Applied Animal Behaviour Science 125 (2010) 30–37 Contents lists available at ScienceDirect Applied Animal Behaviour Science journal homepage: www.elsevier.com/locate/applanim Horses’ learning performances are under the influence of several temperamental dimensions Léa Lansade a,b,c,d,∗ , Faustine Simon a a b c d INRA (National Institute for Agronomical Research), UMR85 Physiologie de la Reproduction et des Comportements, F-37380 Nouzilly, France CNRS, UMR6175 Physiologie de la Reproduction et des Comportements, F-37380 Nouzilly, France Université François Rabelais de Tours, F-37041Tours, France Haras Nationaux, F-37380 Nouzilly, France a r t i c l e i n f o Article history: Accepted 18 February 2010 Available online 30 March 2010 Keywords: Avoidance task Equus caballus Fearfulness Learning Personality Temperament a b s t r a c t Learning performances are influenced by many factors, not only breed, age and sex, but also temperament. The purpose of this study was to understand how different temperamental dimensions affect the learning performance of horses, Equus caballus. First, we carried out a series of behavioural tests on 36 Welsh ponies aged 5–7 years to measure five temperamental dimensions: fearfulness (novel area test and surprise test), gregariousness (social isolation test), reactivity to humans (passive human test), tactile sensitivity (von Frey filament test) and activity level (evaluation of locomotor activity during all the tests). We then presented them with two learning tasks (avoidance and backwards–forwards tasks). In the avoidance task they had to learn to jump over a fence when they heard a sound associated with an aversive stimulus (puff of air). In the backwards–forwards task they had to walk forwards or move backwards in response to a tactile or vocal command to obtain a food reward. There was no correlation between performances on the two learning tasks, indicating that learning ability is task-dependent. However, correlations were found between temperamental data and learning performance (Spearman correlations). The ponies that performed the avoidance task best were the most fearful and the most active ones. For instance, the number of trials required to perform 5 consecutive correct responses (learning criterion) was correlated with the variables aimed at measuring fearfulness (way of crossing a novel area: rs = −0.41, P = 0.01 and time to start eating again after a surprise effect: rs = −0.33, P = 0.05) and activity level (frequency of trotting during all the tests: rs = −0.40, P = 0.02). The animals that performed the backwards–forwards task best were the ones that were the least fearful and the most sensitive. For instance, the learning criterion (corresponding to the number of trials taken to achieve five consecutive correct responses) was correlated with the variables aimed at measuring fearfulness (latency to put one foot on the area: rs = 0.43, P = 0.01; way of crossing a novel area: rs = 0.31, P = 0.06; and time to start eating again after a surprise effect: rs = 0.43, P = 0.009) and tactile sensitivity (response to von Frey filaments: rs = −0.44, P = 0.008). This study revealed significant links between temperament and learning abilities that are highly task-dependent. © 2010 Elsevier B.V. All rights reserved. 1. Introduction ∗ Corresponding author at: INRA (National Institute for Agronomical Research), Physiologie de la reproduction et des comportements UMR85, 37380 France. Tel.: +33 2 4742 7279; fax: +33 2 4742 7743. E-mail address: Lansade@tours.inra.fr (L. Lansade). 0168-1591/$ – see front matter © 2010 Elsevier B.V. All rights reserved. doi:10.1016/j.applanim.2010.02.010 Learning performance varies widely between individuals. This variability is the result of many factors, such as genetics, age, sex, or the environment (for reviews in horses: Nicol, 2002; Murphy and Arkins, 2007), but can L. Lansade, F. Simon / Applied Animal Behaviour Science 125 (2010) 30–37 also be affected by certain temperamental dimensions. A temperamental dimension can be defined as a characteristic of the individual that appears early in life and that is relatively stable across situations and over the course of time (Goldsmith et al., 1987; Bates, 1989). A large number of studies have focused on the influence of the fearfulness dimension. Most have shown that fearfulness impaired performance (e.g. passive avoidance in Japanese quail chicks, Coturnix japonica: Richard et al., 2000; active avoidance or spatial learning in rats, Rattus norvegicus: Brush et al., 1985; Herrero et al., 2006 and in working dogs, Canis familiaris: Svartberg, 2002). Several studies on horses have also suggested that the most fearful or reactive animals take longer to learn various tasks, including instrumental conditioning tasks (Lindberg et al., 1999; Visser et al., 2003), spatial learning tasks (Heird et al., 1986) or discriminative tasks (Fiske and Potter, 1979). However, not all studies support this conclusion. For instance, in guppies, Poecilia reticulata, Budaev and Zhuikov (1998) have shown that it was the most fearful individuals that performed avoidance tasks best. In rodents, Brinks et al. (2007) have reported that high stress-reactive individuals had an advantage over low stress-reactive individuals when learning certain relational tasks. Finally, Lansade (2005) has shown that fearfulness enhances the performance of horses in an associative task. Findings about the influence of the fearfulness dimension (or a similar dimension such as emotionality) are complex: sometimes it seems to improve learning performance, while at other times it has a deleterious effect on cognitive functions. In fact, the findings depend on the species, learning task, kind of reinforcement and level of fearfulness. This area therefore requires further investigation. From a practical point of view, a better understanding of this mechanism would make it possible to determine the ease with which a horse learns according to the task and its level of fearfulness. In view of the complexity of the relationship between fearfulness and learning performance, we would expect that a fearful horse would perform well on one task but not necessarily on another. In this case, we could personalize the training of each horse according to its level of fearfulness. In addition to fearfulness, studies have identified other temperamental dimensions that affect learning performance. For instance, Svartberg (2002) has shown that a shyness/boldness dimension, related to traits such as playfulness, curiosity, chase proneness and sociability, correlates with high success in working dog trials, with bolder dogs being more successful. In the meadow vole, Microtus pennsylvanicus, behavioural strategies displayed in a radial maze were linked to differences in locomotor activity level (Teskey et al., 1998). However, there are very few papers of this type examining the influence of different temperamental dimensions on learning performance. In this study, we investigated the influence of five temperamental dimensions on learning performance in two different instrumental conditioning situations. These dimensions were fearfulness, activity level, gregariousness, sensory sensitivity and reactivity to humans. These dimensions have been described in many species including horses (for a review, see Gosling and John, 1999). We measured them using previously validated tests that have shown 31 good stability over time and across situations (fearfulness: Lansade et al., 2008a; gregariousness: Lansade et al., 2008b; activity: Lansade et al., 2006; reactivity to humans: Lansade and Bouissou, 2008; sensory sensitivity: Lansade et al., 2008c) and also correlate well with a questionnaire assessing the everyday behaviour of a horse (Lansade et al., 2006). Thus, these tests seem to give a reliable and relatively complete estimation of horse temperament. 2. Animals, materials and methods Experiments reported in this paper were conducted under license from the French Ministry of Agriculture (no. 37-125). 2.1. Animals The study involved 36 female Welsh ponies aged 5–7 years, bred at INRA Nouzilly (France) and accustomed to being handled (regularly haltered and tethered). Before the experiment, the animals lived outside in summer and inside in winter. During the experiment, they were housed in seven adjacent loose boxes (6 m × 4 m) in random groups of three or four and spent 4 h per day together in a large outdoor paddock. They had straw bedding and received concentrated feed (pellets) and hay three times a day. Water was available ad libitum. The ponies were assigned to all the tests in random order. 2.2. Temperament tests Five temperamental dimensions (fearfulness, gregariousness, activity level, reactivity to humans and sensory sensitivity) were assessed using the tests developed by Lansade et al. (2008a,b,c, 2006) and Lansade and Bouissou (2008), respectively. We used the behavioural parameters selected during these studies, as they appear to be reliable indicators of temperament due to their stability over time and across situations. 2.2.1. Experimental system All the tests were carried out in a loose box (2.70 m × 8.10 m—Fig. 1). Two observers were hidden behind a one-way mirror. An audience horse was tied up outside the box, visible to the tested pony, to avoid social isolation interfering with the other measured characteristics. This audience horse was chosen for its quietness. The animals had been habituated to the experimental structure by being placed in this situation for 5 min a day for 5 consecutive days. 2.2.2. Experimental procedure Behavioural tests were carried out in strict order for a total period of approximately 30 min per pony. Each pony followed the same test sequence: Habituation to the test pen: The experimenter took the pony into the loose box and left it alone for 5 min. Passive human test: To characterise reactivity to humans, an experimenter (always the same one) entered the loose box and stayed motionless beside a wall for 3 min (Fig. 1). 32 L. Lansade, F. Simon / Applied Animal Behaviour Science 125 (2010) 30–37 Fig. 1. Arrangement for the temperamental tests. The frequency of sniffing or nibbling the human was recorded. Tactile sensitivity test: The aim of this test was to measure tactile sensitivity. An experimenter held the pony on a lunge line throughout the test. A second experimenter applied a von Frey filament to the base of the pony’s withers (von Frey filaments, Stoelting, IL, USA). These filaments consist of a hard plastic body connected to a nylon thread. The principle of the test is to evaluate the response of the individual to mechanical stimuli using different strengths of filaments. Thus, they are calibrated to exert a specific force on the skin, ranging from 0.008 to 300 g. They were applied perpendicularly on the animal’s skin until the nylon filament started to bend. Trembling of the platisma muscle was recorded. The response was coded in a binary form (trembling/not trembling). There were two phases to the test. In the first, which was carried out after the passive human test, a 0.008-g filament was applied to the right side of the pony, followed by a 300-g filament to the left. In the second, after the novel area test, a 0.02-g filament was applied to the pony’s right side and a 1-g filament to its left. We recorded the number of times the ponies responded to the filaments. The most sensitive horses responded whatever the force, whereas the less sensitive ones only reacted to some. Social isolation test: To characterise gregariousness, we removed the audience horse from sight and sound of the tested pony for 1.5 min. The frequency of neighing was recorded. Novel area test: For this test, which characterises reactivity to novelty, a trait underlying the dimension of fearfulness, we divided the floor of the loose box into three zones of 2.7 m × 2.7 m (Fig. 1). The first zone corresponded to a starting zone (on the right in Fig. 1) and the third zone was an arrival zone (on the left in Fig. 1). The arrival zone contained a bucket of pellets with which ponies were familiar. Just before the test, the ponies underwent a habituation phase during which they learnt to go from the starting zone to the arrival zone containing the bucket. To achieve this, an experimenter led the pony by the halter to the starting zone and released it so that it was free to go to the arrival zone to eat. This was repeated three times. During the test, a pink carpet (2 m × 2.7 m) was placed in the second zone. As in the habituation stage, the experimenter released the pony in the starting zone and recorded the time it took to put one foot on the carpet and the manner in which it crossed (walk, trot, jump, did not cross). If the pony did not cross the area within 180 s, the test was terminated and a time of 181 s was assigned. Surprise test: In this test, which characterises reactivity to suddenness, a trait underlying the dimension of fearfulness, the experimenter opened a black umbrella in front of the animal while it was eating. A bucket of pellets was placed near the entrance (Fig. 1). When the animal had been eating with its head in the bucket for 3 s, the experimenter opened the umbrella and started the stopwatch. The test ended when the pony started eating again. The time taken to start eating again was recorded. If the pony did not start eating within 180 s, the experimenter stopped the test and assigned a time of 181 s. Locomotor activity: In order to measure locomotor activity, we divided the test pen into six areas of equal size (Fig. 1). We recorded the number of areas crossed by the pony and the frequency of trotting during the habituation phase, the passive human test and social isolation test. 2.3. Avoidance task Various studies have investigated active avoidance tasks in many species including horses (Haag et al., 1980; Rubin et al., 1980; McCall et al., 1993). In the present study we used a procedure similar to that used by Visser et al. (2003). The avoidance task involves instrumental learning, using negative reinforcement. 2.3.1. Experimental system The testing area consisted of a loose box (5.40 m × 2.70 m) divided into two equal parts by a 40-cm high wooden crossbar (Fig. 2). The same audience horse as the one used in the temperament tests was tied up outside the box, visible to the tested pony. Observers Fig. 2. Arrangement for the avoidance task. L. Lansade, F. Simon / Applied Animal Behaviour Science 125 (2010) 30–37 Table 1 Levels of stimulation used for the avoidance task. 33 corridor 5 min per day, during the 5 days preceding the learning procedure. Level Kind of stimulation Level 0 Level 1 Levels 2, 3, 4, 5 Ringing of bell (duration 1 s) Ringing of bell and puff of air (duration 1 s) Repetition of level 1 remained unseen behind a one-way mirror. Ponies were equipped with an air-pressure system attached to a soft elastic belt (Master Plus Pro® , Dynavet, Switzerland) that could be operated by remote control to emit a bell ring or a puff of compressed air behind the pony’s shoulder. Ponies equipped with this system entered the testing area 5 min per day, for the 5 days preceding the learning procedure. 2.3.2. Learning procedure Ponies had to cross the bar when they heard the bell to avoid the puff of air. The puff of air is negative reinforcement used to encourage the bar-crossing behaviour. There were six sessions, one every 2 days. The pony was led into the area and stayed free for 1 min before carrying out 10 consecutive trials. For each trial, the observer applied the levels of stimulation described in Table 1 until the pony crossed the bar. There was a 3-s break between each level. Whether the pony crossed the bar (success) or not (failure), another trial began after 1.5 min. We recorded two parameters (Table 3). One pony never jumped the bar at any level of stimulation. 2.4. Backwards–forwards task The second learning task was similar to one used for schooling horses, when the handler teaches the horse to be led with a lunge and to walk forwards or move backwards. 2.4.1. Experimental system Ponies, equipped with a halter and a lunge, were tested in the corridor of the barn where they lived (7 m × 3 m). The same audience horse as before was tethered at one extremity of the corridor. To habituate the ponies to the test situation, the experimenter led them individually along the 2.4.2. Learning procedure Ponies were given a food reward when they walked forwards or moved backwards when the experimenter gave them a tactile (step 1) or vocal command (step 2). Ponies were tested six times, once every 2 days. There were 16 consecutive trials in each session (eight walking forwards and eight moving backwards, randomly organised during the session). First step: The experimenter stood at the left side of the pony, holding it with a lunge. Each trial began with a level1 stimulation (Table 2). If the pony did not respond, the stimulation increased as indicated in Table 2 until it did. The correct response (success) was for the pony to take two steps forwards (walk) or backwards (move back). They were rewarded with a handful of pellets. Whether the pony responded or not another trial began after 20 s. Two parameters were recorded (Table 3). If the pony achieved an 80% success rate (respond at level 1), it completed the second step. Second step: The procedure was exactly the same as during the first step, but we added a “level 0”, consisting of a vocal command only (Table 2). Learning under stress (session 6): The procedure of the ‘learning under stress’ (corresponding to session 6) was similar to the one described above, but with a potentially stressful condition. The testing area was unfamiliar and surrounded by 33 plastic bags full of straw (Fig. 3). A white noise was produced and the animal was socially isolated. The number of successes at level 0 or 1 was recorded during this session (Table 3). 2.5. Statistical analyses We used XLSTAT software (Addinsoft Software, Paris, France) to analyse the data. Kolmogorov–Smirnov tests for normality revealed deviation from normality, and so we used nonparametric statistics. We calculated Spearman correlations between the parameters recorded during the two learning tasks, then between the data recorded during the temperament tests and the learning tasks. The Spear- Table 2 Levels of stimulation used for the backwards–forwards task. Level Kind of stimulation Level 0 (step 2 only) To move back: handler said “back” To walk: handler said “walk” Level 1 To move back: handler turned round and placed his hand on the pony’s breast To walk: handler took two steps and exerted a slight pressure on the lunge Level 2 To move back: handler stretched his arm to push on the breast To walk: handler stretched his arm to pull the lunge Level 3 To move back: handler pushed on the breast using his body weight To walk: handler pulled the lunge using his body weight Level 4 To move back: handler pushed on the breast as hard as possible To walk: handler pulled the lunge as hard as possible Level 5 A second handler entered the area and helped push the breast (to move back) or croup (to walk) 34 L. Lansade, F. Simon / Applied Animal Behaviour Science 125 (2010) 30–37 Table 3 Parameters recorded during the two learning tasks. Success in session 1 Learning criteria (number of trials needed to achieve 5 consecutive correct responses) Number of trials to achieve 5 successesa at level 0 Backwards–forwards Step 1 Number of successesa whatever the level during session 1 Number of successesb at level 1 during session 1, step 1 Backwards–forwards Step 2 Number of successesb at level 0 during session 1, step 2 × Parameter not analysed since this criteria was achieved by less than 15% of animals Backwards–forwards Under Stress Number of successesb at level 0 or 1 during stressful session × Redundant parameter with ‘success session 1’, since there was only one session under the stress condition Active avoidance a b Number of trials to achieve 5 successesb at level 1 Crossing the bar. Taking two steps forwards or backwards. 3. Results 3.1. Correlations between the performances on the two learning tasks No significant correlation appeared between the learning criteria measured in the backwards–forwards task and the avoidance task (P > 0.10). 3.2. Correlations between temperamental data and performances on the avoidance task Fig. 3. Arrangement for the backwards–forwards task performed under stress (session 6). man correlation coefficient (rs value) and the P value are shown. We considered correlations as statistically significant when P < 0.05 and tending to be significant when P < 0.1. ‘Success in session 1’ was significantly correlated with the two locomotor activity parameters: the more active the ponies were, the more frequently they jumped the fence during session 1 (P < 0.01, Table 4), and the more sensitive they were, the more they tended to jump the fence (P = 0.09). The ‘learning criterion’ was significantly correlated with fearfulness and locomotor activity: the more fearful and active the ponies were, the faster they learned (correlations Table 4 Spearman correlations between temperamental data and learning performance in the avoidance task (N = 36). Success in session 1 (number of times ponies crossed the bar at any level) Learning criterion (number of trials to achieve 5 consecutive correct responses) Reactivity to humans Frequency of sniffing and licking NS NS Tactile sensitivity Response to von Frey filaments rs = 0.28 P = 0.09 NS Gregariousness Frequency of neighing NS NS Fearfulness—novel area test Latency to put one foot on the carpet NS NS Fearfulness—novel area test Way of crossing NS rs = −0.41 P = 0.01 Fearfulness—surprise test Time to start eating again NS rs = −0.33 P = 0.05 Locomotor activity Number of areas crossed rs = 0.44 P = 0.008 rs = −0.30 P = 0.08 Locomotor activity Frequency of trotting rs = 0.42 P = 0.01 rs = −0.40 P = 0.02 Significant correlations are presented in bold, tendency are presented in italics. L. Lansade, F. Simon / Applied Animal Behaviour Science 125 (2010) 30–37 35 Table 5 Spearman correlations between temperamental data and learning performance in the backwards–forwards task (N = 36). Success in session 1, step 1 Learning criterion step 1 Success in session 1, step 2 Success under stress Reactivity to human Frequency of sniffing and licking NS NS NS NS Tactile sensitivity Responses to von Frey filaments rs = 0.42 P = 0.01 rs = −0.44 P = 0.008 NS NS Gregariousness Frequency of neighing NS NS NS NS Fearfulness—novel area test Latency to put one foot on the carpet rs = −0.42 P = 0.01 rs = 0.43 P = 0.01 NS NS Fearfulness—novel area test Way of crossing rs = −0.29 P = 0.09 rs = 0.31 P = 0.06 NS rs = −0.34 P = 0.04 Fearfulness—surprise test Time to start eating again rs = −0.48 P = 0.004 rs = 0.43 P = 0.009 rs = −0.34 P = 0.04 rs = −0.47 P = 0.004 Locomotor activity Number of areas crossed NS NS NS NS Locomotor activity Frequency of trotting NS NS NS NS Significant correlations are presented in bold, tendency are presented in italics. with: way of crossing: P < 0.01; time to start eating again: P = 0.05; frequency of trotting: P = 0.02, Table 4). 3.3. Correlations between temperamental data and performance on the backwards–forwards task During step 1, the ‘success in session 1’ and the ‘learning criterion’ were significantly correlated with tactile sensitivity (P = 0.01 and P = 0.008, respectively) and fearfulness (correlations with: latency to put one foot on the carpet: P = 0.01 and P = 0.01; time to start eating again: P = 0.004 and P = 0.009, Table 5). The most successful ponies were the most sensitive and the least fearful. During step 2, ‘success in session 1’ was significantly correlated with the time to start eating again during the surprise test: the most successful animals were the least fearful (P = 0.04). Under stressful conditions, success was significantly correlated with the way of crossing (P = 0.04) and the time to start eating again (P = 0.004): the most successful ponies under the stressful condition were the least fearful. 4. Discussion The aim of this study was to investigate the influence of various temperamental dimensions on the learning performance of horses. First, we showed that performance depended on the tasks, with no significant correlation between the performance criteria in the two learning tasks. This result is in line with most previous studies (horses: Wolff and Hausberger, 1996; Visser et al., 2003; for a review: Nicol, 2002). Secondly, we found correlations between some temperamental dimensions and performance. In sum, the animals that performed the avoidance task best were the most fearful and active, whereas in the backwards–forwards task they were the least fearful and the most sensitive. It is particularly noteworthy that the same dimensions were not necessarily involved in each task, and when they were the same (as for fearfulness), they had opposite effects on learning performance. Thus, certain temperamental profiles can enhance the horse’s performance in one task and not in another. Several hypotheses can be put forward to explain this phenomenon and are described below. First, the positive influence of activity on the avoidance task performance could be due to the fact that the most active animals were more inclined to jump the fence spontaneously, and thus to associate the sound with this response. This is in line with other studies such as the one by Teskey et al. (1998) which found that locomotor activity was an important behavioural component of radial maze acquisition by the meadow vole. In a previous study, Lansade (2005) also found that active horses learn to change pace more rapidly when they hear a vocal command predicting an aversive stimulus. By contrast, this dimension was not involved in the backwards–forwards task, probably because the ponies were tethered on a lunge and not free to express movements spontaneously. The involvement of tactile sensitivity in the performance on the ‘backwards–forwards task—step 1’ was also quite logical. The most sensitive ponies are probably more inclined to respond to slight contact, such as the pressure exerted on their breast. In fact, a previous study carried out on 200 horses has shown that this type of horse is sought after by experienced riders because they responded better to aids such as leg contact (Lansade et al., 2007). By contrast, it is interesting to observe that this dimension did not influence performance on the ‘backwards–forwards test—step 2’, which no longer involved a tactile but a vocal command. In the active avoidance task, this dimension only tended to be correlated with success in the first session. In addition to its tactile feature, the main characteristic of the puff of air was undoubtedly its potentially frightening suddenness. The interaction between these two characteristics may explain the weak influence of tactile sensitivity on performance. 36 L. Lansade, F. Simon / Applied Animal Behaviour Science 125 (2010) 30–37 The fearfulness dimension was involved in both tasks, but with opposite effects. In the backwards–forwards task, the least fearful animals performed best, whatever the step or the environment (familiar or stressful). This result is in line with many studies on various species which reported a negative influence of fearfulness on performance (e.g. Japanese quail: Richard et al., 2000; rats: Herrero et al., 2006; dog: Svartberg, 2002). Other studies also confirm this result with horses. For instance, Fiske and Potter (1979) and Heird et al. (1986) found that the least fearful horses, identified using a subjective score or an umbrella test, were the most successful in a discriminative task. Similarly, Mader and Price (1980) and Lindberg et al. (1999) have shown that horses belonging to breeds believed to be particularly reactive learned more slowly in an operant task or in a visual discriminative task than horses belonging to less reactive breeds. To explain this result, Lindberg argues that horses that are nervous and reactive may be more easily distracted and therefore slower to learn. This is in line with a common hypothesis that the most fearful animals shift attention away from the task, resulting in poor performance (for a review, see Mendl, 1999). This hypothesis could also explain our results. In contrast, in the avoidance task, the most fearful animals performed best. As mentioned above, the stimulus used in this test (the puff of air) was potentially frightening because of its suddenness. We can suppose that the most fearful ponies were more inclined to avoid the negative reinforcement and consequently learned faster. This result is not in line with Visser et al.’s study (2003), which also investigated links between emotionality and performance on an avoidance test. Although they did not find any simple relationship, it seems that unsuccessful horses (horses that do not move when an aversive stimulus is used) had a high level of emotionality. In our study, only one pony was unsuccessful. Thus, it is difficult to compare the results. However, it is interesting to note that our ponies were older than the horses in Visser’s study (aged 5–7 years vs. 1–2 years) and belong to different breeds (Welsh ponies vs. Dutch Warmblood horses). For these reasons, it may be possible that our ponies were less fearful than their horses. Consequently, we can postulate that performance in the avoidance test may improve up to a certain level of fearfulness (conclusion of our experiment with older ponies) and may then decrease (as proposed by Visser with younger and probably more fearful horses). This would fit in with the well-known U-shaped curve first described by Yerkes and Dodson (1908). In the future, it would be interesting to validate this hypothesis and describe this curve precisely, assessing at what level fearfulness becomes deleterious to performance. Thus, while fearfulness appears as a major factor influencing performance, its involvement seems to depend on its intensity and the kind of reinforcement. This study provides some indications about this involvement, but further investigation is required for a more thorough understanding of its influence. Finally, the dimensions of gregariousness and reactivity to humans had no impact on performance in any of the tasks. The fact that the animals were tested in the presence of an audience horse may explain the lack of correlation with gregariousness. By contrast, we could have expected reactivity to humans to be involved in the ‘backwards–forwards task’ due to the human presence. However, the fact that the ponies had been largely habituated to humans before the experiment could have masked a possible effect. 5. Conclusion To conclude, the results clearly indicate that the influence of temperament on learning performance is highly task-dependent. We do not think that temperament directly influences acquisition processes, but rather a predisposition to react to stimuli involved in learning situations. For instance, this predisposition could lead the animals to be more motivated to avoid the reinforcement (such as the puff of air or the pressure exerted on their breast), or to be more inclined to move and jump the bar. This could explain why the temperamental profile involved in performance changed according to the characteristics of the tasks. In the future, it would be of interest to identify categories of task that are influenced by the same temperamental profile, describing precisely the kind of reinforcement, the action to be learned, the environment, etc. From a practical point of view, this would enable horse temperament to be matched with suitable training. By evaluating a horse’s temperament it would be possible to determine the most appropriate types and conditions of learning. For instance, some horses would learn better with negative reinforcement, others with positive reinforcement. In this way, an individual training programme could be established for each animal. 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. 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