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.     

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.     

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.     

The development of ordinal numerical knowledge in infancy

A critical question in cognitive science concerns how numerical knowledge develops. One essential component of an adult concept of number is ordinality: the greater than and less than relationships between numbers. Here it is shown in two experiments that 11-month-old infants successfully discriminated, whereas 9-month-old infants failed to discriminate, sequences of numerosities that descended in numerical value from sequences that increased in numerical value. These results suggest that by 11 months of age infants possess the ability to appreciate the greater than and less than relations between numerical values but that this ability develops between 9 and 11 months of age. In an additional experiment 9-month-old infants succeeded at discriminating the ordinal direction of sequences that varied in the size of a single square rather than in number, suggesting that a capacity for non-numerical ordinal judgments may develop before a capacity for ordinal numerical judgments. These data raise many questions about how infants represent number and what happens between 9 and 11 months to support ordinal numerical judgments.     

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.     

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.     

Cone excitation ratios correlate with color discrimination performance in the horse (Equus caballus)

Six horses (Equus caballus) were trained to discriminate color from grays in a counterbalanced sequence in which lightness cues were irrelevant. Subsequently, the pretrained colors were presented in a different sequence. Two sets of novel colors paired with novel grays were also tested. Performance was just as good in these transfer tests. Once the horse had learned to select the chromatic from the achromatic stimulus, regardless of the specific color, they were immediately able to apply this rule to novel stimuli. In terms of the underlying visual mechanisms, the present study showed for the first time that the spectral sensitivity of horse cone photopigments, measured as cone excitation ratios, was correlated with color discrimination performance, measured as accuracy, repeated errors, and latency of approach.     

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.     

Expectations about numerical events in four lemur species (Eulemur fulvus, Eulemur mongoz, Lemur catta and Varecia rubra)

Although much is known about how some primates—in particular, monkeys and apes—represent and enumerate different numbers of objects, very little is known about the numerical abilities of prosimian primates. Here, we explore how four lemur species (Eulemur fulvus, E. mongoz, Lemur catta, and Varecia rubra) represent small numbers of objects. Specifically, we presented lemurs with three expectancy violation looking time experiments aimed at exploring their expectations about a simple 1+1 addition event. In these experiments, we presented subjects with displays in which two lemons were sequentially added behind an occluder and then measured subjects duration of looking to expected and unexpected outcomes. In experiment 1, subjects looked reliably longer at an unexpected outcome of only one object than at an expected outcome of two objects. Similarly, subjects in experiment 2 looked reliably longer at an unexpected outcome of three objects than at an expected outcome of two objects. In experiment 3, subjects looked reliably longer at an unexpected outcome of one object twice the size of the original than at an expected outcome of two objects of the original size. These results suggest that some prosimian primates understand the outcome of simple arithmetic operations. These results are discussed in light of similar findings in human infants and other adult primates.This revised version was published online in March 2005 with corrections to Fig. 6.     

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     

Salamanders (<Emphasis Type="Italic">Plethodon cinereus</Emphasis>) go for more: rudiments of number in an amphibian

Techniques traditionally used in developmental research with infants have been widely used with nonhuman primates in the investigation of comparative cognitive abilities. Recently, researchers have shown that human infants and monkeys select the larger of two numerosities in a spontaneous forced-choice discrimination task. Here we adopt the same method to assess in a series of experiments spontaneous choice of the larger of two numerosities in a species of amphibian, red-backed salamanders (Plethodon cinereus). The findings indicate that salamanders "go for more," just like human babies and monkeys. This rudimentary capacity is a type of numerical discrimination that is spontaneously present in this amphibian. Electronic Publication     

Spontaneous number discrimination of multi-format auditory stimuli in cotton-top tamarins (Saguinus oedipus)

Studies using operant training have demonstrated that laboratory animals can discriminate the number of objects or events based on either auditory or visual stimuli, as well as the integration of both auditory and visual modalities. To date, studies of spontaneous number discrimination in untrained animals have been restricted to the visual modality, leaving open the question of whether such capacities generalize to other modalities such as audition. To explore the capacity to spontaneously discriminate number based on auditory stimuli, and to assess the abstractness of the representation underlying this capacity, a habituation-discrimination procedure involving speech and pure tones was used with a colony of cotton-top tamarins. In the habituation phase, we presented subjects with either two- or three-speech syllable sequences that varied with respect to overall duration, inter-syllable duration, and pitch. In the test phase, we presented subjects with a counterbalanced order of either two- or three-tone sequences that also varied with respect to overall duration, inter-syllable duration, and pitch. The proportion of looking responses to test stimuli differing in number was significantly greater than to test stimuli consisting of the same number. Combined with earlier work, these results show that at least one non-human primate species can spontaneously discriminate number in both the visual and auditory domain, indicating that this capacity is not tied to a particular modality, and within a modality, can accommodate differences in format.     

Exaggerated leftward bias in the mental number line of patients with schizophrenia

Several visuo-motor tasks can be used to demonstrate biases towards left hemispace in schizophrenic patients, suggesting a minor right hemineglect. Recent studies in neglect patients used a new number bisection task to highlight a lateralized defect in their visuo-spatial representation of numbers. To test a possible lateralized representational deficit in schizophrenia, we used the number bisection task in 11 schizophrenic patients compared to 11 healthy controls. Participants were required to orally indicate the central number of an interval orally presented. Whereas healthy subjects showed no significant bias, schizophrenic patients presented a significant leftward bias. Therefore, these results suggest an impairment in higher order representations of the number space in patients with schizophrenia, an impairment that is qualitatively similar to the deficit described in neglect patients.     

Mangabeys (<Emphasis Type="Italic">Cercocebus torquatus lunulatus</Emphasis>) solve the reverse contingency task without a modified procedure

Problem solving often relies on generating new responses while inhibiting others, particularly prepotent ones. A paradigm to study inhibitory abilities is the reverse contingency task (Boysen and Berntson in J Exp Psychol Anim Behav Process 21:82–86, 1995), in which two different quantities of food are offered to an individual who receives the array he did not choose. Therefore, mastery of the task demands selecting the smaller quantity to obtain the larger one. Several non-human primates have been tested in the reverse contingency task. To date, only great apes and rhesus monkeys (Macaca mulatta) have succeeded in the original task, with no need of procedural modifications as the large-or-none contingency, correction trials or symbolic stimuli substituting for actual food quantities. Here, four mangabeys were presented with two stimulus arrays of one and four raisins in the context of the reverse contingency task. Three of them learned to perform the task well above chance without a modified procedure. They also reached above-chance performance when presented with two stimulus arrays of zero and four raisins, despite the initial difficulty of choosing a null quantity. After a period of 7–10 months, in which the animals were not tested on any task, all three subjects continued to perform well, even when presented with novel quantity pairs.     

The role of numerical and non-numerical cues in nonsymbolic number processing: Evidence from the line bisection task

In line bisection tasks, adults and children bisect towards the numerically larger of two nonsymbolic numerosities [de Hevia, M. D., & Spelke, E. S. (2009 de Hevia, M. D., & Spelke, E. S. (2009). Spontaneous mapping of number and space in adults and young children. Cognition, 110, 198–207. doi:10.1016/j.cognition.2008.11.003[Crossref], [PubMed], [Web of Science ®] , [Google Scholar]). Spontaneous mapping of number and space in adults and young children. Cognition, 110, 198–207. doi:10.1016/j.cognition.2008.11.003]. However, it is not clear whether this effect is driven by number itself or rather by visual cues such as subtended area [Gebuis, T., & Gevers, W. (2011 Gebuis, T., & Gevers, W. (2011). Numerosities and space: Indeed a cognitive illusion! A reply to de Hevia and Spelke (2009). Cognition, 121, 248–252. doi:10.1016/j.cognition.2010.09.008[Crossref], [PubMed], [Web of Science ®] , [Google Scholar]). Numbers and space: Indeed a cognitive illusion! A reply to de Hevia and Spelke (2009 de Hevia, M. D., & Spelke, E. S. (2009). Spontaneous mapping of number and space in adults and young children. Cognition, 110, 198–207. doi:10.1016/j.cognition.2008.11.003[Crossref], [PubMed], [Web of Science ®] , [Google Scholar]). Cognition, 121, 248–252. doi:10.1016/j.cognition.2010.09.008]. Furthermore, this effect has only been demonstrated with flanking displays of two and nine items. Here, we report three studies that examined whether this “spatial bias” effect occurs across a range of absolute and ratio numerosity differences; in particular, we examined whether the bias would occur when both flankers were outside the subitizing range. Additionally, we manipulated the subtended area of the stimulus and the aggregate surface area to assess the influence of visual cues. We found that the spatial bias effect occurred for a range of flanking numerosities and for ratios of 3:5 and 5:6 when subtended area was not controlled (Experiment 1). However, when subtended area and aggregate surface area were held constant, the biasing effect was reversed such that participants bisected towards the flanker with fewer items (Experiment 2). Moreover, when flankers were identical, participants bisected towards the flanker with larger subtended area or larger aggregate surface area (Experiments 2 and 3). On the basis of these studies, we conclude that the spatial bias effect for nonsymbolic numerosities is primarily driven by visual cues.     

Disentangling the Mechanisms of Symbolic Number Processing in Adults’ Mathematics and Arithmetic Achievement

A growing body of research has shown that symbolic number processing relates to individual differences in mathematics. However, it remains unclear which mechanisms of symbolic number processing are crucial—accessing underlying magnitude representation of symbols (i.e., symbol‐magnitude associations), processing relative order of symbols (i.e., symbol‐symbol associations), or processing of symbols per se. To address this question, in this study adult participants performed a dots‐number word matching task—thought to be a measure of symbol‐magnitude associations (numerical magnitude processing)—a numeral‐ordering task that focuses on symbol‐symbol associations (numerical order processing), and a digit‐number word matching task targeting symbolic processing per se. Results showed that both numerical magnitude and order processing were uniquely related to arithmetic achievement, beyond the effects of domain‐general factors (intellectual ability, working memory, inhibitory control, and non‐numerical ordering). Importantly, results were different when a general measure of mathematics achievement was considered. Those mechanisms of symbolic number processing did not contribute to math achievement. Furthermore, a path analysis revealed that numerical magnitude and order processing might draw on a common mechanism. Each process explained a portion of the relation of the other with arithmetic (but not with a general measure of math achievement). These findings are consistent with the notion that adults’ arithmetic skills build upon symbol‐magnitude associations, and they highlight the effects that different math measures have in the study of numerical cognition.     

On the language specificity of basic number processing: transcoding in a language with inversion and its relation to working memory capacity

Transcoding Arabic numbers from and into verbal number words is one of the most basic number processing tasks commonly used to index the verbal representation of numbers. The inversion property, which is an important feature of some number word systems (e.g., German einundzwanzig [one and twenty]), might represent a major difficulty in transcoding and a challenge to current transcoding models. The mastery of inversion, and of transcoding in general, might be related to nonnumerical factors such as working memory resources given that different elements and their sequence need to be memorized and manipulated. In this study, transcoding skills and different working memory components in Austrian (German-speaking) 7-year-olds were assessed. We observed that inversion poses a major problem in transcoding for German-speaking children. In addition, different components of working memory skills were differentially correlated with particular transcoding error types. We discuss how current transcoding models could account for these results and how they might need to be adapted to accommodate inversion properties and their relation to different working memory components.