Visual laterality in the domestic horse (Equus caballus) interacting with humans

Most horses have a side on which they are easier to handle and a direction they favour when working on a circle, and recent studies have suggested a correlation between emotion and visual laterality when horses observe inanimate objects. As such lateralisation could provide important clues regarding the horse’s cognitive processes, we investigated whether horses also show laterality in association with people. We gave horses the choice of entering a chute to left or right, with and without the passive, non-interactive presence of a person unknown to them. The left eye was preferred for scanning under both conditions, but significantly more so when a person was present. Traditionally, riders handle horses only from the left, so we repeated the experiment with horses specifically trained on both sides. Again, there was a consistent preference for left eye scanning in the presence of a person, whether known to the horses or not. We also examined horses interacting with a person, using both traditionally and bilaterally trained horses. Both groups showed left eye preference for viewing the person, regardless of training and test procedure. For those horses tested under both passive and interactive conditions, the left eye was preferred significantly more during interaction. We suggest that most horses prefer to use their left eye for assessment and evaluation, and that there is an emotional aspect to the choice which may be positive or negative, depending on the circumstances. We believe these results have important practical implications and that emotional laterality should be taken into account in training methods.     

Cue use by foals (Equus caballus) in a discrimination learning task

Discrimination learning studies suggest that horses learn more easily using spatial than visible object-specific (OS) cues. However, spatial cues have generally confounded intra-array, distal and/or egocentric spatial information. It is also unclear whether conflicting cues compete for association or are redundantly encoded, and furthermore, the influence of prior experiences or training has not been quantified so far. We examined the effect of cue modality on unweaned foals’ performance in a discrimination learning task. After a pilot study confirmed that horses could perform the required OS cue discrimination, nine foals learnt to find food in one of three covered buckets, in any of four positions within a test arena. In Stage 1 the rewarded bucket was signified both by OS cues (pattern) and by relative spatial cues (position). On reaching criterion, cues were separated and foals had to ignore the inappropriate cue (Stage 2). Foals assigned to follow spatial cues (n = 5) completed Stage 2 faster than foals for whom OS cues remained consistent (n = 4). Spatial group foals all reached criterion without delay; no foal in the OS group reached criterion within the testing period. OS group foals initially persisted in responding to the previously correct position, adopting spatially-based strategies when this proved unsuccessful. The findings show for the first time that, even in the absence of distal spatial information, intra-array spatial cues were more salient than OS cues for foals in a food-finding task and learning appeared rather inflexible.     

Shetland ponies (Equus caballus) show quantity discrimination in a matching-to-sample design

Numerical competence is one of the aspects of animal cognition with a long history of research interest, but few results are available for the horse. In the present study, we investigated the ability of three Shetland ponies to discriminate between different quantities of geometric symbols presented on a computer screen in a matching-to-sample arrangement. In Experiment 1, the ponies had to relate two similar quantities to another, paired in contrasts (1 vs. 2, 3 vs. 4 and 4 vs. 5) of the same stimulus (dot). Specific pairs of quantities (all differing by one) of up to five different geometrical symbols were displayed in Experiment 2. In each session, both quantities (more and less) were used as sample in such a way that each of the two quantities presented in one test served as positive and as negative stimulus, respectively. The three Shetland ponies were able to discriminate between the given quantities of dots by showing more than 80 % correct responses in two consecutive sessions. Only one of the ponies distinguished different shapes of geometric symbols at a level of 4 versus 5 items. The results show that all ponies were capable of visual quantity discrimination in the present matching-to-sample design, but task solving seemed more difficult when quantities were composed of heterogeneous stimuli. The present results confirm our hypothesis that the ponies based their decision on the matching concept of sameness and were not biased by a spontaneous preference for higher quantities.     

Lateralized suckling in domestic horses (Equus caballus)

Brain lateralization enables preferential processing of certain stimuli and more effective utilization of these stimuli in either the left or the right cerebral hemisphere. Horses show both motor and sensory lateralization patterns. Our aim was to determine whether a lateralized response could be detected in foals during the naturally side-biased behaviour, suckling. The foals’ preferred suckling side could be the effect of either visual or motor lateralization. In the case of a visual lateralized response, foals are expected to suck more often from the mother’s right side, so potential danger can be detected by the better adapted right hemisphere (i.e. left eye). Motor lateralization can be identified when a foal will suck predominantly from one side, either left or right. We found no population trend in the preferred suckling side, but we detected significant differences amongst individual foals. One-third (35.4 %) of 79 foals showed a strong, either right or left side preference which increased with age. The mothers did not influence the foals’ suckling side preferences either by side-biased rejection or termination of suckling. According to our findings, a general pattern of sucking with the left eye open for better danger detection and recognition is unlikely in foals up to 7 months old. Foals of this age are probably young or fully focused on suckling and rely on their mothers’ vigilance. Individual side preferences amongst foals are suggested to be based on motor lateralization.     

Stereotypic horses (Equus caballus) are not cognitively impaired

Animal Cognition - Stereotypies in animals are thought to arise from an interaction between genetic predisposition and sub-optimal housing conditions. In domestic horses, a well-studied...     

Comprehension of human pointing gestures in horses (Equus caballus)

Twenty domestic horses (Equus caballus) were tested for their ability to rely on different human gesticular cues in a two-way object choice task. An experimenter hid food under one of two bowls and after baiting, indicated the location of the food to the subjects by using one of four different cues. Horses could locate the hidden reward on the basis of the distal dynamic-sustained, proximal momentary and proximal dynamic-sustained pointing gestures but failed to perform above chance level when the experimenter performed a distal momentary pointing gesture. The results revealed that horses could rely spontaneously on those cues that could have a stimulus or local enhancement effect, but the possible comprehension of the distal momentary pointing remained unclear. The results are discussed with reference to the involvement of various factors such as predisposition to read human visual cues, the effect of domestication and extensive social experience and the nature of the gesture used by the experimenter in comparative investigations.     

Socially dependent auditory laterality in domestic horses (Equus caballus)

Laterality is now known to be an ubiquitous phenomenon among the vertebrates. Particularly, laterality of auditory processing has been demonstrated in a variety of species, especially songbirds and primates. Such a hemispheric specialization has been shown to depend on factors such as sound structure, species specificity and types of stimuli. Much less is known on the possible influence of social familiarity although a few studies suggest such an influence. Here we tested the influence of the degree of familiarity on the laterality of the auditory response in the domestic horse. This species is known for its social system and shows visible reactions to sounds, with one or two ears moving towards a sound source. By comparing such responses to the playback of different conspecific whinnies (group member, neighbor and stranger), we could demonstrate a clear left hemisphere (LH) preference for familiar neighbor calls while no preference was found for group member and stranger calls. Yet, we found an opposite pattern of ear side preference for neighbor versus stranger calls. These results are, to our knowledge, the first to demonstrate auditory laterality in an ungulate species. They open further lines of thought on the influence of the social “value” of calls and the listener’s arousal on auditory processing and laterality.     

Horses (Equus caballus) discriminate body odour cues from conspecifics

Knowledge about social recognition and memory in animals can help us to determine appropriate management and husbandry techniques. In this study, we used a habituation–discrimination procedure to investigate the ability of horses (Equus caballus) to distinguish between the body odour samples of unfamiliar conspecifics. To pick up body odour, we rubbed material on the coat of horses and presented these unknown body odours to 16 different conspecifics of the same sex and similar age. The test consisted of two successive two-min presentations of a sample from one individual (e.g. individual ‘A’) and a simultaneous presentation of samples from individual ‘A’ and a novel individual (e.g. individual ‘B’) during a final third presentation. The results showed that horses, regardless of sex, decreased the time they spent investigating conspecific body odour across the initial two presentations—demonstrating habituation. In the final presentation, the results demonstrated successful discrimination of the previously experienced odour because horses investigated the novel olfactory sample (‘B’) significantly more than the pre-exposed sample (‘A’). Taken together, these findings suggest, for the first time, that horses are able to discriminate two stimuli derived from body odours of unfamiliar conspecifics over short period of time.     

Long-term memory for categories and concepts in horses (Equus caballus)

Three horses (Equus caballus) with a history of performing cognitive tasks including discrimination learning, categorization, and concept use were tested to evaluate their long-term memory (LTM) in three experiments. In addition, use of LCD multi-displays for stimulus presentation was incorporated into cognition testing protocol for the first time with horses. Experiment 1 tested LTM for discrimination learning that originally occurred 6 years earlier. Five sets of stimuli were used and the two horses tested showed no decrement in performance on four of the sets; however, both horses did score below chance on one set. Experiment 2 examined long-term categorization recall 10 years after horses had demonstrated the ability to make stimulus selections based on shared characteristics within a given category. The horse tested for LTM after the decade-long interval immediately and consistently applied the previously learned categorization rule to not only familiar but also novel sets of stimuli. Experiment 3 tested another horse for LTM for a relative size concept. This horse had originally demonstrated concept rule use in order to select stimuli based on their relative size to one another. More than 7 years later and without further training, this horse reliably applied the previously established size concept to both familiar and novel sets of stimuli. These findings are the first reports of long-term categorical and conceptual memory in horses and are consistent with observations of domestic and wild horses, which indicate that behavioral and ecological events may be remembered for long periods of time. These studies also demonstrate the adaptive nature of horses with regard to their ability to generalize over several different testing conditions.     

Cross-modal recognition of human individuals in domestic horses (Equus caballus)

This study has shown that domestic horses are capable of cross-modal recognition of familiar humans. It was demonstrated that horses are able to discriminate between the voices of a familiar and an unfamiliar human without seeing or smelling them at the same moment. Conversely, they were able to discriminate the same persons when only exposed to their visual and olfactory cues, without being stimulated by their voices. A cross-modal expectancy violation setup was employed; subjects were exposed both to trials with incongruent auditory and visual/olfactory identity cues and trials with congruent cues. It was found that subjects responded more quickly, longer and more often in incongruent trials, exhibiting heightened interest in unmatched cues of identity. This suggests that the equine brain is able to integrate multisensory identity cues from a familiar human into a person representation that allows the brain, when deprived of one or two senses, to maintain recognition of this person.     

Horse (Equus caballus) whinnies: a source of social information

Many animal species that rely mainly on calls to communicate produce individual acoustic structures, but we wondered whether individuals of species better known as visual communicants, with small vocal repertoires, would also exhibit individual distinctiveness in calls. Moreover, theoretical advances concerning the evolution of social intelligence are usually based on primate species data, but relatively little is known about the social cognitive capacities of non-primate mammals. However, some non-primate species demonstrate auditory recognition of social categories and possess mental representation of their social network. Horses (Equus caballus) form stable social networks and although they display a large range of visual signals, they also use long-distance whinny calls to maintain contact. Here, we investigated the potential existence of individual acoustic signatures in whinny calls and the ability of horses to discriminate by ear individuals varying in their degree of familiarity. Our analysis of the acoustic structure of whinnies of 30 adult domestic horses (ten stallions, ten geldings, ten mares) revealed that some of the frequency and temporal parameters carried reliable information about the caller’s sex, body size and identity. However, no correlations with age were found. Playback experiments evaluated the behavioural significance of this variability. Twelve horses heard either control white noise or whinnies emitted by group members, familiar neighbours or unfamiliar horses. While control sounds did not induce any particular response, horses discriminated the social category of the callers and reacted with a sound-specific behaviour (vigilance and attraction varied with familiarity). Our results support the existence of social knowledge in horses and suggest a process of vocal coding/decoding of information.     

Horses (Equus caballus) select the greater of two quantities in small numerical contrasts

The ability to select the greater numerosity over another in small sets seems to stem from the calculation of which set contains more, and has been taken as evidence of a primordial representation at the roots of the primate numerical system. We tested 56 horses (Equus caballus) in a paradigm previously used with human infants and nonhuman primates. Horses saw two quantities paired in contrasts—2 versus 1, 3 versus 2, 6 versus 4 and a control for volume, 2 versus 1 big—and had to make a choice by snout touching the container holding the numerosity selected. The horses spontaneously selected the greater of the two quantities when the numerosities were small. These results add to evidence showing spontaneous quantity assessment in a variety of species. To my brother Marcelo Uller (1964–2008), for whom the understanding of animals, through biomimetics, was the only way to understand man-made engineering.     

Getting rid of blinkers: the case of mirror self-recognition in horses (Equus caballus)

Animal Cognition - The commentary by Gallup and Anderson (Anim Cogn https://doi.org/10.1007/s10071-021-01538-9 , 2021) on the original article by Baragli, Scopa, Maglieri, and Palagi (Anim Cogn...     

Reinforcement as a mediator of the perception of humans by horses (Equus caballus)

A central question in the interspecific human/animal relationship is how domestic animals perceive humans as a significant element of their environment. In this study, we tested the hypothesis that the use of positive or negative reinforcement in horse training may have consequences on the animals’ perception of humans, as a positive, negative or neutral element. Two groups of ponies were trained to walk backwards in response to a vocal order using either positive or negative reinforcement. Heart rate monitors and behavioural observations were used to assess the animals’ perception of humans on the short (just after training) and long (5 months later) terms. The results showed that the type of reinforcement had a major effect on the subsequent animals’ perception of familiar and unfamiliar humans. Negative reinforcement was rapidly associated with an increased emotional state, as revealed by heart rate measurements and behavioural observations (head movements and ears laid back position). Its use led the ponies to seek less contact with humans. On the contrary, ponies trained with positive reinforcement showed an increased interest in humans and sought contact after training. This is especially remarkable as it was reached in a maximum of 5 sessions of 1 to 3 min (i.e. 5 to 15 min) and had lasting effects (visible after 5 months). Even learning was positively influenced by positive reinforcement. Overall, horses seem capable of associating humans to particular experiences and display extended long-term memory abilities.     

Attributing attention: the use of human-given cues by domestic horses (Equus caballus)

Recent research has shown that domestic dogs are particularly good at determining the focus of human attention, often outperforming chimpanzees and hand-reared wolves. It has been suggested that the close evolutionary relationship between humans and dogs has led to the development of this ability; however, very few other domestic species have been studied. We tested the ability of 36 domestic horses to discriminate between an attentive and inattentive person in determining whom to approach for food. The cues provided were body orientation, head orientation or whether the experimenters’ eyes were open or closed. A fourth, mixed condition was included where the attentive person stood with their body facing away from the subjects but their head turned towards the subject while the inattentive person stood with their body facing the subject but their head turned away. Horses chose the attentive person significantly more often using the body cue, head cue, and eye cue but not the mixed cue. This result suggests that domestic horses are highly sensitive to human attentional cues, including gaze. The possible role of evolutionary and environmental factors in the development of this ability is discussed.     

Use of human-given cues by domestic dogs (Canis familiaris) and horses (Equus caballus)

Sixteen domestic dogs (Canis familiaris) and four horses (Equus caballus) were tested for their ability to use human-given manual and facial cues in an object-choice task. Two of the four horses used touch as a cue and one horse successfully used pointing. The performance of the dogs was considerably better, with 12 subjects able to use pointing as a cue, 4 able to use head orientation and 2 able to use eye gaze alone. Group analysis showed that the dogs performed significantly better in all experimental conditions than during control trials. Dogs were able to use pointing cues even when the cuer’s body was closer to the incorrect object. Working gundogs with specialised training used pointing more successfully than pet dogs and gundog breeds performed better than non-gundog breeds. The results of this experiment suggest that animals’ use of human given communicative signals depends on cognitive ability, the evolutionary consequences of domestication and enculturation by humans within the individual’s lifetime. Received: 15 July 1999 / Accepted after revision: 10 January 2000     

Color vision in horses (Equus caballus): deficiencies identified using a pseudoisochromatic plate test

In the past, equine color vision was tested with stimuli composed either of painted cards or photographic slides or through physiological testing using electroretinogram flicker photometry. Some studies produced similar results, but others did not, demonstrating that there was not yet a definitive answer regarding color vision in horses (Equus caballus). In this study, a pseudoisochromatic plate test--which is highly effective in testing color vision both in small children and in adult humans--was used for the first time on a nonhuman animal. Stimuli consisted of different colored dotted circles set against backgrounds of varying dots. The coloration of the circles corresponded to the visual capabilities of different types of color deficiencies (anomalous trichromacy and dichromacy). Four horses were tested on a 2-choice discrimination task. All horses successfully reached criterion for gray circles and demonstration circles. None of the horses were able to discriminate the protan-deutan plate or the individual protan or deutan plates. However, all were able to discriminate the tritan plate. The results suggest that horses are dichromats with color vision capabilities similar to those of humans with red-green color deficiencies.     

Interplay between environmental and genetic factors in temperament/personality traits in horses (Equus caballus)

The aim of the present study was to broach the question of the relative influence of different genetic and environmental factors on different temperament/personality traits of horses (Equus caballus). The researchers submitted 702 horses to standardized experimental tests and investigated 9 factors, either genetic or environmental. Genetic factors, such as sire or breed, seemed to influence more neophobic reactions, whereas environmental factors, such as the type of work, seemed to play a more dominant role in reactions to social separation or learning abilities. Additive effects were evident, showing how environmental factors may modulate behavioral traits. This study constitutes a first step toward understanding the relative weights of genetic factors and how the environment may intervene in determining individual behavioral characteristics.     

Dynamic averaging and foraging decisions in horses (Equus callabus)

The variability of most environments taxes foraging decisions by increasing the uncertainty of the information available. One solution to the problem is to use dynamic averaging, as do some granivores and carnivores. Arguably, the same strategy could be useful for grazing herbivores, even though their food renews and is more homogeneously distributed. Horses (Equus callabus) were given choices between variable patches after short or long delays. When patch information was current, horses returned to the patch that was recently best, whereas those without current information matched choices to the long-term average values of the patches. These results demonstrate that a grazing species uses dynamic averaging and indicate that, like granivores and carnivores, they can use temporal weighting to optimize foraging decisions.     

Phencyclidine discrimination in the pigeon using color tracking under second-order schedules

Pigeons were trained to track different key colors, depending on whether they had been injected with phencyclidine or saline prior to the session. A second-order schedule was used to generate large numbers of responses prior to the initial food delivery. The procedure offers several advantages over traditional procedures for studying drug discrimination.