air and sniffing other animals), which were most frequent in the

cows separated at 7 days. This seems to indicate that these cows were attempting to use their

olfactory senses to locate their calves. However, it is unclear whether or not these behaviours are

signs of distress.

The behavioural response of the cows was more pronounced and lasted longer when

they could see and hear their calves, particularly for those separated after a longer period of

time. This corresponds well with the study of Marchant-Forde et al. (2002), who found

that auditory contact between mother and young after separation increases the magnitude of

their reaction. However, the cows’ behavioural reactions seemed to wane relatively

quickly after separation. Vocalising, placing head out of the pen and sniffing air dropped

to low levels by 25 h (and for some treatments even by 10 h) after separation. Thus, even

though we found clear evidence for higher stress levels after later separation and after

separation followed by visual and auditory contact with the calf, these differences may be

limited to the first day after separation and therefore may not represent a serious compromise

of the well-being of the cow.

The differences in cows’ response associated with contact with the calf in our study may have

been partly due to differences in housing systems after separation. The cows that had contact with

their calves were housed in individual pens and it may be argued that this relative social isolation

in small pen caused the higher frequency of placing the head outside and more frequent

vocalisation. However, since the cows rarely vocalise and rarely place their heads outside the pen

before separation (Weary and Chua, 2000; Flower and Weary, 2001) and since our cows were

habituated to individual housing before separation, it does not seem that the individual housing

per se was causing the more active and vocal behaviour. On the other hand, the much higher

sniffing of other animals by the cows in the no contact treatments was probably due to the fact that

they had a more opportunity for this behaviour in the loose-housing barn.

The Swedish Red and White cows tended to have a more marked response to separation

compared to the Swedish Holstein cows. Similarly, in a previous study by Loberg and Lidfors

(2001), the Swedish Red and White cows had a stronger reaction to the presence of alien foster

calves.

Similarly to findings of Weary and Chua (2000), the behavioural response of the cows was at

its highest level immediately after the calf was removed and remained high for 9–10 h after

separation. Unlike Flower and Weary (2001), who found a second peak of behavioural response

18 h after separation, we detected a rapid decrease which lasted until the following day. This may

be explained by differences in methodology used in the two studies. In the present study, we

observed cows during the resting period after the morning milking and after feeding whereas

Flower and Weary (2001) recorded the behaviour of the cows at 6:00 h, just prior to the morning

milking.

4.2. Heart rate of the cows

The cows reacted to separation with an obvious increase in heart rate. This finding agrees with

Hopster et al. (1995), who demonstrated a heart rate increased in cows separated from their calves

at 3 days of age and Marchant-Forde et al. (2002), who found that cows had higher heart rates in

response to a playback of calf calls after separation.

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165 161

Although part of the initial heart rate increase could be attributed to the physical exercise

associated with moving the cow, it is improbable that walking 30 or 50 m would alone cause such

a strong and long lasting increase in heart rate. Therefore, we attribute the change in heart rate to

the emotional stress of separation.

Our study is the first to investigate the effect of age of the calves at separation on the heart rate

response of dairy cows. In contrast to the clear differences in behavioural reactions, we found no

effect of the calves age at separation or visual and auditory contact with the calf on the heart rate

of the cows. This was probably because the heart rate was only measured for 1 h after separation

whereas the behavioural measurements were carried out until 51 h after separation. We observed

a dramatic response in heart rate in all of the cows during the first hour after separation which

corresponds with the peak in the behavioural response. However, the strongest differences in

behavioural response between treatments did not occur until the later three observation periods,

when heart rate was no longer recorded.

4.3. The behavioural response of the calves to separation

Similarly to the work of Lidfors (1996), Weary and Chua (2000) and Flower and Weary

(2001), we found that calves separated at older ages were generally more active after separation.

The frequency of putting the head outside the pen was also higher in those calves separated on

days 4 and 7 compared to those separated on day 1, thus supporting the previous findings that

after a few days of contact, the reaction to separation is much more apparent. It is unlikely that the

increases in calf behaviour after separation were a result of natural changes with age in activity

and calling frequency, because Weary and Chua (2000) and Flower and Weary (2001) found that

calves are generally inactive when they are kept with their mothers and they do not show any age

differences in activity and frequency of vocalisation.

Even though the calves that had contact with their mothers were exposed to frequent

vocalisations from their mothers, we did not find any effect of post-separation contact on the

behaviour of the calves. This may seem surprising since Marchant-Forde et al. (2002) reported that

calves are able to recognise the calls of their mother when separated at 24 h of age and show

behavioural response to the calls when they are played back to them. However, the behavioural

response in that study consisted of head and ear movements, and the calves did not react by

standing up or walking during the test. In our study, we only recorded behaviours that involved

movement of the whole body. Thus, if we only consider these behaviours, our results are in

agreement with Marchant-Forde et al. (2002). The activity of the calves was highest immediately

after moving them away from their mothers, yet they were relatively inactive during the following

few hours. A similar behavioural pattern was noted by Lidfors (1996), Weary and Chua (2000) and

Flower and Weary (2001) in the first few hours after calves had been removed from their mothers. It

is possible that calves are adapted to short periods of separation during the first days of life, since in

the wild they would be hidden for such periods when they are young (Webster, 1994). The second

peak of activity occurred approximately 12 h (at 21:00 h) after separation. At this time, there had

been at least a 7 h period since the previous feeding and the calves may have been hungry, because

calves suckling from their dams usually feed 6–10 times per 24 h (e.g. Lidfors, 1996; Albright and

Arave, 1997). A similar response to milk deprivation was described by Flower and Weary (2001)

who observed a peak in activity 18 h after separation. Thomas et al. (2001) found that if calves were

fed five times a day after separation they vocalised approximately six times less compared to those

that were fed only two times a day. Therefore, it seems that the calves’ response to separation could

be minimised by providing a sufficient number of milk feedings.

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165162

4.4. Heart rate of the calves

Calves on all treatments exhibited an increase in heart rate after separation, probably

indicating an immediate stress reaction to separation. The initial rise (both peak and average

bpm first 15 min after separation) was similar in all treatments. Despite this, heart rate was

significantly higher from 15 until 45 min after separation in the calves that were separated on

days 4 and 7. Hence, the change in heart rate lasted longer in the calves that were separated at

older ages. The difference in heart rate may be partly attributed to the fact that the older

calves were generally more active, e.g. standing and moving more often, after separation

compared to calves separated at day 1. We did not detect any effect of post-separation contact

with mother on the heart rate of the calves, even though the cows frequently vocalised.

Contrary to our finding, Marchant-Forde et al. (2002) reported an increase in heart rate in

calves during the playback of the calls of their mothers after separation occurred at 24 h of age.

In contrast, Steinhardt and Thielscher (1999) reported lower heart rates in calves that could

hear the voice of their mothers during an isolation test at 15 days of age. Thus, evidence for the

reaction of the circulatory system of the calves to the calls of their mothers remains

controversial.

4.5. Behaviour of the calves at 3 weeks of age—long term effects of early separation

The calves were observed immediately after they had first contact with unfamiliar calves. The

testing procedure, i.e. putting the focal calf into an already established small group of other

calves, was used to mimic social challenges that the calf would likely encounter later life.

Unfortunately, little is known about how this type of situation affects the calf’s behaviour, and

this lack of information complicates our interpretation of these results, except the quick

habituation of the initial agitation and frequent play which could bee seen as signs of good coping

abilities. The calves that were separated on day 7 seemed to cope better in a novel situation since

they demonstrated high activity immediately after they were placed into the group of unfamiliar

calves and spent more time resting during the second day of observations. These results closely

resemble those of Krohn et al. (1999), where calves separated at 4 days of age spent more time

within a 2 m distance of unfamiliar calves during a test at 3 and 11 weeks of age than calves that

were separated within 24 h. Flower and Weary (2001) also found that calves that were kept with

their mothers during the first 2 weeks after birth exhibited more intense social behaviour (licking,

butting, rubbing of the head) towards unfamiliar calves, which could indicate lower fearfulness of

unknown conspecifics (Krohn et al., 1999). This conclusion is only valid for the calves which are

reared individually during the milk feeding period, because additional social contact during the

rearing period could substantially influence the calves’ social behaviour. There may also be

longer lasting benefits of keeping cows and calves together for longer periods of time. A study by

Le Neindre and Sourd (1984) demonstrated that heifer calves kept with their mothers were more

socially dominant in adulthood compared to those that were separated immediately from their

mother and fed from a bucket. Two results of our study also suggest that the social environment

during the first week of life affects the calf’s social behaviour over time. First, calves that were

separated on day 4 engaged in more social play than calves that were separated on day 1 or 7.

Second, social play occurred more often in calves that were kept without any post-separation

contact. However we do not currently understand why these changes are inconsistent over time

(e.g. different only on day 4, but not day 7) nor do we understand the mechanism behind these

differences in social play.

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165 163

In sum, it seems clear that the social behaviour of calves is influenced by the age at separation

and by post-separation contact with their mother. Further research is needed to establish in more

detail the specific effects of both the age of separation and post-separation contact on the social

behaviour of the calves.

4.6. Weight gain of the calves

In the present study, the weight gain of calves during the first 3 weeks of age was not

influenced by the age at separation or contact with their mothers after separation. These

results agree with Weary and Chua (2000), who did not find any weight gain advantages from

delayed separation at 4 days of age. In a study by Krohn et al. (1999), calves separated

at the age of 4 days showed twice the rate of gain until separation, even if they were not

allowed to feed from the dam, than those separated immediately after birth, but the effects

on weight gain were only transitory as they were restricted to the period of differential

treatment. In contrast, other studies found that delaying separation to 10 or 14 days resulted in

a 2–3 times increase in weight gain and that these effects persisted up to 4 weeks of age

(Flower and Weary, 2001) or 2 months (Metz, 1987) after separation. It seems likely that

differences in weight gain associated with age of separation are dependent on the amount of

time spent with mother, and these differences may explain some of the discrepancies between

studies.

5. Conclusions

Marked behavioural reactions indicate that separating dairy cows from their calves is more

stressful for both parties if it is accomplished on days 4 and 7 than on day 1. The behavioural

response to separation was more intense and lasted longer in both of the animals when they had

visual and auditory contact after separation. The heart rate response of cows was not influenced

by the age of the calves at separation or post-separation contact. However the heart rate increase

in the later separated calves lasted longer. Further, the age of the calves at separation and contact

with the mother influenced the behaviour of the calves during social challenges at 3 weeks of age.

From the welfare point of view, there seems to be a trade-off between the more stressful option of

separating calves from their mothers several days or weeks after birth and the potential longer

lasting benefits of prolonged contact between the cow and calf, particularly for the calves. Further

research should focus on the magnitude and duration of the longer lasting benefits of additional

cow–calf contact.

Acknowledgements

The authors thank the owners of the organic farm Kartorp for allowing us to carry out this

experiment in their herd of dairy cows. Further, we thank the farm animal manager Olof Andren

and the other dairy farm staff who helped us very much with all animal manipulations and the

observations. Funding was provided by FOOD21 (MISTRA and FORMAS) in Sweden. This

research project has been also co-financed by the European Commission, within the 6th

Framework Programme, contract no. FOOD-CT-2004-506508. The text represents the authors’

views and does not necessarily represent a position of the Commission who will not be liable for

the use made of such information. Trevor DeVries and Cassandra Tucker kindly helped with the

English of the final version of the text.

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165164

References

Albright, L.L., Arave, C.W., 1997. The Behaviour of Cattle. CAB International, Wallingford, UK.

Flower, F.C., Weary, D.M., 2001. Effects of early separation on the dairy cow and calf. 2. Separation at 1 day and 2 weeks

after birth. Appl. Anim. Behav. Sci. 70, 275–284.

Flower, F.C., Weary, D.M., 2003. The effects of early separation on the dairy cow and calf. Anim. Welfare 12, 339–348.

Hartmann, C., 1994. Att lata mjolkkornas kalvar dia. Specialarbete 17. Swedish University of Agricultural Sciences,

Skara (in Swedish).

Hopster, H., O’Connell, J.M., Blokhuis, H.J., 1995. Acute effects of cow–calf separation on heart-rate, plasma-cortisol

and behaviour in multiparous dairy-cows. Appl. Anim. Behav. Sci. 44, 1–8.

Hudson, S.J., Mullord, M.M., 1977. Investigations of maternal bonding in dairy cattle. Appl. Anim. Ethol. 3, 271–276.

KRAV Standards, 1999. KRAV, Uppsala, Sweden, p. 50.

Krohn, C.C., Foldager, J., Mogensen, L., 1999. Long-term effect of colostrum feeding methods on behaviour in female

dairy calves. Acta Agric. Scand. Sect. A: Anim. Sci. 49, 57–64.

Le Neindre, P., Sourd, C., 1984. Influence of rearing conditions on subsequent social-behaviour of Friesian and Salers

heifers from birth to 6 months of age. Appl. Anim. Behav. Sci. 12, 43–52.

Lidfors, L.M., 1996. Behavioural effects of separating the dairy calf immediately or 4 days post-partum. Appl. Anim.

Behav. Sci. 49, 269–283.

Loberg, J., Lidfors, L., 2001. Effect of stage of lactation and breed on dairy cows’ acceptance of foster calves. Appl. Anim.

Behav. Sci. 74, 97–108.

Marchant-Forde, J.N., Marchant-Forde, R.M., Weary, D.M., 2002. Responses of dairy cows and calves to each other’s

vocalisations after early separation. Appl. Anim. Behav. Sci. 78, 19–28.

Metz, J., 1987. Productivity aspects of keeping dairy-cow and calf together in the postpartum period. Livest. Prod. Sci. 16,

385–394.

Price, E.O., Harris, J.E., Borgwardt, R.E., Sween, M.L., Connor, J.M., 2003. Fenceline contact of beef calves with their

dams at weaning reduces the negative effects of separation on behaviour and growth rate. J. Anim. Sci. 81, 116–121.

Steinhardt, M., Thielscher, H.H., 1999. Reaktionsmuster von Tieren auf gewohnte und ungewohnte Ereignisse. Transport

and temporare Separation von Saugkalbern aus der Mutterkuhhaltung in verschiedenen Altersperioden wahrend der

fruhen Aufzuchtperiode sowie Effekte der Prasentation von Muttertierrufen auf hormonelle variablen, Herzfrequenz

und Lautaußerungen der Tiere. Landbauforschung Volkenrode Heft. 49, 153–166.

Thomas, T.J., Weary, D.M., Appleby, M.C., 2001. Newborn and 5-week-old calves vocalise in response to milk

deprivation. Appl. Anim. Behav. Sci. 74, 165–173.

Weary, D.M., Chua, B., 2000. Effects of early separation on the dairy cow and calf. 1. Separation at 6 h, 1 day and 4 days

after birth. Appl. Anim. Behav. Sci. 69, 177–188.

Webster, J., 1994. Animal Welfare: A Cool Eye Towards Eden. Blackwell Science, Oxford, UK.

I. Stehulova et al. / Applied Animal Behaviour Science 110 (2008) 144–165 165