The motivation-based calving facility: Social and cognitive factors influence isolation seeking behaviour of Holstein dairy cows at calving
The motivation-based calving facility: Social and cognitive factors influence isolation seeking behaviour of Holstein dairy cows at calving
Maria Vilain Rørvang 0 1 2
Mette S. Herskin 0 1 2
Margit Bak Jensen 0 1 2
0 Aarhus University, Department of Animal Science , Tjele , Denmark
1 Demonstration Programme of the Danish Ministry of Food , Agriculture & Fisheries, Copenhagen, Denmark , and Graduate School of Science & Technology, Aarhus University , Denmark, for funding to MVR. The funders had no role in the
2 Editor: Juan J. Loor, University of Illinois , UNITED STATES
In order to improve animal welfare it is recommended that dairy farmers move calving cows from the herd to individual pens when calving is imminent. However, the practicality of moving cows has proven a challenge and may lead to disturbance of the cows rather than easing the process of calving. One solution may be to allow the cow to seek isolation prior to calving. This study examined whether pre-parturient dairy cows will isolate in an individual calving pen placed in a group calving setting and whether a closing gate in this individual calving pen will cause more cows to isolate prior to calving. Danish Holstein cows (n = 66) were housed in groups of six in a group pen with access to six individual calving pens connected to the group area. Cows were trained to use one of two isolation opportunities i.e. individual calving pens with functional closing gates (n = 35) allowing only one cow access at a time, or individual calving pens with permanently open gates allowing free cow traffic between group area and individual pen (n = 31). The response variables were calving site, calving behaviour and social behaviour. Unexpectedly, a functional gate did not facilitate isolation seeking, perhaps because the cows were not able to combine a learnt response with the motivation to isolate. Dominant cows had the highest chance of calving in an individual calving pen. If an alien calf was present in the group pen or any of the individual pens, cows were less likely to calve in an individual calving pen. Future studies should allow cows easy access to an individual calving pen and explore what motivates pre-parturient cows to seek isolation in order to facilitate voluntary use of individual calving pens.
Calving is an essential part of commercial milk production and during the transition period
(typically defined as three weeks before and three weeks after calving), there are several threats
to dairy cow welfare. Calving itself places high demands physically and is associated with pain
]. Disease and mortality rates are high during this period, with nearly 75% of disease cases
occurring within the first month after calving (reviewed in [
]) and 30±50% of post-partum
cows being affected [
]. When cows calve in group calving pens they are frequently disturbed
design of this study, data collection/analysis or
preparation of the manuscript.
by other cows and the risk of mismothering is high [
]. In order to protect the calving cow, it
is currently recommended that cows calve in individual calving pens to which they are moved
well in time before calving (e.g. by law in Denmark by Ministry of Environment and Food of
Denmark, Danish Veterinary and Food Administration, Law number. 520, Chapter 4, 26/05/
] and by recommendation of The Canadian Dairy Code of Practice [
]). Cows are,
however, often moved too late due to the difficulty of determining the onset of the first stages
of labour, which have been suggested to be an appropriate time to move cows to individual
calving pens [
]. Moving cows later may disturb the process of calving and prolong the
second stage of labour , leading to an increased risk of calving complications (dystocia) and
related diseases [
]. The current international trend towards increased herd size [
that farmers have more cows to supervise, making it more difficult to identify the correct time
to move cows to individual calving pens. Increased herd size may also result in farmers using
group calving facilities more often, which may not be consistent with the choice of the cows.
In order to aid identification of imminent calving, devices have been developed to monitor
behavioural and physiological changes before calving, such as reduced rumination [
increased number of lying bouts [
]. In addition, vaginal temperature has been reported to
decrease prior to calving  and sensors to detect this physiological change are now available.
However, these changes are only measurable in the hours prior to calving meaning that many
cows are unlikely to be moved in time. Hence, there is a need for a practical solution
facilitating the moving of cows well in time before calving. One possible solution may be to develop a
motivation-based calving facility, taking advantage of the natural motivation of calving cows
to seek isolation.
To design motivation-based calving facilities, knowledge of the behaviour and preferences
of calving cows are essential. The natural behaviour of cattle is to stay within the proximity of
the herd, and typically cattle synchronize their behaviour [
]. However, as parturition
approaches, cows become restless [
] and move away from the group and isolation
seeking behaviour have been reported among cows kept in semi-natural  and production
]. The underlying evolutionary strategy may be to ensure that the female
bonds to her offspring without disturbance from group members, but isolation seeking
behaviour may also have developed to protect the newborn against predation as suggested by .
In a practical setting, motivation for isolation seeking may be utilized when cows are to be
moved into individual calving pens before calving, by allowing cows to move into these pens
on their own, and at the same time ensure a timely move to individual housing, as well as
ensuring an undisturbed environment.
One aspect of isolation seeking behaviour is the social factors inevitably arising from
housing pregnant cows in groups. In addition, individual differences between cows are apparent.
Although sparsely studied, dairy cows have different levels of sociability [
] and individual
behavioural characteristics [
]. Making a choice of where to calve, may also depend on the
personality of the cow. To our knowledge, neither the effects of social factors, nor personality,
has been studied previously. Edwards [
] noted that when calving in groups, cows close to
parturition were exceedingly attracted to calves (as 14 out of 16 cows licked an alien calf before
giving birth themselves). Calves are, however, not the only distraction. In intensive indoor
dairy production, pre-parturient cows are often moved to a new group, where the level of
aggression can be relatively high [
], especially in relation to accessing limited resources, such
as an individual calving pen. Adding to this, maternal aggression (expression of defensive and
aggressive behaviour to protect the offspring [
]) is well documented in a number of
ungulates (cattle: [
] sheep: [
], pigs: [
]), which may lead to even higher levels of aggression
among peri-parturient females. Within a group of pre-parturient cows, social dominance
determines the outcome of competition for resources [
]. If individual calving pens are
2 / 19
perceived as a limited resource by the cows, the chance of gaining access to a pen is likely
higher for dominant individuals. However, possible interactions between social dominance
and stage of pregnancy cannot be ruled out. The motivation to seek isolation may also cause
cows to avoid social confrontations and become more submissive as calving approaches. What
happens in relation to personality and dominance in the pre-parturient cows and whether
other factors affect the motivation to isolate before calving, must be determined in order to
develop motivation-based calving facilities.
The aim of this study was to examine whether cows kept under conventional dairy farming
conditions will isolate in an individual calving pen when they are trained to access this
beforehand. We housed 13 groups of six pregnant dairy cows in group pens with access to individual
calving pens, allowing either free cow traffic (the gate was permanently open) or access for
only one cow at a time (functional mechanical gate). We hypothesized that cows housed with
functional gates would be more likely to seek isolation in an individual calving pen prior to
calving due to their experience of being alone, behind the closed gate compared to cows
housed in groups where calving pens had free cow traffic and thus no gate to keep other cows
out. Additionally, we investigated whether other factors, such as personality, dominance,
group structure, and presence of alien calves influenced the choice of calving site during 12 to
8 h pre-calving. A personality assessment tests was incorporated into the experimental plan by
using Human approach tests and recording social interactions. We hypothesized that
subordinate cows would be more likely to seek isolation behind the gates due to increased risk of
aggression from dominant cows and that the presence of an alien calf would lower the
probability of cows moving into an individual calving pen.
Materials and method
The experiment complied with the current Danish law, except for calving in individual calving
pens and except for the cow-calf pairs being separated before 12 hours after calving. The
procedures were evaluated prior to the experimental start by the responsible laboratory animal
veterinarian from the institutional animal ethics committee at The Department of Animal
Science, Aarhus University, Tjele, Denmark. It was confirmed that no ethical oversight was
The experiment took place at the Danish Cattle Research facility at Aarhus University
(Foulum, Denmark) September 2015 to June 2016.
The experimental barn had 3 sections (Fig 1), each consisting of a group area (9 x 9 m)
connected to 6 adjacent individual calving pens (4.5 x 3 m each). The individual calving pens were
built from 1.3 m high tubular metal bars covered by 1.8 m high grey plastic barriers (barrier
width: 10mm, Jyden, Vemb, Denmark), on the two 4.5 m sides as well as 1.5 m of one other
side (half of the pen side facing the group area), leaving a 1.5 m opening towards the group
area. The fourth side (facing the outer walls of the barn) had a 3.0 m feed bunk (Jyden, Vemb,
Denmark). This construction was chosen to offer the cows an opportunity to isolate (Fig 2A
and 2D). In the opening into the group area (1.8 x 1.5 m), a mechanical cow gate was installed.
The gate was designed to allow access for just one cow at a time by locking mechanically after
one cow had entered the pen i.e. opened the gate and passed through (Prototype designed by
Jyden, Vemb, Denmark). The cow was always able to go back to the group area, after which,
the gate would unlock mechanically from the manipulation when the cow pushed the gate
open, allowing another individual to enter. In half of the sections, the experimental calving
pens were equipped with open gates (gate permanently open), while the other half of the
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Fig 1. The experimental barn. Top view of the experimental barn, including all three sections. Thick lines around
individual calving pens represent covered sides, and the ªwindowº illustrates where the mechanical gate (grey insertion
in the ªwindowº) was installed in all individual calving pens. Vertically attached brushes, drinking cups and feed tables
sections had a functional gate (i.e. gate) (as illustrated in Fig 2A and 2D). The two experimental
treatments (permanently open gate; functional gate) were applied to one whole section in a
balanced order to control for possible effects of section placement within the barn. The order
Fig 2. The levels of closing the gate. A: The view from an individual calving pen with open gate, representing the start
point for all cows (initial training, step 1) and the treatment termed ªpermanently open gateº. B: The top bar of the gate
being closed (corresponding to training step 2 for cows housed with ªfunctional gatesº). C: The trainer holding the
gate, half way open, in order for the cow to see the way out (corresponding to training step 3 for cows housed with
functional gates). D: The view from inside the gated individual calving pens with the gate fully closed (corresponding
to training step 4 for cows housed with ªfunctional gatesº).
PLOS ONE | https://doi.org/10.1371/journal.pone.0191128
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was (section; treatment: 1; permanently open, 2; functional, 3; functional, 1; functional, 2;
permanently open, 3; permanently open, 1; functional, 2; permanently open, 3; functional, 1;
permanently open, 2; functional, 3; permanently open, 1; functional).
The floor of all group areas and individual calving pens was covered by approximately 15
cm of sand (Kosand, brand; Dansand, Brñdstrup, Denmark; mean grain size 0.322 mm) and
approximately 15 cm of barley straw on top as bedding to ensure good lying comfort and a
non-slip surface for getting up and lying down behaviour of the cows. New straw was added
on a daily basis in the morning between 9.30 and 12.00 am after cleaning (removal of faeces
and wet and soiled straw). Cleaning was done to maintain similar bedding quality in the
different areas. After each calving, the place where the amniotic sac broke, and the place where the
calf was born, was located using video surveillance (described below) and straw as well as sand
was fully removed in an area of Ø: 1 m. Afterwards new sand and straw were added. This
procedure was used in order to limit the potential influence from leftover birth fluids [
Each group area had 6 individual feed bins constituting the feed table (bin width: 75 cm,
model 1318±8210, Jyden, Vemb, Denmark) and two self-filling, automatic drinking cups
(model 2177±4010, Jyden, Vemb, Denmark). Each individual calving pen had one water cup
similar to the ones in the group area.
Cows were only fed in the group area to avoid confounding of feed and isolation motivation
in the individual calving pens. During the whole experimental period, a total mixed ration
with a forage-to concentrate ratio of 80:20 (%, dry matter basis) was provided for ad libitum
intake. Feed was allocated twice daily, in the morning between 9.30 and 12.00 am, and in the
afternoon between 5.30 and 6.00 pm.
Initially, the experiment included a total of 78 multiparous Danish Holstein cows all provided
from the Danish Cattle Research facility at Aarhus University (Foulum, Denmark). This
sample size was chosen based on a pre hoc power analysis, utilizing the probability of calving in a
pen of 0.71 (found in a pilot experiment). Based on the power analysis, we decided to include 6
animals per section meaning that there was one individual pen per cow. The probability of
detecting a difference in the experiment was 94% (at least 80% is recommended [
including 72 animals. The total sample size thus consisted of 72 animals determined from the
power analysis plus an additional 6 animals included as a buffer. The buffer of animals was
included due to the possibility of having to exclude cows due to sickness, calving difficulty or
other calving related issues (see exclusion criteria below).
Thirteen section groups were constructed based on expected calving dates in order to
ensure a similar dispersion of calvings and allow barn staff time to clean between calvings.
Section group cows had at least 1 day between expected calvings and the average dispersion
between expected calvings was 6.9 days (range: 1 to 15 days).
Training and tests
Each group of six cows was moved to the experimental section and placed in the group area
approximately two weeks prior to the first expected calving. All cows were allowed 24 h to
acclimatize to the experimental housing before initiation of training (Fig 3). During this
period, all gates were kept permanently open regardless of treatment, and cows could move
freely in and out of the individual calving pens. Depending on the gate treatment, a specific
training procedure was used (see below).
Personality assessment. On the day of training initiation, a personality assessment of all
cows was done based on the cowsÂ immediate reaction in terms of exploration-avoidance of
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Fig 3. Time line. Days prior to experimental start, indicating all initial procedures and assessments. Moved to the barn (day -14), temperament test (day -12) after a
settling period (day -14 to -12), initial training (day -12 to -3), entering and re-entering tests (day -3 and -2) leading to either inclusion or exclusion (day -1) and
experimental start (day 0).
the trainer and the training procedures. This was done in a ªsection entering testº and a
ªhuman approach testº, conducted prior to the initiation of training as well as during the first
steps of the training procedures (described below) and termed ªfearfulness during the initial
trainingº. The ªsection entering testº assessed the cow's immediate reaction to the trainer.
When the trainer entered the barn for the first time, the trainer would stand in front of the
group area for a few minutes (2 to 5 min) before entering the section. The trainer then climbed
the outer walls and entered the middle of the group area. Inside the group area, the trainer
walked to the centre, making sure that all cows paid attention, and then scored the cows
according to the definitions shown in Table 1. The ªhuman approach testº was initiated by
identifying a cow in the group area not eating, drinking or engaging in social interactions, and
which was attentive towards the trainer. The trainer would then approach the focal cow from a
distance of two cow lengths (approximately 5 m) from an angle of 45Ê of the front of the cow,
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with an outstretched arm and an even pace (as described by ). When the cow moved away
from the trainer, the avoidance distance was noted. If the cow did not move, the avoidance
distance was left censored as zero (Table 1). The evaluation of ªfearfulness towards the initial
trainingº was carried out during the first training bout and consisted of evaluating the reaction
of the cow towards the initial training procedure, and the level reached according to the initial
training steps described below. Finally, each cow was evaluated as being ªexplorativeº or
ªavoidingº according to the definitions in Table 1. A cow was determined `shy’ when scored in
the ªavoidingº category for at least two out of three assessments, and likewise determined
`bold’ when scored in the ªexplorativeº category for at least two out of three assessments.
Boldness was thus defined as an explorative animal, i.e. willing to explore a novel person (the
trainer) and a novel situation (the initiation of training) [
Initial training. Cows were trained section-wise i.e., trained individually, but with the
other five cows of the section present during training. Each training day had two distinct
periods of training chosen based on expected level of feeding motivation and disturbances: 08 to
10 am just before feeding, and 01 to 03 pm just before topping up the feed.
Each training lasted a maximum of 8 minutes per cow (determined during a pilot training
session with pilot cows prior to the experiment) and the average training duration was
4.8 ± 2.6 min per bout. Training bouts were separated by either one day or 3 to 7 hours. A
training bout was always initiated from within the group area, and each cow was trained to
enter a randomly chosen individual calving pen (i.e. decided beforehand by a third person
rolling a dice). If the chosen individual calving pen was already occupied, the neighbouring pen to
the right was chosen. Throughout the successive training bouts, cows were always trained in
the pen to the right, during the proceeding training, in order to allow experience with all
individual calving pens. No cow was chased out of an individual calving pen in order to be trained,
i.e. if a cow was inside an individual calving pen, the trainer would wait until the cow
reentered the group area before initiating her training.
All cows were gradually trained to follow the trainer by reinforcing the approach by positive
reinforcement i.e. eating concentrates from a black plastic bucket (12 litres). The trainer would
stand in front of the cow and at first allow the cow to eat a mouthful from the bucket without
having to move (step 0). If the cow was unwilling, or too fearful to eat from the bucket, she was
allowed to eat from the bucket while it was standing on the bedding without the trainer present
(watching from a 3 m distance). In such cases, the trainer would gradually approach the cow
while she was eating from the bucket, until the trainer was able to touch and later lift the
bucket. Shortly after completing step 0, the trainer moved a few steps away from the cow
towards the individual calving pen while still facing the cow and displaying the bucket. When
the cow followed, she was again allowed to eat a mouthful upon reaching the bucket and the
trainer. At first, the reward would be given after just a few steps in order for the cow to learn to
follow. The demand would then rise as training progressed. Step 1 consisted of the trainer
moving backwards through the opening of the individual calving pen, spending a few seconds
inside the individual calving pen rewarding the cow and then leaving. When the cow walked
willingly through the open gate three times, she had completed step 1.
Cows housed with functional gates (n = 35): The gate was gradually closed in three steps:
First the top bar was closed (step 2), secondly, the gate was closed, but the trainer held the gate
halfway open in order for the cow to see the way in and out (step 3) and lastly, the gate was
fully closed (step 4) (Fig 2B, 2C and 2D, respectively). In these steps, the trainer passed through
the gate opening, and then faced the cow to entice her to pass. Each time the degree of closing
the gate was changed, the procedure of going in and out was repeated until the cow walked
willingly through, three times. During the training, the trainer ensured that no other cows
followed the focal cow into an individual calving pen at any point. The success criterion for the
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initial training was that the cow should be able to open the closed gate on her own three times
(i.e. two times going into the individual calving pen, and one time going out) without showing
any fear-related behaviour (Table 2). For visual of cows being trained to perform step 4, see S1
Cows housed with permanently open gates (n = 31): All cows were gradually trained to
walk with the trainer through the gate opening of the individual calving pen and back again, as
described above (step 1). The duration of a training bout was determined as the mean duration
of a corresponding training bout for cows trained to the functional gates. Likewise, the number
of training bouts corresponded to the number of training bouts for cows trained to functional
gates, and thus all cows, irrespective of the treatment, were trained for the same duration of
time. In this treatment, other cows were allowed to follow the trained cow when she entered
an individual calving pen, and likewise allowed to enter the individual calving pen with the
focal cow and the trainer. The success criterion for this training was that the cow should be
able to enter the individual calving pen (two times) and re-enter the group area (one time)
without showing any fear-related behaviour (Table 2).
Entering- and re-entering tests. When a cow complied with either success criterion, she
was left inside the individual calving pen to which she was trained as the trainer left through
the gate. The cow then had a maximum of 4 hours to re-enter the group area on her own. This
test was called the re-entering test.
When the re-entering test was passed, each cow was observed for 48 hours (on video) to
check if she entered any individual calving pen(s) on her own. This period constituted the
When initial training, the re-entering as well as the entering test was passed successfully, a
cow was considered ready for the experiment and thus complied with the overall success
criterion (Fig 3).
Training progress and exclusion during training. The training progress was supervised
for all 78 cows to make sure all cows complied with the success criterion and to ensure that all
were trained for approximately the same amount of time. Seventy-four cows reached the
criterion after 4 successive days of training. However, four cows still showed strong fear reactions
towards the trainer and the feed bucket upon their fourth training day (eighth training
session). These were given two extra training days. Thus, median of training duration was 4
successive days (range 4 to 6 days) until compliance (Fig 4). One cow did not reach the success
criterion (housed with functional gates) within these extra days, and was excluded from the
experiment. The remaining three cows managed to reach the criterion and were included.
Two cows calved during the training period (i.e. before complying with the criterion) and
were excluded. In total, 75 cows; 41 housed with functional gates and 34 housed with
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Fig 4. Graphs of mean training level per training bout for each treatment illustrating the progress during the
initial training period (divided onto treatment: ªFunctional gatesº and ªpermanently open gatesº). Training level:
Step 0 (not able to follow the trainer), step 1 (following the trainer in and out of individual calving pen = success
criterion for ªpermanently open gatesº) and step 4 (following the trainer in and out of individual calving pen while
opening the gate without any help or encouragement = success criterion for ªfunctional gatesº). Training bouts 1±9 (2
bouts per day) leading to experimental start (shown in light grey) and subsequent weekly post training checks 1.1 to 1.6
(1 bout per day) during the experiment (shown in dark grey).
permanently open gates, were included after the training sessions, and they all complied with
the success criterion in the subsequent weekly post-training checks (Fig 4).
Measures and analysis
Data editing and video recordings. For each cow the experimental period ended when
she calved and each cow-calf pair was removed 5 to12 h after calving, in order to allow the cow
time to nurse the calf and for the staff to have time to move them. This relatively rapid removal
limited the period where potentially attractive alien calves were present [
]. Cows were
excluded from the study if they had a stillborn calf (n = 2), gave birth to twins (n = 1), or had
clinical milk fewer, mastitis or retained placenta (diagnosed by the herd veterinarian on the
day of calving (n = 4)). Delivery of the calf was assisted if the calf was not born within 4 h after
the appearance of the amniotic sac. Assisted calvings also resulted in the cow being excluded
from the study (n = 2). Thus, 66 cows could be included in the analysis (35 housed with
functional gates (from 7 groups having 5, 5, 5, 5, 4, 5, and 6 cows per group) and 31 housed with
permanently open gates (from 6 groups having 4, 6, 5, 5, 6, and 5 cows per group); and 19 and
8 cows entering their second parity, 10 and 16 cows entering their third parity, and 6 and 7
cows entering fourth or later parities, respectively, for the two treatments). The behaviour of
the cows was monitored by digital video cameras (model: TVCCD-624, Monacor, Bremen,
Germany) mounted above the sections: Two cameras covered each end of each group area and
one camera was mounted above each individual calving pen, leading to a total of 8 cameras per
section. Recordings were made continuously throughout the whole experimental period, and
an experienced technician monitored and stored all video material. The behaviour of each cow
was observed continuously for 12 h prior to calving by an experienced observer after
termination of the data collection. The same observer performed all video analyses.
Use of the individual calving pens and calving. Location of the cow and the occurrence
of behavioural elements (Table 3) were monitored in order to keep track of the calving process
and the use of the individual calving pens. The response variable was the location of the birth
place (the placing of the cow when the hips of the calf were fully expelled from the birth canal),
which also determined the end-point of the observation period. The location of the cow was
determined as either within the group area or within an individual calving pen. If a cow was
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located between the two areas, the head of the cow determined the outcome. This definition
was also used when determining where the cow experienced her first sequence of rhythmical
abdominal contractions. The duration of the second stage of labour was defined as initiated at
the first rhythmical abdominal contraction bout (Table 1) and continuing until the hips of the
calf were fully expelled ([
Social behaviour. Social behaviour includes displacements with physical contact and
displacements without contact (described in Table 3). For each focal cow all displacements given
and received were summarized over the 12 h video observation period. Furthermore, it was
noted if an alien calf was present within the last 8 h before calving as this is the approximate
period where cows have been reported to isolate [
]. Within each group, the order of
calving was noted. For each calf born, a pregnant heifer (average age and body weight: 21.6 months
and 575 ± 48 kg) was added to the group pen after the removal of the cow and her calf.
Gait scores and body weight. Cows were gait-scored when entering and exiting the
experiment. The scoring was done by two experienced observers according to the 5-point
scoring system developed by Thomsen et al. [
]. Two cows were scored as obviously lame (above
score 3) when entering the experiment, but treated for laminitis (hoof washing followed by
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application of a bandage with salicylic acid treatment) upon entering, and scored 2 and 1,
respectively, before calving. Median gait score was 1, when entering as well as exiting the
experiment (range 1 to 4 at entry, and range 1 to 3 when exiting). Cows were weighted upon
entering the experiment and when leaving the experiment 5 to12 hours after calving, by the
use of the same automatic scale (Danvñgt, Hinnerup, Denmark). The cows weighed on
average (mean ± s.d) 687 ± 75 kg and 715 ± 61 kg when entering the experiment, and 674 ± 56 kg
and 692 ± 65 kg when leaving the experiment, for cows housed with functional gates and for
cows housed with permanently open gates, respectively. After calving, cows were machine
milked once in the experimental pen before being moved from the barn. Rectal temperature
was measured at this milking (mean ± s.d.: 38.7 ± 0.8ÊC).
Maintenance of bedding and gates
Bedding quality of each individual calving pen and group area was evaluated daily at 08:30±
10:00 am prior to removal of manure. The quality of the bedding was scored by the barn staff
according to a 5-point scale [
]: (0) dry, no faeces or urea, (1) moist, some parts with faeces
(n < 3), (2) slightly wet, both dark and light straw and more than 3 parts with faeces, (3) wet,
mostly dark straw and faeces, (4) very wet, only dark straw and faeces and urea spread over the
whole pen (median: 1, range: 0±4).
The force needed to open a functional gate from inside the individual calving pens and
from the group area was measured by an electronic scale (model OCSF300, Scale House 1,
3711 NW 36th St., Miami, FL 33142) at the end of the experiment (5 measures per gate),
showing that 12.5 ± 1.8 kg and 12.2 ± 3.6 kg force was needed, respectively.
Behavioural variables. We calculated the social dominance as a so-called `rank ratio' for
each cow based on the displacements summarized over the 12 h before calving: the number of
individuals being displaced by the focal cow, and the number of individuals displacing the
focal cow; the higher the ratio the higher the rank [
]. Afterwards, the ratio was converted
into an index (x 100) indicating if the cow was in the dominant or subordinate end of the scale
from 1±100 within the group. Calving order was identified as a categorical variable with 6
levels. The presence of an alien calf (a calf born from another cow which was not yet removed
before the next calving) during the last 8 hours before calving was categorized as one or zero
(present or absent). Similarly, the location where the cow experienced her first sequence of
rhythmical abdominal contractions was categorized as `Group area' or `Individual Pen'. A
preliminary analysis showed positive correlations (based on Mann-Whitney U test; [
personality score (shy, bold) and dominance and thus only one of these two variables could be
included in the model. Social dominance was chosen over personality assessment (`shy' or
`bold') as this measurement was based on observations of the cows close to time of calving.
Based on Shapiro-Wilk normality test and visual assessment of histograms, normality of the
recorded variables could not be assumed. Hence, all variables were analysed using models and
tests not assuming normality. Statistical analyses were performed using the R software, version
3.1.2 (R Core Team 2014, Vienna, Austria) and all p-values evaluated according to a
significance level of 5% and 10% for tendencies.
Modelling. Due to the primary response variable having a binary outcome (i.e. calving
site), data were analysed using logistic regression and package lme4 [
] in the R software. It
was not possible to include season or other time-related measures in the analyses, as the
section groups were temporally overlapping to ensure suitable inter-calving intervals within
group. We did, however, include the placement of each section group within the barn as a
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random effect in a mixed effects model. Thus, the full model included the random effect of
placement within barn (i.e. section number 1, 2 or 3 in Fig 1) as well as the fixed effects;
dominance ratio within group at calving (mean ± s.d.: 64 ± 33), duration of second stage labour
(mean ± s.d. (min): 108 ± 51), calving order within group (1, 2, 3, 4, 5 or 6), parity (second,
third or older), presence of alien calf (0 or 1) and the location where the cow showed her first
rhythmical abdominal contractions (individual pen, or group area). The full model did not
show any effect of the random variable ªsectionº (variance and s.d. = 0), which was
subsequently excluded. A normal logistic regression model was then fitted to test effects of the
explanatory variables. The normal logistic regression model was reduced using a backwards
stepwise procedure with p < 0.1 as inclusion criteria. The final model included the fixed
effects; rank ratio within group at calving, presence of alien calf and the location where the
cow showed her first rhythmical abdominal contractions. The model fit was checked using the
Hosmer Lemeshow Goodness of fit test [
]. Odds ratios and 95% confidence intervals were
constructed from the final parameter estimates by back-transforming, applying the inverse
Calving location. In order to examine whether cows calved in random locations within
each section, and thus were not affected by the location of previous calvings (as suggested by
]), a 1-sample proportion test with continuity correction was used [
All cows displaced other cows but also received displacements from other cows, and thus the
social dominance of each cow within each group could be determined through rank ratio
calculation without any cows sharing the same level of dominance within a group. Bold cows
(n = 34) had significantly higher rank ratios whereas shy cows (n = 32) had lower rank ratios
(Mann Whitney U-test: (median; range (rank ratio index)) bold: 93; 6±100 vs. shy: 41; 0±100,
W = 806, r = 0.8, p < 0.001).
Factors affecting isolation seeking behaviour at calving
Effect of being housed with functional gates. Across the two treatments 34 cows calved
in an individual calving pen and 32 in the group area. The odds of calving in an individual
calving pen tended to increase when cows were housed in sections with permanently open
gates compared to cows housed in sections with functional gates (Table 4).
`Treatment' i.e. having a functional gate or a permanently open gate, `Rank' i.e. the rank ratio index within each group at calving, `Presence of alien calf' i.e. having an
alien calf present within the last 8 h before calving or not and `where 1st sequence of rhythmical abdominal contractions occurred' i.e. in the group area or in an
individual calving pen. Odds ratio for each variable with corresponding 95% confidence intervals and p-values are presented along with coefficients `b' and stand errors
12 / 19
Effect of dominance. Rank ratio significantly affected choice of calving site; the higher
the rank within a group at calving, the higher the odds of calving inside an individual calving
pen. For every increase in rank order of 1 the odds of calving inside the individual pen
increased by 1.14 (Table 4), and due to the assumption of proportional odds in the model this
was the same for each level of rank e.g. rank 33 to 34 and rank 66 to 67.
Effect of location of first rhythmical abdominal contractions. Another factor,
influencing the choice of calving site, was the location of the first sequence of rhythmical abdominal
contractions. Fifty-one percent of the cows had the first sequence of rhythmical abdominal
contractions while being in an individual calving pen and these cows had significantly higher
odds of calving inside the individual calving pen than cows showing their first sequence of
rhythmical abdominal contractions within the group area (Table 4). Additionally, 10 cows (4
housed with functional gates and 6 housed with permanently open gates) had their first
rhythmical abdominal contraction in the group area and then moved to an individual calving pen
during the second stage of labour, whereas 6 cows (3 cows housed with functional gates and 3
cows housed with permanently open gates) moved from an individual pen to the group area
during the second stage of labour.
Effect of presence of an alien calf. An alien calf was present within the last 8 hours prior
to another calving for 41% of the cows. For these cows the odds of calving in an individual
calving pen decreased by 0.46 (Table 4).
Three variables had no effect in the full model and were thus stepwise removed from the
model: First parity (pparity2 = 0.85 and pparity3 = 0.62), then duration of second stage labour
(mean ± s.d.: 108.5 ± 51.4 min) (p = 0.65) and lastly calving order within group (1 to 6) (p2 =
0.89, p3 = 0.16, p4 = 0.49, p5 = 0.38 and p6 = 0.41). These had no effect on the odds of calving
inside an individual calving pen (p > 0.1) and were thus not included in the final model.
The median dispersion between calvings within group was 6 (range: 0 to 20) days. For 8 out of
13 groups, one or two cows (n = 10) calved in close proximity to the location of the last calving
(within one cow length from the site of last calving). When testing these groups separately in
the 1-sample proportion test, however, no evidence for selection of calving site due to the site
of a previous calving could be found, neither for groups with one cow calving close to another
or groups having two cows calving close to a third cow.
In this study, we gave cows an opportunity to move to another pen to calve and examined
whether isolation seeking behaviour was facilitated by this opportunity to isolate (seek
seclusion as well as increase the distance to group members). Additionally, the study included social
aspects of housing pre-parturient cows in groups and the effects of these on choice of calving
site. We found that 50% of the cows moved away from the group and isolated in an individual
calving pen regardless of the presence of a functional gate in the pen or not. Unexpectedly,
cows housed with permanently open gates tended to be more likely to calve in the individual
calving pens than cows with functional gates. Dominance increased the odds of calving in an
individual calving pen whereas the presence of an alien calf, during the last 8 hours prior to
calving, decreased the odds.
Contrary to the hypothesis, cows housed with permanently open gates tended to be more
likely to calve in the individual calving pens compared to cows housed with functional gates.
This implies that the cows experienced the gates as obstacles rather than an advantage for
isolating before calving. As we have previously found that cows seeking maximum isolation (75%
13 / 19
versus 50%) at calving had longer second stage labour [
], the fact that there was no effect of
the duration of second stage labour on the choice of isolation in this study further supports
that the gate may have been an obstacle for the cows. All cows received prior training and
complied with a learning criterion for being able to use the gate before being included in the
experiment, suggesting that the cows had learnt to use the gate. In none of the treatments did the
choice deviate from random. In this study, the cows had to combine a learnt and rather
conspicuous response (opening the gate and knowing that no one could follow) with the
motivation to isolate. Research on the cognitive abilities of cattle is generally limited, but cows do
possess the ability to perform rather complex instrumental conditioning tasks [
], have quite
good spatial memory and are able to navigate quite complicated mazes which they memorize
for up to 6 weeks [
]. Furthermore, all cows used the pens on their own several times before
calving, and approx. 50% ended up calving in one of the pens. Therefore, there is no particular
reason to argue that the cows did not learn and memorise the task of opening the gate.
Memorising a conspicuous task compared to memorising e.g. a route to a preferred grass patch may
however, be of less biological relevance (as argued for horses by [
]). Furthermore, being in
labour and experiencing pain may have made it difficult for the cows to recall this knowledge.
Not because the task of isolating was not biologically relevant, but because the task of opening
a rather conspicuous gate was less biologically relevant in this situation. The gate may also
have been perceived as too heavy to manoeuvre, or uncomfortable to pass, especially once the
cow was in labour. Being in labour, the abdomen of a pregnant cow may become increasingly
sore as contractions arise [
] and thus opening a gate which place pressure on the abdomen
may be unpleasant. Furthermore, being in labour pain, cows may have tried to avoid potential
conflicts with conspecifics and thus they may have favoured to be able to control their visual
field and have the opportunity to withdraw from conspecifics. Therefore, entering a gated pen
may have been too risky also because it required effort to leave. In terms of the evolutionary
history of ungulates, the trade-off between isolation seeking and having less control in term of
executing a flight response to avoid a predator also supports this. Lastly, the distance between
the group area and the individual calving pen may have influenced whether cows used the
individual pens for calving or not. In a semi-natural setting, Lidfors et al. [
] noted that cows
walked a considerable distance away from the group and it is thus possible, that in the present
study, the cows perceived the individual calving pens as situated too close (maximum 9 m
away) to the rest of the group in order for the pens to be perceived as offering isolation.
Future studies on motivation-based calving facilities may examine whether gates that are
less conspicuous and require less effort to open result in more cows calving in them.
Potentially, a gate that closes behind the cow when entering the calving pen, without a need for
pushing or manipulation may be more appropriate. Also combining a simpler and less heavy gate
with a pen that offers a higher level of isolation could be one option to facilitate the use of
individual calving pens. In addition, potential effects of distance between individual calving pens
and the group-members should be examined. Lastly, more research on learning and
memorising of more complex instrumental tasks in cattle is needed in order to understand the capacity
of cattle to learn how to perform a task and recall it in the peri-parturient period, where other
underlying motivations may be strong.
In the present study social order within the group influenced whether cows chose to calve
in the individual calving pens (Table 4). When analysing the variable ªrank ratioº an
assumption of proportional odds (i.e. that the odds for each increase in rank by a factor 1 would be the
same) had to be made, and thus this assumption should be kept in mind when the results are
interpreted. Furthermore, when using the term ªrank ratioº we had to assume that the social
dominance in these groups was linear, which may or may not be the case. However, if groups
of pre-calving cows are newly established, or dynamic, there is a risk of aggression due to the
14 / 19
establishment of dominance relations [
], especially near defendable resources. Cows become
maternally motivated due to hormonal change prior to calving (reviewed in [
]) and as
calving approaches an individual calving pen may become a valued site. In the present study,
dominant cows may have guarded the individual calving pens leaving them a higher chance of
calving there. Also being dominant they were less likely to be displaced from the pens with
permanently open gates. This suggestion may be supported by the finding that cows positioned
inside an individual calving pen when the second stage of labour was initiated, had a
significantly higher probability of calving in the pen. It is possible, that the choice of where to calve
was already taken when the second stage of labour was initiated, and thus, if a cow had been
guarding an individual calving pen, she was more likely to end up calving there. Moreover,
cows that changed their position after initiation of the second stage of labour mainly changed
from the group area to an individual calving pen (10 out of 16 cows). These cows were mainly
dominant cows (8 out of 10) and all cows moving from an individual calving pen to the group
area (n = 6) were subordinate (happened in sections of both experimental treatments, 3 from
each), further implying that dominance continued to play a role, even during the second stage
of labour. These findings have implications for the design of a motivation-based calving facility
as in this case, the facility only succeeded in proving an isolation opportunity for the dominant
cows even though there was always one individual calving pen per cow. More research is
needed in order to outline how to account for dominance in a motivation-based calving pen
design. Additionally, this highlights the role of the social hierarchy in a pre-calving group and
points out that farmers may have to consider that some cows need more protection if they are
to move away from the group at calving. Another, perhaps surprising result, in relation to this,
is that social dominance (ªrank ratioº) strongly correlated with the personality assessment
made prior to the experiment and at least one week prior to calving. This correlation showed
that bolder and more explorative cows were also the more dominant while shy cows were
more subordinate. This information combined with the above results may be useful for
farmers trying to identify cows needing assistance in order to enter individual calving pens and shy
cows may even have to be moved manually. In this experiment most shy cows calved in the
group area, and protection from competition for individual calving pens may be required.
Personality assessment may be a practical tool to assess dominance and to identify cows that may
have to be moved manually to an individual calving pen.
Another social aspect of pre-parturient groups, which farmers also need to be aware of, is
the presence of alien calves in the group. In this study, the presence of an alien calf lowered the
probability of the cows calving inside the individual calving pens, potentially because of an
attracting effect of either the calf as such [
] or the amniotic fluids in the fur of the new-born
]. Combining this knowledge with the described effects of dominance, these social aspects
may make a motivation-based calving system difficult to apply unless subordinate cows are
moved manually to an adjacent maternity pen.
The fact that not all cows calved in the calving pens does thus not imply a lack of learning
or knowledge of this opportunity, but rather a combination of competition and motivational
conflict. For a motivation-based calving facility to function, these issues have to be solved. As
discussed above, improving or increasing the quality of the isolation opportunity is important
especially when aiming to facilitate isolation seeking at the time where they are motivated to
do so. Competition and distractions are problems that may not be easily solved, but if a cow
moves to an isolated area before giving birth and if her calf is subsequently retained there, this
would nonetheless decrease the level of competition and distractions in the environment of
the cows. From this perspective, another possible approach may be to use the presumably
attracting effect of birth fluids [
] or amniotic fluid at specific points inside the calving pens.
As these substances have previously been shown to attract cows no earlier than 12 h prior to
15 / 19
giving birth [
], this may be an option to attract only the cows that are close to calving and
move them away from the herd before they (or their calves) become a distraction.
Additionally, this attraction may function as a somewhat neutral stimuli for cows that are not yet close
In this study we could not show that use of a functional gate combined with prior training
facilitated isolation seeking, as measured by the use of individual calving pens at the time of
calving. This may be due to the rather conspicuous instrumental task of opening the gate or
because the cows were trying to avoid a potentially confined situation where executing a flight
response may be difficult. Alternatively, the cows may not have been able to combine the learnt
response with the motivation to isolate. Social factors had a strong influence on the odds of a
cow calving inside an individual calving pen, with dominant cows having the highest odds of
occupying an individual calving pen in the moment of calving and the presence of an alien calf
reducing the likelihood of cows calving inside the individual calving pen. Therefore, social
factors have implications for the functionality of a motivation-based system. More research is
needed to fully outline the cognitive basis from which cows are able to learn and recall learnt
responses and whether these responses are overruled by other motivations once calving is in
S1 Video. Cow performing the last step in the training procedure for cows housed with
functional gates. The video mimics the training situation as this particular cow is not an
experimental cow. She was trained on randomly chosen pens and with the five other group
members present. Prior to this take, the cow has been successful in all the previous training
steps. At this stage the cow is trained to open the fully closed gate on her own and subsequently
enter the pen to reach the reward from the trainer facing the cow from inside the pen/the
group area (step 4 in the training procedure). The trainer ensured that no other cows followed
the focal cow into the individual calving pen at any point.
S1 File. Gate errors.
S2 File. Data. The data sheet provided for this study includes all necessary information in
order to recalculate the results of the study. The sheet has one horizontal line for each
individual cow in the experiment and each cow has its own identification number (`cow_number').
All subsequent columns has an explanation line on the top of the sheet and whenever data is
not available for the particular cow this is marked with a `.'. Two variables are one/zero
variables corresponding to 1 = ªyes and 0 = ªnoº.
We thank Jyden Bur A/S, Vemb, Denmark, for assisting in the development and test of the
one-cow gates and for providing equipment for the experimental sections. The farm staff at
Aarhus University, Foulum, Denmark and all involved students are gratefully acknowledged
for assistance throughout the study. In addition, we thank John Misa Obidah, Aarhus
University, Denmark, Frederik Rønne Pachler, Aarhus University, Denmark, Eva Garre, Ecole
16 / 19
Superieure DÂAgriculture, France and Paul Lasagesse, LyceÂe Faidherbe, Lille, France, for all
the technical assistance and help during the training sessions.
Conceptualization: Maria Vilain Rørvang, Mette S. Herskin, Margit Bak Jensen.
Data curation: Maria Vilain Rørvang.
Formal analysis: Maria Vilain Rørvang.
Funding acquisition: Margit Bak Jensen.
Investigation: Maria Vilain Rørvang, Margit Bak Jensen.
Methodology: Maria Vilain Rørvang, Mette S. Herskin, Margit Bak Jensen.
Project administration: Maria Vilain Rørvang, Margit Bak Jensen.
Software: Maria Vilain Rørvang.
Supervision: Mette S. Herskin, Margit Bak Jensen.
Visualization: Maria Vilain Rørvang.
Writing ± original draft: Maria Vilain Rørvang.
Writing ± review & editing: Maria Vilain Rørvang, Mette S. Herskin, Margit Bak Jensen.
17 / 19
18 / 19
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