Orgãos Sensitivos Dos Cavalos

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APPLAN-2933; No of Pages 20
Available online at www.sciencedirect.com
Applied Animal Behaviour Science xxx (2008) xxx–xxx

www.elsevier.com/locate/applanim
Sensory sensitivities: Components of a horse’s

temperament dimension
Léa Lansade *, Gaëlle Pichard, Mathilde Leconte
Laboratoire de Comportement, Neurobiologie et Adaptation, UMR 6175 Physiologie de la Reproduction et des
Comportements, INRA-CNRS-Université de Tours-Haras Nationaux, F-37380, Nouzilly, France
Accepted 27 February 2008
Abstract
Temperament is an important factor when working with horses. Behavioural tests have already been
developed to measure certain dimensions of a horse’s temperament (fearfulness, gregariousness, etc.). In
order to measure the temperament more precisely, our work aimed to identify a dimension which has already
been described in several species but not yet in horses, namely sensory sensitivity. Our study was based on
the definition of a temperament dimension as ‘‘a behavioural characteristic stable across situations and over
time’’. We designed several tests for each sense and then determined whether the responses observed were
correlated between situations and in time. The principle of the tests was to generate two stimuli of different
intensities for each sense (e.g. two different sounds) and to measure the intensity of the horse’s response
(N = 26). Using Spearman rank correlations, we tested whether the responses to these different stimuli were
inter-correlated. We repeated the same tests 5 months later to determine whether the responses were
correlated over time.
Within each sense, results show that the greater the horses’ response to one stimulus, the greater
their response to the other. For example, the reaction to the odour of cinnamon (time spent near
the source of the odour) was significantly correlated to the reaction to lavender (R = 0.53, p = 0.004).
The reactions to two different sounds or to two different tactile stimuli (von Frey filaments, or
contact of a brush on the body), were also significantly correlated (R = 0.59, p < 0.0001; R = 0.38,
p = 0.029). Finally, the reactions to two different tastes or to two visual stimuli tended to be correlated
(R = 0.27, p = 0.09; R = 0.27, p = 0.09). However, there was no significant correlation between the
responses to stimuli relating to different senses. Finally, except for the responses to odour, the responses
to other sensory stimuli showed stability over a 5-month period (e.g. tactile stimulation: R = 0.71,
p < 0.0001).
In conclusion, our study revealed characteristics which were stable across situations and over time. The
absence of links between the characteristics measured for the different senses suggests that a dimension for
* Corresponding author. Tel.: +33 2 4742 7279; fax: +33 2 4742 7743.
E-mail address: lansade@tours.inra.fr (L. Lansade).
0168-1591/$ – see front matter # 2008 Elsevier B.V. All rights reserved.
doi:10.1016/j.applanim.2008.02.012
Please cite this article in press as: Lansade, L., et al., Sensory sensitivities: Components of a horse’s
temperament dimension, Appl. Anim. Behav. Sci. (2008), doi:10.1016/j.applanim.2008.02.012
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2 L. Lansade et al. / Applied Animal Behaviour Science xxx (2008) xxx–xxx
each sense exists (e.g. tactile sensitivity) rather than a general sensory sensitivity dimension covering all the
senses.
# 2008 Elsevier B.V. All rights reserved.
Keywords: Horse; Equus caballus; Temperament; Sensory sensitivity; Behavioural tests
1. Introduction
Temperament is generally defined as a set of individual differences in behavioural tendencies,

called ‘‘traits’’ or ‘‘dimensions’’, that are relatively stable across various kinds of situations and
over the course of time (Goldsmith et al., 1987; Bates, 1989). These dimensions should be
independent from each other. According to Gosling’s review (2001), over 40 different
temperamental dimensions have been described in animals. The dimensions which have been
studied the most are emotionality, fearfulness, aggression, timidity, confidence, activity and
exploration. In horses, numerous dimensions have also been described. For instance, factor
analyses of responses to a temperament questionnaire completed by people working with horses
have identified dimensions such as anxiety, novelty seeking, understanding, trainability and
affability (Momozawa et al., 2003, 2005). Using experimental tests, some authors have found
other dimensions such as flightiness, sensitiveness, patience, willingness to perform (Visser et al.,
2001), fearfulness, gregariousness (Wolff et al., 1997; Lansade et al., 2006, 2007), activity and
reactivity to humans (Lansade et al., 2006).
In order to measure equine temperament more precisely, it is possible to identify dimensions
which have already been described in several species but not, or only partially, in horses. One
example is sensory sensitivity, which can be defined as the propensity of an individual to react to
a greater or lesser extent to sensory stimulations in its environment (olfactory, gustatory—often
grouped as oro-sensory characteristics, visual, auditory and tactile).
In his review of human sensory sensitivity, Dunn (2001) explains that ‘‘people have different
thresholds for noticing, responding to, and becoming irritated with sensations’’. He observes
that people with high-sensory sensitivity ‘‘notice sensory stimuli quite readily and more
sensory events in daily life than do others; they are easily distracted by movements, sounds or
smells while in groups of people, such as in class or at the movie theatre. They notice food
textures, temperatures and spices more rapidly than others. They may be uncomfortable with
clothing tags, elastic or certain fabric textures’’. Similar dimensions of sensory sensitivity have
been included in several human temperament models. For instance, Plomin (1983) has
described a dimension called ‘‘threshold of responsiveness’’, defined as the intensity level of
stimulation needed to evoke a response. Thomas and Chess (1977) observed ‘‘sensory
threshold’’ as one of nine basic traits that distinguish children. Finally, Aron and Aron (1997)
developed the highly sensitive person scale (HSPS) to evaluate the sensory sensitivity to
internal stimuli (pain, etc.) or external stimuli (noise, light, etc.). According to these authors,
‘‘sensory processing refers to a difference not in the sense organs per se but to something that
occurs as sensory information is transmitted to or processed in the brain’’. Therefore, sensory
sensitivity seems to differ from perception in that an individual can perceive stimulation but
does not respond to it.
Similar dimensions have been described in dogs. Among the 20 dimensions assessed by the
trainers at the Royal Guide Dogs Association of Australia, two are linked to sensory sensitivity:
body sensitivity (the dog’s level of responsiveness to tactile stimuli) and hearing sensitivity (the
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L. Lansade et al. / Applied Animal Behaviour Science xxx (2008) xxx–xxx 3
dog’s level of responsiveness primarily to auditory stimuli) (Murphy, 1998). These dimensions
are used to select dogs using behavioural tests.
At least two authors have mentioned a similar dimension in horses. In his paper on horse
personality, Mills (1998) defined sensitiveness as the ease with which performance is affected by
environmental disturbance. In their study on temperament, Visser et al. (2001) identified a
dimension of sensitiveness from responses to a novel object test. Very sensitive horses are defined
as those which focus strongly on the novel object. The authors also hypothesised that horses that
were very sensitive to novel objects were the most fearful. However, since this dimension refers
only to visual sensoriality and is linked to fearfulness, it differs from the dimensions in humans
and dogs described above, which are multisensory and are not linked to fearfulness. In this paper,
our purpose was to identify a multisensory dimension, called sensory sensitivity, which is
independent of other temperament traits.
Our study had the following five aims:
(1) To create several behavioural tests to measure a horse’s responses to sensory stimulations for
each of the five senses (e.g. olfactory and tactile) and to describe its level of response
according to the stimulus intensity.
(2) To test the primary criteria of the definition of a temperament dimension: are the responses to
these tests stable over time?
(3) To test the secondary criteria of the definition of a temperament dimension: are the responses
to these tests stable across situations?
(4) To determine whether there are sensitivities for each sense (e.g. auditory and olfactory), or
whether there is a single overall sensitivity covering all the senses.
(5) To determine whether the sensitivity tests created in the study measure a new dimension
or one already described (such as fearfulness, gregariousness, activity or reactivity to
humans).
2. Animals, materials and methods
2.1. Animals
Twenty-six 3- and 4-year-old Welsh female ponies, bred at INRA Nouzilly and accustomed to being
handled (they were regularly haltered and tethered), were used. Before the present experiment, they lived
outdoors during summer and indoors during winter. They were individually identified with ear tags. During
the experiment, animals were kept at pasture except during the tests (3 h per day). They were fed daily with
concentrated feed (pellets). Water was available ad libitum.
2.2. Experimental system and habituation
All the tests used in this study were designed during a preliminary study conducted on 10 Welsh
female ponies prior to data collection for the study itself. The study was carried out in straw-bedded
loose boxes (4 m 6 m). There were two horses per box because many of them panicked when
alone. Fourteen ponies were tested in the morning and 12 in the afternoon. Animals were tied up
for 3 h one behind the other with two tethers fixed to either side of the halter and attached to two walls
(Fig. 1). The tethers kept the ponies parallel to the walls, 60 cm from one wall and 3.40 m from the
other. The loose boxes were equipped with separation planks so that the ponies could not see the
reactions of their conspecifics. A fixed feeding trough was placed in front of each pony and a
video camera beside it to film its reactions. The position of the ponies was determined randomly
every day.
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Fig. 1. Overhead view of the horses’ position during the tests. ( ) Video camera; (—) full walls; ( ) separation
planks (1.80-m high); ( ) feeding trough; (- - - -) tether; ( ) entrance.
The animals were tied up in this way for three consecutive hours for the 5 days preceding the first and
third series of tests (see below). At the end of these 5 days, all the ponies were accustomed to being tied up
and stood quietly throughout this period.
2.3. Experimental procedure
The general principle of the tests was to generate the same stimulation for each sense (odour, taste,
visual, sound or tactile) at five different intensities and to observe the responses of the animals. Different
intensities of a single stimulus were tested to increase the probability of observing inter-individual
variability more precisely. The five intensities were selected during the preliminary study so that they
produce a satisfactory inter-individual variability of responses.
For each type of stimulus (e.g. odour), one intensity was tested per day, except for the sound test (see
below). The intensities were generated randomly. The intensities were presented in the same order for all
animals. If the test order was balanced through all individuals, we could not compare the individuals
between them, whereas this is the main aim of this study.
To assess response stability over time, two series of identical tests were repeated at a 5-month interval, in
May and October (series A and A0 , Table 1).
To assess the response stability for each sense across situations, the ponies were tested with two
successive series of different stimuli, at a 2-week interval (Table 1). For example, for the odour test, the
ponies were first subjected to a cinnamon-odour series (series A; 5 days) and a week later to a lavender-
odour series (series B; 5 days).
All of the tests were filmed except for the food and tactile tests. Different behaviours (latency and
frequency) were recorded, but only one was kept per test to limit the number of correlations in the statistical
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Table 1
L. Lansade et al. / Applied Animal Behaviour Science xxx (2008) xxx–xxx

Intensity distribution order of the different sensory stimulus
Series A Series B (after 1 week) Series A0 (after 5 months)
Scented compound volumes (ml) Cinnamon Lavender Cinnamon
5 2 4 1 3 5 2 4 1 3 5 2 4 1 3
Repulsive preparation volumes (ml) Spicy preparation Meat preparation Spicy preparation
8 2 1 6 4 8 20 30 15 25 8 2 1 6 4
Visual stimuli sizes (mm) Blue string diameter Orange sticky tape Blue string diameter
25 5 15 5 10
3 0.5 1 2 4 3 0.5 1 2 4
40 10 25 5 15
Sound intensities (dB) Beep Shrill beep Beep
7 35 55 33 67 70 35 55 33 67 70 35 55 33 67
Tactile stimuli intensities (g/matter) Filaments Filaments
300 0.6 0.02 0.008 8 300 0.6 0.02 0.008 8
Stifle-haunch axis stimulation Stifle-haunch axis stimulation Stifle-haunch axis stimulation
W HPB P R C W HPB P R C W HPB P R C
W: wood, HPB: hard paintbrush, P: plastic, R: rubber, C: cotton.
5
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analysis. Using many different behaviour responses for the correlation analysis, would have increased the
risk of finding significant results by chance (Martin and Bateson, 1993). The most relevant variable was
therefore selected. They are indicated following the description of each test.
2.3.1. Odour test

The aim of this test was to evaluate the ponies’ response to an odour source. A 2-cm diameter hole was
made in the bottom of the feeding trough in front of each pony. A Petri dish without a lid and containing a
compress soaked with a scented compound was fixed under this opening, out of the pony’s sight to avoid any
visual stimulation. One scented compound volume was presented to the ponies on each of the 5 days in a
series (Table 1). Cinnamon was used in series A and A0 , and lavender in series B. Each test lasted 2 min. The
time each animal spent with its nose in a zone near the odour source (30 cm maximum from the source) was
recorded. This time was called ‘‘time spent in the odour zone’’. The 30-cm distance was chosen because it
was half the length of the tether used to tie the pony up near the feeding trough.
2.3.2. Food test

The aim of this test was to assess each pony’s response to a repulsive flavour mixed with the usual feed
(pellets). First, we verified that all the ponies ingested 100 g of pellets (quantity used by Goodwin et al.
(2005a,b)) in less than 3 min. Then, on each of the 5 days, each individual received 100 g of pellets to which
a specific volume of a repulsive spicy or liquid meat preparation had been added at five different
concentrations (Table 1). The experimenter left the box immediately after pouring the food into the
feeding trough. After 3 min, the unconsumed pellets were removed from the feeding trough and weighed.
The choice of repulsive flavour was made after the preliminary study. The tested flavours were quinine,
vinegar, pepper, essential oil of cinnamon and thyme, a meat preparation (Viandox1), a spicy sauce
containing chilli pepper (Tabasco1), a dehydrated vegetable stock (Kubor1) and a spicy preparation based
on the latter two flavours (base mixture: 0.4 g of dehydrated vegetable stock per millilitre of spicy sauce,
made up to 10 ml with water). The meat and spicy preparations were selected because they were the only
flavours that were refused by some ponies. In accordance with Goodwin et al. (2005a,b), the quantity of
unconsumed pellets at the end of the test was measured.
2.3.3. Visual stimulus test

This test consisted in lowering a visual stimulus in front of the pony’s eyes, 1.2 m from the floor. In series
A and A0 , 20-cm lengths of blue twisted baler string of different diameters were used, and in series B orange
sticky tape of different sizes. One stimulus was presented to the ponies every day (Table 1). To do this, an
experimenter stood in the adjacent box behind the separation plank, out of view of the tested subject, and
slowly lowered the visual stimulus in front of the subject with the help of a nylon thread. The frequency with
which the pony looked at the stimulus (ears pricked forward in the direction of the stimulus) was recorded
for 2 min, from the moment the stimulus started to be lowered.
2.3.4. Sound test

This test consisted in generating a brief sound near the pony and assessing its responses. A transmitter
was placed in the adjacent loose box, 6 m from each pony. In series A and A0 , a beep (fundamental
Table 2
Scale of response intensity marks during the sound emission
Mark Reaction
0 No reaction
1 Ear movements (the pony turned one or two ears)
2 Lateral head movement (the pony turned the head to look for the sound origin)
3 Head straightening up in vigilance position (the head raised high and fixed, the two ears pointed
forward, the position held for more than 2 s)
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Fig. 2. Photography of the instruments used during the tactile tests. On the left, a von Frey filament; on the right, the
instrument used for the stifle-haunch axis stimulation test.
frequency: 1000 Hz, maximal frequency: 1000 Hz, duration: 1.5 s) was used, and in series B a shrill beep
(fundamental frequency: 1400 Hz, maximal frequency: 9800 Hz, duration: 0.23 s). For this test, the ponies had
to stay totally still to avoid acoustic interference that could bias the results. To meet this requirement, the
animals were tested in groups of just two ponies in the test stable (because it is easier to keep two rather than 12
ponies totally motionless at the same time). To avoid moving the ponies, several sound intensities were
generated on the same day, three the first day and two the second. These intensities are presented in Table 1.
There was a minimum of 40 s between each stimuli. A mark was allocated according to the response (Table 2).
2.3.5. Tactile tests

In contrast to the previous stimulations, the two tactile tests were based on different principles and are
therefore described separately below. They were both carried out during series A and A0 to verify their
stability over time and were always performed by the same experimenter.
2.3.5.1. Von Frey filaments test. For this test, a von Frey filament was applied at the base of the withers
(von Frey filaments, Stoelting, IL, USA; already used on horses by Redua et al. (2002) to evaluate the pain
response after a sutured skin incision). These filaments consist of a hard plastic body extended by a nylon
thread (Fig. 2). They are calibrated to exert a specific force on the skin. A different force was applied every
day (Table 1). They were applied perpendicularly on the animal’s skin until the nylon thread started to bend.
Trembling of the platisma muscle (behaviour used by horses to drive away flies—see Saslow, 2002) was
recorded. The response was coded in a binary form (trembling/not trembling).
Table 3
Scale of bodily reaction intensity during the stifle-haunch axis stimulation
Mark Reaction
0 No reaction
0.5 Stomach trembling
1 Stomach tensing
1.5 Stomach tensing and breathing interrupted
2 Stomach and croup tensing and breathing interrupted
2.5 General tensing of all muscles and breathing interrupted
3 General tensing of all muscles, breathing interrupted and step to the side
3.5 General tensing of all muscles, breathing interrupted and kicking
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Table 4
Definitions of tested dimensions
Dimension name Definition
Fearfulness Propensity to react to a variety of potentially threatening situations
Social motivation Propensity to react to social separation
Locomotor activity Propensity to demonstrate locomotor activity in various situations
Reactivity to humans Propensity to react to a passive or active human
2.3.5.2. Stifle-haunch axis stimulation test. This test consisted in stimulating the stifle-haunch axis with 3-
cm wide instruments of different hardness (Fig. 2). A different hardness of instrument was used every day
(Table 1). The stimulation caused a bodily reaction. The experimenter quickly moved the instrument
upwards along the tuft of the stifle-haunch axis, keeping an even pressure. The intensity of the bodily
reaction was assessed on a pre-defined scale (Table 3).
2.3.6. Description of the temperament tests

To check that the sensory tests presented above measured a new temperament characteristic and did not
correspond to dimensions already described, tests used by Lansade et al. (2006) to characterise fearfulness,
gregariousness, reactivity to humans and locomotor activity were also carried out. These dimensions are
defined in Table 4. These tests were carried out in the week between series A and B.
Except for the social isolation test, all the tests were carried out in a loose box (4 m 6 m) familiar to the
animals. An audience pony was tied up outside the box, visible to the tested pony, to avoid social isolation
interfering with the other measured characteristics. The behavioural variables analysed had been selected by
Fig. 3. Novel object and novel area tests experimental system.
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Lansade and Bouissou (2003), Lansade (2005), Lansade et al. (2007) because they are considered to be the
best indicators of the dimensions studied, in particular because of their stability.
2.3.6.1. Fearfulness tests. To characterise fearfulness, three test situations were used involving novel
stimuli (novel object, crossing an unknown surface) and a sudden stimuli (opening an umbrella).
2.3.6.1.1. Novel object test. A novel object (traffic cone topped with a plume of red and white striped
plastic bands) was placed near the entrance (Fig. 3) for 5 min. The frequency of sniffing and nibbling the
object was recorded.
2.3.6.1.2. Crossing an unknown surface test. For this test, the floor of the loose box was divided into
three zones (Fig. 3). The first zone (1.5 m 4 m) corresponded to a starting zone and the third zone (towards
the audience pony, 1.5 m 4 m) was an arrival zone. The second zone measured 3 m 4 m. The arrival
zone contained a bucket of pellets, familiar to the animals.
Before the test, the ponies underwent a habituation stage during which they learnt to go from the starting
zone to the arrival zone containing the bucket. To do so, an experimenter led the pony to the starting zone
using a halter. The experimenter released the pony so that it was free to go to the arrival zone to eat. The time
between crossing the starting line and taking a mouthful of pellets was measured. A line was considered
crossed when the two front hooves stepped over the line. As soon as the pony had eaten a mouthful of pellets
it was taken back to the starting zone. If the animal did not eat pellets within 40 s, it was taken back to the
starting zone and the attempt was considered invalid. Two consecutive attempts had to be validated to start
the test.
During the test, a white sheet (2.4 m 4 m) was placed in the second zone and the procedure was the
same as before except that there was only one attempt. The time taken to go and eat was recorded. A
maximum time of 300 s was allocated. If the pony did not eat any pellets in this time, the test was terminated
and a time of 301 s was assigned.
2.3.6.1.3. Umbrella opening. This test consisted in opening a black umbrella in front of the animal
when it was eating. Before the animal entered the loose box, a bucket of pellets was placed near the entrance
and the umbrella was placed in a closed position 1 m from the bucket, 1 m off the ground. When the animal
had been eating with its head in the bucket for more than 3 s, the umbrella was opened and the chronometer
was started. The test stopped when the pony started eating again. The time taken to start eating again was
recorded. A maximum time of 300 s was allocated. If the pony did not eat pellets within this time, the test
was terminated and a time of 301 s was assigned.
2.3.6.2. Social isolation test. This test consisted in isolating the pony from its conspecifics with no
possibility of seeing or communicating with them. The pony was taken from its group and led to a familiar
corridor (2 m 15 m) where the animals were herded daily between tests. During this test the pony was
totally alone for 5 min and the frequency of neighing was recorded.
2.3.6.3. Reactivity to humans test. To characterise reactivity to humans, two tests were performed.
2.3.6.3.1. Known passive human test. An experimenter (always the same one) entered the loose box
and stayed motionless beside a wall for 5 min. The frequency of sniffing and nibbling the human was
recorded.
2.3.6.3.2. Known active human test. At the end of the known passive human test, the same
experimenter moved slowly towards the pony and when 1 m away tried to put his hand on the pony’s nose
(for 2 s). The time to touch the nose was recorded. A maximum time of 120 s was allocated. If the
experimenter did not manage to touch the pony’s lower front part of the face in this time, the test was
terminated and a time of 121 s was assigned.
2.3.6.4. General locomotor activity assessment. During the novel object test, the social isolation test and
the passive human test, the loose box was divided into six areas of equal size (Fig. 3) and the total number of
areas the horse crossed was recorded. It has been shown that this locomotor activity reflects another
dimension than the three previous ones (Lansade, 2005; Lansade et al., 2006).
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2.4. Statistical analysis
Due to the small number of subjects and the nature of measurements, non-parametric statistics were
used. Data were analysed with the XLSTAT software.
2.4.1. Description of the animals’ responses according to stimulus intensity

For the first series of tests (series A), the responses to each of the five intensities for each stimulus type
were compared using a Friedman test. When the Friedman test was significant, intensity responses were
compared in pairs with Wilcoxon tests. When responses were continuous data (all the responses except for
the filament test), the changes in the responses according to the stimulus intensity were shown in box–plot
form (in accordance with Tukey’s model, 1977). When the responses were recorded in binary form (filament
test), the numbers of individuals who responded to each intensity were analysed and shown in histogram
form.
2.4.2. Stability over time

All of the following analyses were performed on the sum of the responses of the five tested intensities for
each series and each pony. For example, for the odour test, the five ‘‘times in the odour zone’’ measured for
each series were summed per individual. The sum of responses was used to avoid having too many variables
for correlation analysis. In addition, these sums present a greater variability than the responses to a single
intensity (when looking at the responses to a single intensity, many animals show no response and have a
‘‘0’’ score; in this case, correlation analyses are not possible).
To test the stability of the responses over time, Spearman correlations were calculated between the sums
of the responses obtained during series A and A0 .
2.4.3. Stability across situations

To test the stability of the responses across situations, Spearman correlations were calculated between
the sums of the responses obtained during series A and B.
2.4.4. Correlations between the sensory tests

Spearman correlations were calculated between the sums of the responses to the different sensory tests of
series A (e.g. between the sums of the responses to the odour test and the sound test).
2.4.5. Correlations between the sensory tests and the pre-existing temperament tests
Lastly, possible links between the sensory tests and the other temperament tests were investigated.
Spearman correlations were calculated between the sums of the responses to the sensory tests of series A and
the different variables measured during the three fearfulness tests, the social isolation test, the reactivity to
human test and the general locomotor activity assessment.
3. Results
3.1. Description of the animals’ responses according to stimulus intensity (series A)
3.1.1. Odour test

The ‘‘time spent in the odour zone’’ did not change significantly between the first four
concentrations but increased for the last one (Fig. 4).
3.1.2. Food test

The quantity of unconsumed pellets increased significantly with the concentration of the
repulsive flavour, except between 2 and 4 ml (Fig. 5).
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Fig. 4. Odour test, changes in the ‘‘time spent in the odour zone’’ (s) according to the scented compound volume (ml).
Legend of the box–plots presented in Figs. 4–9: (&) 25 (Q1); 75% (Q3) ;(*) >3 (Q3 Q1); (*) <3 (Q3 Q1);
value >(Q1 1.5 (Q3 Q1), Q1); value <(Q3, Q3 + 1.5 (Q3 Q1)); — median value. Different letters (X, Y)
indicate significant difference ( p < 0.05) in the Wilcoxon test.
Fig. 5. Food test, changes in the unconsumed pellet quantity (g) according to the repulsive flavour concentration (ml).
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Fig. 6. Visual stimulus test, changes in the number of ‘‘looks at the stimulus’’ according to the string size (mm).
Fig. 7. Sound test, changes in the response mark according to the sound intensity (dB).
3.1.3. Visual stimulus test

There was no significant change in the number of ‘‘looks at the stimulus’’ according to the size
of the stimulus (Fig. 6).
3.1.4. Sound test

The response mark significantly increased with the intensity of the sound, except between 55
and 67 dB (Fig. 7).
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Fig. 8. Von Frey filaments test, changes in the number of horses producing the response according to the force of the
filament (g).
Fig. 9. Stifle-haunch axis stimulation test, changes in the response mark according to the instrument hardness.
3.1.5. Tactile tests

3.1.5.1. Von Frey filaments test. There was a significant increase in the number of horses
producing the response between the 0.008 and 8 g filaments, and then a significant decrease
between the two higher forces (8 and 300 g, Fig. 8).
3.1.5.2. Stifle-haunch axis stimulation test. There was no significant change in the bodily
reaction mark with different instrument hardness (Fig. 9).
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Table 5
Spearman correlations between series A and A0 (stability over time) and between series A and B (stability across
situations)
Tests Stability over time Stability across situations
(series A and A0 (series A and B correlations)
correlations)
Food test R = 0.50 R = 0.27
p = 0.005 p = 0.09 tendency
Odour test R = 0.18 R = 0.53
NS p = 0.004
Visual test R = 0.38 R = 0.27
p = 0.028 p = 0.09 tendency
Sound test R = 0.45 R = 0.59
p = 0.01 p < 0.0001
Tactile tests Filaments: R = 0.71, p < 0.0001 R = 0.38
Stifle-haunch axis stimulation: R = 0.53, p = 0.003 p = 0.029
3.2. Stability over time
The correlations between series A and A0 were significant for the sum of the responses to the
food, visual and sound stimulus tests, and the von Frey filament and stifle-haunch axis stimulation
tests, but not to the odour test (Table 5).
3.3. Stability across situations
The correlations between series A and B were significant for the sum of the responses to the
odour, sound and tactile tests. The correlations tended to significance (**p < 0.1) for the food
and visual stimulus tests (Table 5).
3.4. Correlations between sensory tests
The only significant correlation appeared between the responses to the food and odour test
(R = 0.40, p = 0.022). The more the ponies stayed in the area near the scented compound the
more they ate flavoured pellets and vice versa.
3.5. Correlations between the sensory tests and the other temperament tests
No significant correlations were found between the sums of responses calculated for the
different sensory tests and the variables measured during the other temperament tests.
4. Discussion
4.1. Description of the animals’ responses according to the stimulus intensity
Since this is the first time these tests have been performed, it is of interest to describe the
pony’s response curve according to the intensity of the different sensory stimuli. The results
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presented below give only an indication of the phenomenon. As the intensities of the stimuli were
presented in the same test order for all individuals, it is in fact possible that the difference in
responses observed for each intensity were due to the test order.
The response to odour was observed to increase as the intensity increased, but only for the
higher concentration. It is possible that the horses did not detect the presence of the odour below a
concentration of 5 ml. However, this hypothesis is improbable, as although few studies have
investigated horse olfaction, several anatomical particularities indicate that horses have a
particularly well-developed sense of smell. For instance, the brain contains extremely large
olfactory bulb with a convoluted surface, the nose can move large volumes of air at one breath
and trap large numbers of molecules (Saslow, 2002), and the nostrils are separated and pointed in
opposite directions, allowing stereo olfaction in the localization of olfactory sources (Stoddart,
1980). Thus, it is more likely that the horses were able to detect the odour for concentrations
below 5 ml, but were only attracted by a strong odour (5 ml). It should also be noted that the 5-ml
intensity was presented first in the series and it is possible that the animals reacted less to the later
presentations because they were already familiar with the odour.
For the food tests, the animals’ responses increased with the stimulus intensity. The stronger
the repulsive flavour, the more the animals refused to eat it. Several studies have examined the
effects of adding different flavours to concentrates (e.g. Goodwin et al., 2005a,b), but did not
investigate the effect of the concentration of the flavour on consumption, although they did
observe a variability of consumption between individuals.
For the visual tests, the responses did not change with stimulus intensity. The lack of
difference between stimuli sizes might have prevented the horses from responding differentially.
A greater variety of stimulus sizes was used during the preliminary study, from very small to very
large, and the responses increased with size. However, for the large stimuli, a ceiling effect was
observed: all the horses looked frequently at them, with very low inter-individual variability. For
this reason, the size of the visual stimuli was reduced. However, it is interesting to notice that
some horses responded to the smallest stimuli, indicating sufficient acuity to see them, although a
number of papers (Timney and Keil, 1992; Saslow, 1999) have suggested that horses do not have
very good visual acuity (the ability to resolve details of a visual pattern), especially in a
motionless scene. The fact that the object moves can help them to detect it.
For the sound tests, the animals’ responses increased with the stimulus intensity. The louder
the sound, the more they adopted vigilant postures. According to Heffner and Heffner (1983),
horses’ hearing ranges from 55 Hz to 33.5 kHz, with their optimum hearing between 1 and
16 kHz, and the lowest average threshold of 7 dB. The intensities of the sounds used in this
experiment were comprised between 33 and 70 dB, with a frequency of 1 kHz for the first sound
and between 1.4 and 9.8 kHz for the second. Thus, each of the sounds emitted during the
experiment was perfectly audible to horses. This indicates that horses can perceive a sound but
not react to it, demonstrating that the behavioural responses we observed were not directly linked
to their ability to perceive the stimulus.
Finally, for the tactile tests, the responses to the stifle-haunch stimulation test did not change
with stimulus intensity. The pressure exerted by the different instruments was probably too
similar to cause different intensities of reaction. By contrast, for the von Frey filament test, the
number of horses which reacted to the filament increased up to a pressure of 8 g and then
decreased when the pressure became too high (300 g). This result could be connected with the
biological function of the platisma muscle trembling: to drive flies away through a skin rippling
(Saslow, 2002). The pressure exerted by the smallest filament may have been too light to be
detected or associated with an insect by all the horses. On the other hand, the pressure exerted by
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the biggest filament may have been too strong to be associated with an insect. In addition, 73% of
the ponies were observed to react to the smallest filament whereas no humans perceived it when it
was applied to the pad of their index finger (personal observation of 26 men and women, tested
according to the touch-test sensory evaluator instructions; North Coast Medical, Morgan Hill,
CA). This result is in accordance with Saslow (2002) who reported in her review on the
perceptual world of horses that ‘‘horses can react to pressures that are too light for the human
feel’’.
4.2. Are the responses stable over time?
The second aim of this study was to examine whether the responses to the tests were stable
over time. To achieve this, two identical series of tests were performed at a 5-month interval and
correlations between them were investigated. Except for the odour test, the more animals
responded to the stimulus in the first series of tests, the more likely they were to respond to the
stimulus in the second series test performed 5 months later. This result is in line with Dunn (2001)
who found that sensory processing patterns in humans are stable across the life span.
This result also shows that the stability over time of the responses to sensory stimuli may not
be affected by the season, since the first series of tests was performed in spring and the second in
autumn. It could have been thought that the stability of the reactions, particularly for the tactile
stimuli, would be influenced by the thickness of the coat (thicker in autumn than in spring), the
mare’s hormonal cycle or the presence of many flies in spring (which could have caused a
habituation to the tactile touch), but this is not supported by our results.
4.3. Are the responses stable across situations—within each sensoriality?
The third aim was to determine whether the responses were stable across situations. For that
purpose, two different stimulations were generated for each sense (e.g. two different sounds), and
we assessed whether the responses were correlated.
Significant correlations were found between the two different series of odour, sound and
tactile tests. These correlations show that within each of these sensorialities, the same animals are
always more sensitive, whatever the stimulation used. Thus, for the auditory sense, the more the
horses reacted to the first sound, the more they reacted to the second. For the tactile sense, the
more the horses trembled with the von Frey filament, the more they reacted to the stifle-haunch
axis stimulation. Finally, the longer the horses spent in the zone near the cinnamon source, the
longer they spent near the lavender source. In addition, this last result shows a stability of the
response to odours over a period of 2 weeks, indicating stability over a short if not a long period.
The correlations between the two different series of food and visual tests tended towards
significance. For the food tests, in contrast to the second flavour (meat preparation), the spicy
preparation produced a sensation of irritation due to the capsaicin in the chilli pepper. This
difference between the two flavours could explain the low correlation between the two tests. It is
possible that for high concentrations of the spicy preparation, the ponies did not react to a novel
taste, but to the sensation of irritation, which involves the trigeminal system (Verhagen and
Engelen, 2006). It would be of interest to repeat this experiment, using two flavours which do not
involve the trigeminal system.
The moderate correlation between the responses to the two visual tests can be explained by the
fact that the first stimulus (blue string) was more familiar than the second (orange sticky tape), as
the horses had frequently seen the blue string used to tie up bales of straw, whereas they had never
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seen the tape before. Thus, the parameter of ‘‘novelty’’ was more prevalent in the second stimulus
than in the first, which could bias the result and reduce the response stability. Moreover, it may be
possible that these two stimuli have not the same sensory characteristics (e.g. not the same
odour).
In summary for the last two sections (4.2 and 4.3), the responses to most of the tests show
stability over time and across situations. Thus, these tests measure not only sensitivity to a given
stimulus at a given time, but also sensitivity to different stimuli relating to a single sense,
whatever the period of testing.
However, although the two stimulations sought to evaluate stability across situations are
different, the context in which they were performed was usually the same (the horse tethered in a
loose box). To explore the stability across situations of these sensitivities in more detail, it would
be interesting to perform the stimulations in different contexts. However, to standardize the tests,
a very quiet environment is required, without external stimuli which could disturb the test, and
such a situation is hard to find.
4.4. Is there a sensitivity for each sense or a global sensitivity, whatever the sense?
The fourth aim was to examine whether there is a sensitivity for each sense or a global
sensitivity covering all the senses. For that purpose, we examined whether there were links
between the responses to the tests relating to each of the five senses. Results only show a
significant correlation between the responses to the odour test and the food test. Ponies that
stayed a long time in the odour zone ate more flavoured pellets and inversely. This link is as to be
expected as both are oro-sensory characteristics. Thus, the food test measures a gustato-olfactory
sensitivity rather than a gustatory sensitivity. However, that does not mean that a gustatory
sensitivity independent of olfactory sensitivity does not exist, but this could not be demonstrated
by our food test. To do so would require using a taste without odour that is also repulsive to
ponies, something we have been unable to do so far.
The absence of other links between tests suggests that there are independent dimensions of
sensitivity for each sense (except for the gustatory sense) rather than a general sensory sensitivity
dimension. For example, horses may be very sensitive to tactile stimulation, but not to auditory
ones. This result is in line with Dunn’s study on humans (2001) in which he found that ‘‘people
may be more sensitive for some types of sensory input, while being less attentive for other types
of sensory input’’.
4.5. Are these sensitivities linked to other temperament dimensions already described?
Finally, we investigated whether there were any differences between these dimensions of
sensitivities and those classically reported. Correlation analyses performed between responses to
sensitivity tests and parameters measured during fearfulness, gregariousness and reactivity to
humans tests and locomotor activity assessment do not permit to identify links between them.
Thus, the different sensitivities may be independent of each other. However, this conclusion must
be confirmed, particularly because our sensitivity tests do not cover the whole range of sensibility
and thresholds of detection for each sense.
However, if visual sensitivity is in fact independent of other dimensions, this would prove that
it is different from the ‘‘sensitiveness’’ dimension found by Visser et al. (2001) which was linked
to fearfulness. More generally, the possible independence of the sensitivities from other
dimensions is in line with Murphy’s study (1998) on dogs in which he distinguishes two different
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temperament dimensions independent of the others: hearing sensitivity and body sensitivity. In
humans too, Thomas and Chess (1977) and Plomin (1983) consider sensitiveness as an
independent dimension. However, Dunn (2001) takes a different point of view, whereby sensory
sensitivity may underlie temperament and personality manifestations. For instance, sensory
sensitivity may be a facet underlying the temperament traits of persistence, ‘‘because without
being distracted by external stimuli, persons can persist at task performance’’. This question
requires further investigation.
4.6. Are these results show more temperament traits or sensitivity traits related to
physiology of each individual?
Finally, the question arises as to whether our data show more temperament or sensitivity traits
related to the physiology (and not the psychology) of each individual. For instance, for visual
sensitivity, was the variability of responses observed between individuals due to differences in
their visual acuity or to psychological differences? In other words, the different levels of response
may have been due to the relatively poor vision of some animals. From our data, it is not possible
to draw a firm conclusion for all the tests. However, for the sound test, a previous experiment
(Heffner and Heffner, 1983) indicates that all the sounds emitted during the tests are audible to
horses, showing that animals can perceive a sound but not respond to it. This provides evidence
for the notion that the variability of responses observed was probably due to temperamental
rather than physiological differences. Aron and Aron (1997) discussed this point at length,
concluding that ‘‘sensory processing refers to a difference not in the sense organs per se but to
something that occurs as sensory information is transmitted to or processed in the brain’’.
However, further experiments are required to determine precisely whether the level of
responsiveness is due to physiology, psychology, or both. It would be of particular interest to test
the visual evoked response (VER), the brainstem auditory evoked response (BAER) and the
somatosensory evoked response (SSER) of the horse when presented with different stimuli. A
method of testing the BAER in non-anesthetized horses and ponies has already been developed
(Marshall, 1985). The aim would be to determine whether these evoked responses are correlated
with the differences observed in behavioural responses, from which it would be possible to
determine whether there is a physiological basis for the behavioural differences observed
between animals. The physiological basis of certain human temperament dimensions (such as
extraversion) have already been reported (for a review, see Stelmack, 1990).
4.7. Practical application
This study could have practical relevance. Generally, temperament is an important factor
when working with horses; an Australian study has shown that temperament is the most
important characteristic of a Pony Club horse (Buckley et al., 2004). In particular, it would be
interesting to determine whether visual, auditory and tactile sensitivities are linked to sensitivity
to the rider’s aids, such as reins or leg aids, tongue clicking or other sound or visual indications. If
these links exist, the corresponding tests could be used to predict the ease with which the horses
would naturally respond to the aids. It is also possible that very sensitive horses for these senses
could be easily disturbed by various stimuli when they are ridden. To be able to select horses
according to these sensitivities would enable them to be directed towards certain equestrian
disciplines or certain types of rider. Finally, the gustato-olfactory sensitivity could also have an
impact on horse management; horses that are more sensitive to these senses can be difficult to
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feed or water, particularly when travelling (e.g. for show jumping competitions) when food and
water can have unfamiliar tastes or odours. Such horses may require careful management.
5. General conclusion
To conclude, our study revealed four temperament dimensions for horses (tactile sensitivity,
gustato-olfactory sensitivity, auditory sensitivity and visual sensitivity), which are stable across
situations and over time. An olfactory sensitivity dimension may also exist but with only short-
term stability. The absence of links between the four dimensions suggests that a dimension of
sensitivity specific to each sense may exist rather than a general sensory sensitivity dimension.
However, this possibility needs to be confirmed. Moreover, these dimensions seem to be
independent of dimensions already described such as fearfulness, gregariousness, activity or
reactivity to humans. Finally, it could be now interesting to examine if the differences of
behavioural responses observed between animals have a physiological basis or not.
In addition to its fundamental interest, this study could have practical implications for horses.
Identifying different types of horses according to their level of sensitivity would allow them to be
ridden and managed more appropriately.
Acknowledgments
The National French Stud: ‘‘les Haras Nationaux’’ funded this experiment. The authors are
grateful to Guy Duchamp (INRA Nouzilly, France) and his staff for allowing the use of the
animals and facilities. We would also like to thank Chantal Moussu and Fabien Cornilleau for
their participation in collecting the data.
References
Aron, E.N., Aron, A., 1997. Sensory-processing sensitivity and its relation to introversion and emotionality. J. Pers. Soc.
Psychol. 73, 345–368.
Bates, J.E., 1989. Concepts and measures of temperament. In: Kohnstamm, G.A., Bates, J.E., Rothbart, M.K. (Eds.),
Temperament in Childhood. Wiley, New York, pp. 3–26.
Buckley, P., Dunn, T., More, S.J., 2004. Owners’ perceptions of the health and performance of Pony Club horses in
Australia. Prev. Vet. Med. 63, 121–133.
Dunn, W., 2001. The sensations of everyday life: empirical, theoretical, and pragmatic considerations. Am. J. Occup.
Ther. 55, 608–620.
Goldsmith, H.H., Buss, A.H., Plomin, R., Rothbart, M.K., Thomas, A., Chess, S., Hinde, R.A., McCall, R.B., 1987.
Roundtable: what is temperament? Four approaches. Child Dev. 58, 505–529.
Goodwin, D., Davidson, H.P.B., Harris, P., 2005a. Selection and acceptance of flavours in concentrate diets for stabled
horses. Appl. Anim. Behav. Sci. 95, 223–232.
Goodwin, D., Davidson, H.P.B., Harris, P., 2005b. Sensory varieties in concentrate diets for stabled horses: effects on
behaviour and selection. Appl. Anim. Behav. Sci. 90, 337–349.
Gosling, S.D., 2001. From mice to men: what can we learn about personality from animal research? Psychol. Bull. 127,
45–86.
Heffner, R., Heffner, H., 1983. Hearing in large mammals: the horse (Equus caballus) and cattle (Bos taurus). Behav.
Neurosci. 97, 299–309.
Lansade, L., Bouissou, M.F., 2003. Caractérisation du tempérament chez le poulain après sevrage, 29ème Journée de la
Recherche Equine, Paris, pp. 49–60.
Lansade, L., 2005. Le tempérament du cheval : étude théorique, applications à la sélection des chevaux destinés à
l’équitation. Ph.D. Thesis. University of Tours, France. http://wcentre.tours.inra.fr/prc/internet/resultats/theses/lansade/
these_lea_lansade.pdf.
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Lansade, L., Bouissou, M.F., Erhard, H., 2007. Reactivity to isolation and association with conspecifics: a temperament
trait stable across time and situations. Appl. Anim. Behav. Sci., doi:10.1016/j.applanim.2007.03.003.
Lansade, L., Lévy, F., Bouissou, M.F., 2006. Horse’s temperament and suitability for riding activity can be predicted from
8 months of age. In: 40th International Congress of the International Society for Applied Ethology, Bristol, p. 234.
Marshall, A.E., 1985. Brain stem auditory-evoked response in the nonanesthetized horse and pony. Am. J. Vet. Res. 46,
1445–1450.
Martin, P., Bateson, P., 1993. Measuring Behaviour: An Introductory Guide. Cambridge University Press, Cambridge.
Mills, D.S., 1998. Personality and individual differences in the horse, their significance, use and measurement. Equine
Vet. J. 27 (Suppl.), 10–13.
Momozawa, Y., Kusunosce, R., Kikusui, T., Takeuchi, Y., Mori, Y., 2005. Assessment of equine temperament
questionnaire by comparing factor structure between two separate surveys. Appl. Anim. Behav. Sci. 92, 77–84.
Momozawa, Y., Ono, T., Sato, F., Kikusui, T., Takeuchi, Y., Mori, Y., Kusunosce, R., 2003. Assessment of equine
temperament by a questionnaire survey to caretakers and evaluation of its reliability by simultaneous behavior test.
Appl. Anim. Behav. Sci. 84, 127–138.
Murphy, J.A., 1998. Describing categories of temperament in potential guide dogs for the blind. Appl. Anim. Behav. Sci.
58, 163–178.
Plomin, R., 1983. Developmental behavioral genetics. Child Dev. 54, 253–259.
Redua, M.A., Valadao, C.A.A., Duque, J.C., Balestrero, L.T., 2002. The pre-emptive effect of epidural ketamine on
wound sensitivity in horses tested by using von Frey filaments. Vet. Anaesth. Analg. 29, 200–206.
Saslow, C.A., 1999. Factors affecting stimulus visibility for horses. Appl. Anim. Behav. Sci. 61, 273–284.
Saslow, C.A., 2002. Understanding the perceptual world of horses. Appl. Anim. Behav. Sci. 78, 209–224.
Stelmack, R.M., 1990. Biological bases of extraversion: psychophysiological evidence. J. Pers. 58, 293–311.
Stoddart, D.M., 1980. The Ecology of Vertebrate Olfaction. Chapman & Hall, London, p. 25.
Thomas, A., Chess, S., 1977. Temperament and Development. Brunner Mazel, New York.
Timney, B., Keil, K., 1992. Visual acuity in the horse. Vision Res. 32, 2289–2293.
Tukey, J.W., 1977. Box-and-whisker plots. In: Exploratory Data Analysis, Addison, Wesley, Reading, pp. 39-43.
Verhagen, J.V., Engelen, L., 2006. The neurocognitive bases of human multimodal food perception: sensory integration.
Neurosci. Biobehav. Rev. 30, 613–650.
Visser, E.K., van Reenen, C.G., Hopster, H., Schilder, M.B.H., Knaap, J.H., Barneveld, A., Blokhuis, H.J., 2001.
Quantifying aspects of young horses’ temperament: consistency of behavioural variables. Appl. Anim. Behav. Sci. 74,
241–258.
Wolff, A., Hausberger, M., Le Scolan, N., 1997. Experimental tests to assess emotionality in horses. Behav. Process. 40,
209–221.

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Applied Animal Behaviour Science xxx (2008) xxx–xxx

Sensory sensitivities: Components of a horse’s

2 L. Lansade et al. / Applied Animal Behaviour Science xxx (2008) xxx–xxx

Keywords: Horse; Equus caballus; Temperament; Sensory sensitivity; Behavioural tests

Temperament is generally defined as a set of individual differences in behavioural tendencies,

L. Lansade et al. / Applied Animal Behaviour Science xxx (2008) xxx–xxx 3

2. Animals, materials and methods

2.2. Experimental system and habituation

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2.3. Experimental procedure

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2.3.1. Odour test

2.3.2. Food test

2.3.3. Visual stimulus test

2.3.4. Sound test

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2.3.5. Tactile tests

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2.3.6. Description of the temperament tests

Fig. 3. Novel object and novel area tests experimental system.

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10 L. Lansade et al. / Applied Animal Behaviour Science xxx (2008) xxx–xxx

2.4. Statistical analysis

2.4.1. Description of the animals’ responses according to stimulus intensity

2.4.2. Stability over time

2.4.3. Stability across situations

2.4.4. Correlations between the sensory tests

3.1. Description of the animals’ responses according to stimulus intensity (series A)

3.1.1. Odour test

3.1.2. Food test

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12 L. Lansade et al. / Applied Animal Behaviour Science xxx (2008) xxx–xxx

3.1.3. Visual stimulus test

3.1.4. Sound test

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3.1.5. Tactile tests

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3.2. Stability over time

3.3. Stability across situations

3.4. Correlations between sensory tests

4.1. Description of the animals’ responses according to the stimulus intensity

L. Lansade et al. / Applied Animal Behaviour Science xxx (2008) xxx–xxx 15

16 L. Lansade et al. / Applied Animal Behaviour Science xxx (2008) xxx–xxx

4.2. Are the responses stable over time?

4.3. Are the responses stable across situations—within each sensoriality?

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18 L. Lansade et al. / Applied Animal Behaviour Science xxx (2008) xxx–xxx

4.7. Practical application

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20 L. Lansade et al. / Applied Animal Behaviour Science xxx (2008) xxx–xxx

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