Are domesticated animals dumber than their wild relatives? A comprehensive review on the domestication effects on animal cognitive performance.

Animal domestication leads to diverse behavioral, physiological, and neurocognitive changes in domesticated species compared to their wild relatives. However, the widely held belief that domesticated species are inherently less "intelligent" (i.e., have lower cognitive performance) than their wild counterparts requires further investigation. To investigate potential cognitive disparities, we undertook a thorough review of 88 studies comparing the cognitive performance of domesticated and wild animals. Approximately 30% of these studies showed superior cognitive abilities in wild animals, while another 30% highlighted superior cognitive abilities in domesticated animals. The remaining 40% of studies found similar cognitive performance between the two groups. Therefore, the question regarding the presumed intelligence of wild animals and the diminished cognitive ability of domesticated animals remains unresolved. We discuss important factors/limitations for interpreting past and future research, including environmental influences, diverse objectives of domestication (such as breed development), developmental windows, and methodological issues impacting cognitive comparisons. Rather than perceiving these limitations as constraints, future researchers should embrace them as opportunities to expand our understanding of the complex relationship between domestication and animal cognition.

Horses discriminate between human facial and vocal expressions of sadness and joy.

Communication of emotions plays a key role in intraspecific social interactions and likely in interspecific interactions. Several studies have shown that animals perceive human joy and anger, but few studies have examined other human emotions, such as sadness. In this study, we conducted a cross-modal experiment, in which we showed 28 horses two soundless videos simultaneously, one showing a sad, and one a joyful human face. These were accompanied by either a sad or joyful voice. The number of horses whose first look to the video that was incongruent with the voice was longer than their first look to the congruent video was higher than chance, suggesting that horses could form cross-modal representations of human joy and sadness. Moreover, horses were more attentive to the videos of joy and looked at them for longer, more frequently, and more rapidly than the videos of sadness. Their heart rates tended to increase when they heard joy and to decrease when they heard sadness. These results show that horses are able to discriminate facial and vocal expressions of joy and sadness and may form cross-modal representations of these emotions; they also are more attracted to joyful faces than to sad faces and seem to be more aroused by a joyful voice than a sad voice. Further studies are needed to better understand how horses perceive the range of human emotions, and we propose that future experiments include neutral stimuli as well as emotions with different arousal levels but a same valence.

Environmental enrichment improves cognitive flexibility in rainbow trout in a visual discrimination task: first insights.

Research on fish cognition provides strong evidence that fish are endowed with high level cognitive skills. However, most studies on cognitive flexibility and generalization abilities, two key adaptive traits for captive animals, focused on model species, and farmed fish received too little attention. Environmental enrichment was shown to improve learning abilities in various fish species, but its influence on cognitive flexibility and generalization abilities is still unknown. We studied farmed rainbow trout (Oncorhynchus mykiss) as an aquaculture model to study how environmental enrichment impacts their cognitive abilities. Using an operant conditioning device, allowing the expression of a motivated choice, we measured fish cognitive flexibility with serial reversal learning tests, after a successful acquisition phase based on two colors discrimination (2-alternative forced choice, 2-AFC), and their ability to generalize a rewarded color to any shape. Eight fish were divided into two groups: Condition E (fish reared from fry stages under enriched conditions with plants, rocks and pipes for ~9 months); Condition B (standard barren conditions). Only one fish (condition E) failed in the habituation phase of the device and one fish (condition B) failed in the 2-AFC task. We showed that after a successful acquisition phase in which the fish correctly discriminated two colors, they all succeeded in four reversal learnings, supporting evidence for cognitive flexibility in rainbow trout. They were all successful in the generalization task. Interestingly, fish reared in an enriched environment performed better in the acquisition phase and in the reversal learning (as evidenced by fewer trials needed to reach the learning criterion), but not in the generalization task. We assume that color-based generalization may be a simpler cognitive process than discriminative learning and cognitive flexibility, and does not seem to be influenced by environmental conditions. Given the small number of individuals tested, our results may be considered as first insights into cognitive flexibility in farmed fish using an operant conditioning device, but they pave the way for future studies. We conclude that farming conditions should take into account the cognitive abilities of fish, in particular their cognitive flexibility, by allowing them to live in an enriched environment.

First report of reindeer (Rangifer tarandus tarandus) response to human-given cues.

Many argue that the animal understanding of human referential communication is a by-product of domestication. However, the domestication hypothesis is not unanimously supported as some nondomesticated species such as sea lions, dolphins, or African elephants perform well in the understanding of human pointing gesture. There is a need to study species with different levels of domestication across different taxa to understand the emerging communicative sociocognitive skills in animals that provide them with the ability to comprehend human-given cues. We conducted a pilot study to assess the performance of eight sledging reindeer following a commonly used human-given cue (a pointing gesture associated with gaze at the target and local enhancement) in a two-way choice task. Domestic reindeer are considered semicaptive, because of their husbandry system in free-ranging conditions, with limited control of their reproduction. We observed that the willingness of the reindeer to participate in the test was age-related, with the younger individuals which lack experience with humans being reluctant to approach the experimenters. Within the more experienced working sledging reindeer, two individuals showed excellent skills and followed the human-given cues 9 out of 10 times. Reindeer show great potential in following a human indication to locate hidden food with minimal training when properly tamed. The effect of previous experience with humans requires further investigation. This is the first demonstration in cervids of an ability to make use of experimenter-given cues in an object-choice task. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

What drives horse success at following human-given cues? An investigation of handler familiarity and living conditions.

Cues such as the human pointing gesture, gaze or proximity to an object are widely used in behavioural studies to evaluate animals' abilities to follow human-given cues. Many domestic mammals, such as horses, can follow human cues; however, factors influencing their responses are still unclear. We assessed the performance of 57 horses at a two-way choice task testing their ability to follow cues of either a familiar (N = 28) or an unfamiliar informant (N = 29). We investigated the effects of the length of the relationship between the horse and a familiar person (main caregiver), their social environment (living alone, in dyads, or in groups) and their physical environment (living in stalls/paddocks, alternating between paddocks and pastures, or living full time in pastures). We also controlled for the effects of horses' age and sex. Our results showed that horses' success rate at the task was not affected by the familiarity of the informant and did not improve with the relationship length with the familiar informant but did increase with the age of the horses. Horses living in groups had better success than the ones kept either in dyads or alone. Finally, horses housed in small paddocks had lower success than those living on pasture. These results indicate that with age, horses get better at following human-given indications regardless of who the human informant is and that an appropriate living and social environment could contribute to the development of socio-cognitive skills towards humans. Therefore, such aspects should be considered in studies evaluating animal behaviour.

Horses cross-modally recognize women and men.

Several studies have shown that horses have the ability to cross-modally recognize humans by associating their voice with their physical appearance. However, it remains unclear whether horses are able to differentiate humans according to different criteria, such as the fact that they are women or men. Horses might recognize some human characteristics, such as sex, and use these characteristics to classify them into different categories. The aim of this study was to explore whether domesticated horses are able to cross-modally recognize women and men according to visual and auditory cues, using a preferential looking paradigm. We simultaneously presented two videos of women and men's faces, while playing a recording of a human voice belonging to one of these two categories through a loudspeaker. The results showed that the horses looked significantly more towards the congruent video than towards the incongruent video, suggesting that they are able to associate women's voices with women's faces and men's voices with men's faces. Further investigation is necessary to determine the mechanism underlying this recognition, as it might be interesting to determine which characteristics horses use to categorize humans. These results suggest a novel perspective that could allow us to better understand how horses perceive humans.

Horses discriminate human body odors between fear and joy contexts in a habituation-discrimination protocol.

Animals are widely believed to sense human emotions through smell. Chemoreception is the most primitive and ubiquitous sense, and brain regions responsible for processing smells are among the oldest structures in mammalian evolution. Thus, chemosignals might be involved in interspecies communication. The communication of emotions is essential for social interactions, but very few studies have clearly shown that animals can sense human emotions through smell. We used a habituation-discrimination protocol to test whether horses can discriminate between human odors produced while feeling fear vs. joy. Horses were presented with sweat odors of humans who reported feeling fear or joy while watching a horror movie or a comedy, respectively. A first odor was presented twice in successive trials (habituation), and then, the same odor and a novel odor were presented simultaneously (discrimination). The two odors were from the same human in the fear or joy condition; the experimenter and the observer were blinded to the condition. Horses sniffed the novel odor longer than the repeated odor, indicating they discriminated between human odors produced in fear and joy contexts. Moreover, differences in habituation speed and asymmetric nostril use according to odor suggest differences in the emotional processing of the two odors.

Captive Blue-and-yellow macaws (Ara ararauna) show facial indicators of positive affect when reunited with their caregiver.

In mammals, human-animal bonding is recognized as a source of positive affect for companion or farm animals. Because this remains unexplored in birds, we investigated captive parrots' perspective of the human-animal relationship. We used a classical separation-reunion paradigm and predicted that variations in parrots' facial displays and behaviours would indicate their appraisal of the relationship. The test was divided into three phases of two minutes each: the bird was placed in an unfamiliar environment with a familiar caregiver (union), then the bird was left alone (separation) and finally, the caregiver returned (reunion). The test was repeated 10 times for each bird and video recorded in order to analyze their behaviour. The data show significantly higher crown and nape feather heights, higher redness of the skin and higher frequency of contact-seeking behaviours during the union and reunion phases than during the separation phase during which they expressed long distance contact calls. We observed the expression of eye pinning during the union and reunion phases in one out of five macaws. We argue that variation in facial displays provides indicators of parrot's positive appraisal of the caretaker presence. Our results broaden the scope for further studies on parrots' expression of their subjective feelings.

Short photoperiod modulates behavior, cognition and hippocampal neurogenesis in male Japanese quail.

The mechanisms underlying the photoperiodic control of reproduction in mammals and birds have been recently clarified. In contrast, the potential impact of photoperiod on more complex, integrative processes, such as cognitive behaviors, remains poorly characterized. Here, we investigated the impact of contrasted long and short photoperiods (LP, 16 h light/day and SP, 8 h light/day, respectively) on learning, spatial orientation abilities, and emotional reactivity in male Japanese quail. In addition, we quantified cell proliferation and young cell maturation/migration within the hippocampus, a brain region involved in spatial orientation. Our study reveals that, in male quail, SP increases emotional responses and spatial orientation abilities, compared to LP. Behaviorally, SP birds were found to be more fearful than LP birds, exhibiting more freezing in the open field and taking longer to exit the dark compartment in the emergence test. Furthermore, SP birds were significantly less aggressive than LP birds in a mirror test. Cognitively, SP birds were slower to habituate and learn a spatial orientation task compared to LP birds. However, during a recall test, SP birds performed better than LP birds. From a neuroanatomical standpoint, SP birds had a significantly lower density of young neurons, and also tended to have a lower density of mature neurons within the hippocampus, compared to LP birds. In conclusion, our data reveal that, beyond breeding control, photoperiod also exerts a profound influence on behavior, cognition, and brain plasticity, which comprise the seasonal program of this species.

Early castration in foals: Consequences on physical and behavioural development.

BACKGROUND: The impact of very early castration of foals has not yet been studied despite the many positive effects observed in dogs and cats. OBJECTIVES: The objective of the study was to compare castration at 3 days and 18 months and assess their subsequent morphological and behavioural development. STUDY DESIGN: This was a randomised, blinded clinical study. METHODS: Twenty-two Welsh ponies underwent either early (3-day old, EC group, n = 11) or traditional (18-month old, TC group, n = 11) castration. Animals were followed up to 3 years of age. All ponies were castrated using a primary closure technique under general anaesthesia. Weight and morphometric measurements were monitored monthly from birth until 8 months of age in both groups. Then, measurements were taken every 3 months until 2 years of age and then every 6 months until 3 years of age. Temperament tests were performed on all animals when they were 1- and 3-years old. RESULTS: No differences were observed between the EC and TC groups in terms of physical development from birth until 40 months of age or in terms of temperament and behaviour at either 1 or 3 years of age. MAIN LIMITATIONS: The study included only one breed (Welsh ponies) and only 22 animals that were castrated before 2 years of age, precluding comparison with castration performed at older ages. CONCLUSIONS: We demonstrate that early castration at 3 days does not interfere with morphological or behavioural development.

Horses form cross-modal representations of adults and children.

Recently, research on domestic mammals' sociocognitive skills toward humans has been prolific, allowing us to better understand the human-animal relationship. For example, horses have been shown to distinguish human beings on the basis of photographs and voices and to have cross-modal mental representations of individual humans and human emotions. This leads to questions such as the extent to which horses can differentiate human attributes such as age. Here, we tested whether horses discriminate human adults from children. In a cross-modal paradigm, we presented 31 female horses with two simultaneous muted videos of a child and an adult saying the same neutral sentence, accompanied by the sound of an adult's or child's voice speaking the sentence. The horses looked significantly longer at the videos that were incongruent with the heard voice than at the congruent videos. We conclude that horses can match adults' and children's faces and voices cross-modally. Moreover, their heart rates increased during children's vocalizations but not during adults'. This suggests that in addition to having mental representations of adults and children, horses have a stronger emotional response to children's voices than adults' voices.

How Useful Are Existing Protocols in the Quick Assessment of the Welfare of Semi-Feral Horses? Pilot Study on Konik Polski Horses Living in the Forest Sanctuary.

Scientifically validated and standardised methods for the evaluation of the welfare of free-living horses are urgently needed by both the owners and managers of these populations and those responsible for implementing national welfare legislation. The aim of the study was to test the feasibility and usefulness of two welfare protocols that could be applied to semi-feral populations: a prototype of welfare assessment template (WAT) for Carneddau semi-feral ponies and the IFCE/INRAE Horse Welfare Protocol. Additionally, the body condition scale designed by Henneke (BCS-H) was employed. The study took place in July/August 2022 and April 2023 to evaluate the welfare of a pilot population of nineteen semi-feral Konik polski horses. The horses scored high or satisfactory under indicators across both protocols; only body condition scores were significantly lower in early spring (BCS-WAT: 1.11 ± 0.57; BCS-H: 3.84 ± 1.17) than in the summer (BCS-WAT: 1.58 ± 0.61; BCS-H: 5.63 ± 1.01). Our study confirmed the feasibility of utilising most of the WAT and IFCE/INRAE welfare indicators in semi-feral horses. Some adaptations, such as considering validation of scales, positive welfare indicators and animals' free-choice of conditions, have been suggested for future in-field application.

Weaned horses, especially females, still prefer their dam after five months of separation.

Under natural conditions, foals stop nursing from their dam at approximately-9 months old, but their bond persists until 1.5-2.5 years of age. In contrast, in horse breeding, foals are generally artificially weaned and totally separated from their dam at 5-7 months. However, it is not known whether the bond between the dam and her foal is maintained after artificial weaning. The aim of this study was (1) to assess whether foals still recognise and prefer their dam over other familiar mares several months after weaning and (2) to evaluate whether the preference for the dam is more pronounced in fillies or colts. Fifteen fillies and 19 colts were weaned at the age of 7 months old (complete separation from the mother). At the age of one year (i.e., 5 months after the separation), they underwent a test evaluating their preference for their dam or a familiar mare from their natal group. Significantly more foals first approached their dam; they also sniffed and tended to look more often at her. This finding indicates that artificially weaned horses remember and still exhibit a preference for their dam, suggesting that the bond persists even after 5 months of separation. Moreover, fillies exhibited a stronger preference for both mares than colts: they looked at them more frequently, sniffed them for a longer duration and spent more time in proximity to both mares than colts. This suggests that fillies generally have an even stronger attachment to their dam as well as to other mares from their natal group. This study calls into question the practice of artificial weaning at 5-7 months of age.

Domestic hens succeed at serial reversal learning and perceptual concept generalisation using a new automated touchscreen device.

Improving the welfare of farm animals depends on our knowledge on how they perceive and interpret their environment; the latter depends on their cognitive abilities. Hence, limited knowledge of the range of cognitive abilities of farm animals is a major concern. An effective approach to explore the cognitive range of a species is to apply automated testing devices, which are still underdeveloped in farm animals. In screen-like studies, the uses of automated devices are few in domestic hens. We developed an original fully automated touchscreen device using digital computer-drawn colour pictures and independent sensible cells adapted for cognitive testing in domestic hens, enabling a wide range of test types from low to high complexity. This study aimed to test the efficiency of our device using two cognitive tests. We focused on tasks related to adaptive capacities to environmental variability, such as flexibility and generalisation capacities as this is a good start to approach more complex cognitive capacities. We implemented a serial reversal learning task, categorised as a simple cognitive test, and a delayed matching-to-sample (dMTS) task on an identity concept, followed by a generalisation test, categorised as more complex. In the serial reversal learning task, the hens performed equally for the two changing reward contingencies in only three reversal stages. In the dMTS task, the hens increased their performance rapidly throughout the training sessions. Moreover, to the best of our knowledge, we present the first positive result of identity concept generalisation in a dMTS task in domestic hens. Our results provide additional information on the behavioural flexibility and concept understanding of domestic hens. They also support the idea that fully automated devices would improve knowledge of farm animals' cognition.

Pet-directed speech improves horses' attention toward humans.

In a recent experiment, we showed that horses are sensitive to pet-directed speech (PDS), a kind of speech used to talk to companion animals that is characterized by high pitch and wide pitch variations. When talked to in PDS rather than adult-directed speech (ADS), horses reacted more favorably during grooming and in a pointing task. However, the mechanism behind their response remains unclear: does PDS draw horses' attention and arouse them, or does it make their emotional state more positive? In this study, we used an innovative paradigm in which female horses watched videos of humans speaking in PDS or ADS to better understand this phenomenon. Horses reacted differently to the videos of PDS and ADS: they were significantly more attentive and their heart rates increased significantly more during PDS than during ADS. We found no difference in the expressions of negative or positive emotional states during PDS and ADS videos. Thus, we confirm that horses' perception of humans can be studied by means of video projections, and we conclude that PDS attracts attention and has an arousing effect in horses, with consequences on the use of PDS in daily interactions with them.

Foraging Behavior Shows Individual-Consistency Over Time, and Predicts Range Use in Slow-Growing Free-Range Male Broiler Chickens.

Recent research on free-range chickens shows that individual behavioral differences may link to range use. However, most of these studies explored individual behavioral differences only at one time point or during a short time window, assessed differences when animals were out of their social group and home environment (barn and range), and in specific tests or situations. Therefore, it is yet unclear how different behaviors relate to range use and how consistent these behaviors are at the individual level. To fill this gap, we here aimed to describe the behavioral budget of slow-growing male broiler chickens (S757N) when in their social group and home environment during the whole rearing period (from the second week of life to the twelfth week, before slaughter), and to relate observed behavioral differences to range use. For this, we followed a sample of individuals in two flocks (n = 60 focal chickens out of 200 chickens per flock), over two seasons, during three periods: before range access (from 14 to 25 days old), during early range access (first weeks of range access, from 37 to 53 days old), and during late range access (last weeks of range access, from 63 to 87 days old). By the end of each period, individual tests of exploration and social motivation were also performed, measuring exploration/activity and sociability propensities. Our results show that foraging (i.e., pecking and scratching at the ground) was the only behavior that correlated to range use for all three rearing periods, independent of the season. Foraging was also the only behavior that showed within-individual consistency from an early age and across the three rearing periods. Foraging may, therefore, serve as a useful behavioral predictor of range use in free-range broiler chickens. Our study increases the knowledge of how behaviors develop and relate to each other in a domesticated and intensely selected species, and improves our understanding of the biology of free-range broiler chickens. These findings can, ultimately, serve as a foundation to increase range use and improve chicken welfare.

Human-controlled reproductive experience may contribute to incestuous behavior observed in reintroduced semi-feral stallions (Equuscaballus).

Equine reproductive behavior is affected by many factors, some remaining poorly understood. This study tested the hypothesis that a period of captivity during the juvenile period and human-controlled reproduction may potentially be involved in the disruption of the development of incestuous mating avoidance behavior in sanctuary-reintroduced male Konik polski horses. Between 1986 and 2000, cases of incestuous behavior in harem stallions born and reared until weaning in the sanctuary were studied. Eight males lived in the sanctuary's feral herd for the rest of their lives (the non-captive group; nC). They gained their own harem of mares without human intervention (no human-controlled reproductive activity, nHC). Another five stallions were removed as weanlings, reared in captivity and then reintroduced as adults (captive, C). Three of these C stallions were used as in-hand breeding stallions, one as a "teaser" (human-controlled reproductive activity, HC) and one was not used for reproduction in captivity (nHC). Reproductive records for 46 mares, daughters of all 13 harem stallions, were scrutinized and cases of incestuous breeding were recorded by interrogation of foal parentage records. C stallions failed to expel more daughters than nC stallions (33% vs. 18%, P = 0.045), and mated with significantly more of them (28% vs. 11%, P = 0.025). Interestingly, HC stallions expelled fewer (60%) and successfully mated with more (33%) daughters that nHC stallions (84% expelled, P = 0.013, and 10% successful mating with daughters, P = 0.010). All HC stallions bred incestuously at least once. We propose that human intervention during a critical period of development of social and reproductive behavior in young stallions, by enforced separation from their natal herd and in-hand breeding, may contribute to their later aberrant behavior and disruption of inbreeding avoidance mechanisms in these stallions. The previous occurrence of human-controlled breeding may be one of the factors promoting incestuous behavior of stallions in natural conditions. The uninterrupted presence of stallions in their harems and herd member recognition may also play important roles in inbreeding avoidance in horses.

Cognition and the human-animal relationship: a review of the sociocognitive skills of domestic mammals toward humans.

In the past 20 years, research focusing on interspecific sociocognitive abilities of animals toward humans has been growing, allowing a better understanding of the interactions between humans and animals. This review focuses on five sociocognitive abilities of domestic mammals in relation to humans as follows: discriminating and recognizing individual humans; perceiving human emotions; interpreting our attentional states and goals; using referential communication (perceiving human signals or sending signals to humans); and engaging in social learning with humans (e.g., local enhancement, demonstration and social referencing). We focused on different species of domestic mammals for which literature on the subject is available, namely, cats, cattle, dogs, ferrets, goats, horses, pigs, and sheep. The results show that some species have remarkable abilities to recognize us or to detect and interpret the emotions or signals sent by humans. For example, sheep and horses can recognize the face of their keeper in photographs, dogs can react to our smells of fear, and pigs can follow our pointing gestures. Nevertheless, the studies are unequally distributed across species: there are many studies in animals that live closely with humans, such as dogs, but little is known about livestock animals, such as cattle and pigs. However, on the basis of existing data, no obvious links have emerged between the cognitive abilities of animals toward humans and their ecological characteristics or the history and reasons for their domestication. This review encourages continuing and expanding this type of research to more abilities and species.

Horses with sustained attention follow the pointing of a human who knows where food is hidden.

When interacting with humans, domesticated species may respond to communicative gestures, such as pointing. However, it is currently unknown, except for in dogs, if species comprehend the communicative nature of such cues. Here, we investigated whether horses could follow the pointing of a human informant by evaluating the credibility of the information about the food-hiding place provided by the pointing of two informants. Using an object-choice task, we manipulated the attentional state of the two informants during food-hiding events and differentiated their knowledge about the location of the hidden food. Furthermore, we investigated the horses' visual attention levels towards human behaviour to evaluate the relationship between their motivation and their performance of the task. The result showed that horses that sustained high attention levels could evaluate the credibility of the information and followed the pointing of an informant who knew where food was hidden (Z = - 2.281, P = 0.002, n = 36). This suggests that horses are highly sensitive to the attentional state and pointing gestures of humans, and that they perceive pointing as a communicative cue. This study also indicates that the motivation for the task should be investigated to determine the socio-cognitive abilities of animals.

Handler familiarity helps to improve working performance during novel situations in semi-captive Asian elephants.

Working animals spend hours each day in close contact with humans and require training to understand commands and fulfil specific tasks. However, factors driving cooperation between humans and animals are still unclear, and novel situations may present challenges that have been little-studied to-date. We investigated factors driving cooperation between humans and animals in a working context through behavioural experiments with 52 working semi-captive Asian elephants. Human-managed Asian elephants constitute approximately a third of the remaining Asian elephants in the world, the majority of which live in their range countries working alongside traditional handlers. We investigated how the familiarity and experience of the handler as well as the elephant's age and sex affected their responses when asked to perform a basic task and to cross a novel surface. The results highlighted that when novelty is involved in a working context, an elephant's relationship length with their handler can affect their cooperation: elephants who had worked with their handler for over a year were more willing to cross the novel surface than those who had a shorter relationship with their handler. Older animals also tended to refuse to walk on the novel surface more but the sex did not affect their responses. Our study contributes much needed knowledge on human-working animal relationships which should be considered when adjusting training methods and working habits.