The recognition oftheiremotional lives, cognitive abilities, and a certain form of consciousness in animals under human care has profoundly changed the way their well-being is considered. While it is essential to provide living conditions that meet their behavioural needs, it is equally important to offer stimuli that allow the animal to actively interact with its environment in order to develop its adaptive capacities, particularly through environmental enrichment. Due to the much later recognition of fish as sentient beings compared to terrestrial vertebrates, current knowledge on ways to improve their well-being through environmental enrichment remains fragmented. This issue is becoming a major societal concern due to the growing interest of society and various sectors in fish welfare. This non-exhaustive review aims to present recent research avenues concerning fish under human care, whether for commercial, recreational, conservation, or experimental purposes. Afterdefining what is meant by enrichment, we will discuss the physical, sensory, social, and cognitive enrichment strategies currently being explored, before addressing the limitations of their implementation as well as future perspectives. Fish represent a far greater number of species than terrestrial vertebrates. We will see that while some enrichment strategies show promising potential, much knowledge still needs to be acquired to develop relevant strategies adapted to each species and life stage.

In mammals, facial expressions serve as a window into a variety of affective states. Emotional facial expressions in birds have received little scientific attention. Juvenile hens showed variations in their facial display, facial redness and head feather position, depending on their affective states. When sexually mature, domestic hens develop fleshy coloured outgrowths such as the comb and the wattles, which are considered as functional visual signals of the individual's health or fertility. Here, we investigated, in adult hens, whether bare facial skin (cheeks, ear lobes, comb and wattles) redness and head feather position (sleeked/fluffed) also vary with affective states. We hypothesized little or no redness variations for the wattles and comb because of their involvement as functional signals. Twelve adult hens were filmed during naturally occurring situations and controlled tests varying in their valence and arousal level: situations associated with fear/frustration (negative valence/high arousal), situations associated with appetitive motivational states (positive valence/high arousal) and situations associated with calm/contentment (positive valence/low arousal). We showed that positive situations of low arousal were associated with the lowest redness in all areas of bare skin. A significantly higher level of redness was found in high arousal compared with low arousal positive situations, except for the comb. The highest redness was observed in all skin areas in negative situations of high arousal. Head feathers were observed more frequently fluffed in most of the positive situations and sleeked in all feedrelated situations. Thus, facial skin redness, including secondary sexual characters and head feather position are potential markers of the affective states of adult hens. Studying facial displays opens new research opportunities to increase our knowledge on how avian species perceive their environment. Further research is needed to clearly differentiate between the effects of valence and arousal. (c) 2025 The Authors. Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour. This is an open access article under the CC BY license (http://creativecommons.org/licenses/ by/4.0/).

La reconnaissance d’une sensibilité émotionnelle, de capacités cognitives et d’une certaine forme de conscience chez les animaux sous la responsabilité des humains a profondément modifié la manière d’appréhender leur bien-être. S’il est impératif de fournir des conditions de vie répondant à leurs besoins comportementaux, il est également essentiel d’offrir des stimulations permettant à l’animal d’interagir activement avec son environnement afin de développer ses capacités d’adaptation, notamment par l’enrichissement du milieu de vie. En raison de la reconnaissance bien plus tardive des poissons en tant qu’êtres sensibles par rapport aux vertébrés terrestres, les connaissances actuelles sur les moyens d’améliorer leur bien-être par l’enrichissement du milieu restent parcellaires. Ce sujet devient un enjeu sociétal majeur en raison de l’intérêt croissant de la société et des filières pour le bien-être des poissons. Cette synthèse, non exhaustive, vise à exposer les pistes de recherche récentes concernant les poissons sous la dépendance des humains, que ce soit à des fins commerciales, récréatives, de conservation ou d’expérimentation. Après avoir défini ce que l’on entend par un enrichissement, nous aborderons les stratégies d’enrichissement physique, sensoriel, social ou cognitif actuellement explorées, avant d’évoquer les limites à leur mise en place ainsi que les perspectives futures. Les poissons représentent un bien plus grand nombre d’espèces que les vertébrés terrestres. Nous verrons que certains enrichissements offrent des pistes prometteuses, mais que de nombreuses connaissances restent à acquérir afin de développer des stratégies pertinentes adaptées à chaque espèce et stade de vie.

Broiler chickens are reared in very large groups and are submitted to repeated encounters with unknown conspecifics. Our aim was to assess the consequences of these encounters on broiler chickens learning abilities, social behaviour and emotional reactivity. Groups of 5 male medium-growing broiler chickens (JA 957) were reared under stable or unstable social conditions (n=16 groups in each condition). Under the unstable conditions, the 5 birds of the groups changed every 3-4 days from 1 to 52 days of age. Broiler chickens were tested in an associative learning task (conditioned place preference) between 14 and 18 days of age. A highly palatable food (mealworms) was delivered in a particular environment with coloured stripes and preference for this environment was then tested. Social behaviours were analysed between 21 and 43 days of age. Emotional reactivity of the birds when faced with novelty (reactivity to unknown food, object and human) was assessed at the end of the rearing period (46-51 days of age). Only birds from the stable condition were able to associate the palatable food to the environment in which it was delivered (p = 0.02). Furthermore, these birds had a higher social proximity than those from the unstable condition (p <0.01) and showed less aggressive interactions such as aggressive pecking (p = 0.06), threat (p < 0.01) or facing each other with wing flapping in front of another bird (p = 0.03). The social conditions tested had no significant effect on their emotional reactivity. Our results evidenced that chickens exposed to social instability have some learning abilities impaired compared to chickens maintained in stable social conditions. In addition, social instability increases aggressive interactions between conspecifics, while social stability favours bird proximity. In conclusion, being reared in unstable social conditions likely impairs bird adaptation to their rearing environment and increases deleterious consequences of aggressive interactions with conspecifics.

Animal welfare is a result of accumulated negative and positive emotions. Therefore, it is important to limit the former and promote the latter. Doing so requires a precise identification of these emotions. The aim of this study was to characterize the behaviours, facial expressions and locomotor parameters of 21 horses in two conditions with opposite valence, presumed to induce positive anticipation and frustration. In the positive anticipation situation, the horse was led to a bucket of food, knowing that they would be allowed to eat it. In the frustration situation, experimenters indicated to the horse that food was available without allowing them to eat it. In the positive anticipation situation, horses exhibited a lower neck position with the ears forward and upper lip advanced and went faster by increasing their stride frequency accompanied by increased global locomotor activity. In the frustration situation, horses exhibited a higher neck position with the ears backward or to the side, accompanied by ear movements and eye blinks, and interacted more with the experimenters. This study describes new possible indicators of positive anticipations and frustration in horses.

The study of facial expressions in mammals provided great advances in the identification of their emotions and then in the comprehension of their sentience. So far, this area of research has excluded birds. With a naturalist approach, we analysed facial blushing and feather displays in domestic fowl. Hens were filmed in situations contrasting in emotional valence and arousal level: situations known to indicate calm states (positive valence / low arousal), have rewarding effects (positive valence / high arousal) or induce fear-related behaviour (negative valence / high arousal). Head feather position as well as skin redness of comb, wattles, ear lobes and cheeks varied across these situations. Skin of all four areas was less red in situations with low arousal compared to situations with higher arousal. Furthermore, skin redness of the cheeks and ear lobes also varied depending on the valence of the situation: redness was higher in situations with negative valence compared to situations with positive valence. Feather position also varied with the situations. Feather fluffing was mostly observed in positively valenced situations, except when hens were eating. We conclude that hens have facial displays that reveal their emotions and that blushing is not exclusive to humans. This opens a promising way to explore the emotional lives of birds, which is a critical step when trying to improve poultry welfare.

Prenatal maternal stress (PMS) is known to shape the phenotype of the first generation offspring (F1) but according to some studies, it could also shape the phenotype of the offspring of the following generations. We previously showed in the Japanese quail that PMS increased the emotional reactivity of F1 offspring in relation to (i) a variation in the levels of some histone post-translational modification (H3K27me3) in their brains and (ii) a modulation of the hormonal composition of the eggs from which they hatched. Here, we wondered whether PMS could also influence the behaviour of the second (F2) and third (F3) generation offspring due to the persistence of the specific marks we identified. Using a principal component analysis, we found that PMS influenced F2 and F3 quail profiles with subtle differences between generations. It increased F2 neophobia, F3 fearfulness and F3 neophobia but only in females. Interestingly, we did not find any variations in the level of histone post-translational modification in F3 brains and we observed inconsistent modulations of androstenedione levels in F1 and F2 eggs. Although they may vary over generations, our results demonstrate that PMS can have phenotypical effects into the third generation.

To improve the early perinatal conditions of broiler chicks, alternative hatching systems have been developed. On-farm hatching (OFH) with an enriched microbial and stimulating environment by the presence of an adult hen is a promising solution. Day-old chicks were allotted within five hatching and rearing conditions: OFH, conventional hatchery (CH), CH and post-hatching treatment with antibiotics (CH + AB), as well as both hatching systems with an adult hen at hatching (OFH + H, CH + H). To challenge the robustness of chickens, they were exposed on D27 to suboptimal rearing conditions by combining for 4 h transport in boxes in a new room at a lower temperature and fasting. On their return to the original room, the chicken density was increased, and birds were orally vaccinated with the Gumboro vaccine. The impacts of these conditions on hatchability, chick quality score, performance, health and robustness were determined. The OFH chick body weights (BWs) were significantly greater than those of CH chicks at hatching. Whereas there was no effect of hatching conditions, the presence of hens decreased the hatchability rate, the quality score of OFH chicks and increased mortality at hatching. Treatment of CH chicks with antibiotics (CH + AB) temporarily decreased chicken BW at D19, but the feed conversion ratio (FCR) was not modified. At D19, OFH chicks had the highest BW compared to the other groups, and the presence of hens at hatching harmed chicken BW regardless of the hatching condition and FCR. An interaction between the effect of experimental rearing conditions and chicken sex was observed later for BW. In males, the OFH chickens were the heaviest compared to the other groups at D34 but not at D56. The presence of hens negatively impacted CH chicken BW at D56. In females, there was no effect of hatching condition on the BWs at D34 and D56, and the presence of hens had a positive impact on OFH chicken BW. There was no effect of hatching conditions on health parameters. In conclusion, the OFH system was a hatching system at least equivalent to the CH system. The presence of the hen at hatching and during the chick start-up phase on performance interacted with the hatching condition and the sex of the chickens.

Non-invasive markers of affective states can help understanding animals' perception of situations and improving their welfare. These markers are scarce in avian species. In this study, we investigate the potential relation between alterations in facial skin redness in hens and their corresponding affective states. Six hens were filmed in both naturally unfolding scenarios and controlled tests designed to elicit various affective states. The facial skin redness was measured from images extracted from the videos. Our observations revealed that hens exhibited the highest degree of facial skin redness in negative situations of high arousal, a high redness in positive situations of high arousal, and the lowest in positive situations of low arousal. In a second study, we further examined whether facial skin redness and secretory immunoglobulin A (S-IgA) can serve as markers for the quality of the human-animal relationship. Two groups of hens, one habituated to humans (n=13) and one non-habituated (n=12), were compared for general fearfulness in an open field test and for fear of humans in a reactivity to human test. In the open-field test, there were no statistical differences in general fearfulness, facial skin redness or S-IgA concentrations between both groups. However, habituated hens exhibited significantly lower fearfulness and facial skin redness in the presence of humans compared to non-habituated hens in the reactivity to human test. Additionally, habituated hens showed significant lower S-IgA concentration in lachrymal fluid in the presence of humans, with no significant differences in saliva or cloacal samples. We propose that changes in facial skin redness reflect variations in affective states and can be used as a marker for assessing the quality of the human-hen relationship. The relationship between S-IgA concentrations and affective states requires further investigation.

As human influence reshapes communities worldwide, many species expand or shift their ranges as a result, with extensive consequences across levels of biological organization. Range expansions can be ranked on a continuum going from pulled dynamics, in which low-density edge populations provide the “fuel” for the advance, to pushed dynamics in which high-density rear populations “push” the expansion forward. While theory suggests that evolution during range expansions could lead pushed expansions to become pulled with time, empirical comparisons of phenotypic divergence in pushed vs. pulled contexts are lacking. In a previous experiment using Trichogramma brassicae wasps as a model, we showed that expansions were more pushed when connectivity was lower. Here we used descendants from these experimental landscapes to look at how the range expansion process and connectivity interact to shape phenotypic evolution. Interestingly, we found no clear and consistent phenotypic shifts, whether along expansion gradients or between reference and low connectivity replicates, when we focused on low-density trait expression. However, we found evidence of changes in density-dependence, in particular regarding dispersal: populations went from positive to negative density-dependent dispersal at the expansion edge, but only when connectivity was high. As positive density-dependent dispersal leads to pushed expansions, our results confirm predictions that evolution during range expansions may lead pushed expansions to become pulled, but add nuance by showing landscape conditions may slow down or cancel this process. This shows we need to jointly consider evolution and landscape context to accurately predict range expansion dynamics and their consequences.

In mammals, positive tactile contact is recognized as an effective tool for triggering positive affective states. In this study, we investigated the benefits of providing automatic rotative brushes for group-housed horses. Our three aims were: i. to determine whether horses used automatic brushes and if so how they used them; ii. to investigate whether the presence of these brushes induced positive social behaviors (allogrooming), or aggressiveness, as can be observed when there is competition for a desired resource; iii to provide a preliminary explanation of the role of the positive facial expression displayed by some horses while being groomed. Two brushes were installed in a large stable with free access to a paddock in which 40 horses were housed 7h30/day. For four days, video-cameras placed above the brushes continuously filmed the horses. First, analysis of the video footage demonstrated that brushes were used by almost 90% of the horses, mainly on areas that are not easily accessible to another horse during allogrooming, such as the head. Secondly, it revealed that among the horses that used the brush, 25.7% expressed positive social behavior (allogrooming) at the same time, while none expressed aggressive behaviors. The brush thus seems to act as a catalyst for affiliative behaviors rather than competition for a desired resource. Thirdly, we observed their facial expression, especially a positive one, described when the horse are groomed (neck moderately raised, eyes open or half-closed, upper lip extended, ears turned backwards almost in line with the nose). More than half of the horses displayed this while using the brushes. Interestingly, when a horse expressed this face, it had a six times greater chance to subsequently start allogrooming with a conspecific. Several possible explanations are discussed, including a social function. Further research is needed to validate other criteria to help qualify whether or not this positive facial expression is a signal of intentional communication. From a practical point of view, the results show that the brushes provide both a direct benefit to the horses enabling them to conduct self-grooming, and also in all likelihood, an indirect benefit by facilitating positive social interactions. This study encourages the installation of automatic brushes in places where horses are kept to improve their welfare and provide comfort.

As the sensory systems of vertebrates develop prenatally, embryos perceive many environmental stimuli that can influence the ontogeny of their behaviour. Whether the nature and intensity of prenatal stimuli affect this ontogeny differently remains to be investigated. In this context, this study aimed to analyse the effects of prenatal auditory stimulation (natural stimulation, NS; predator vocalisations or artificial stimulation, AS; metallic sounds) on the subsequent behaviour of young Japanese quail (Coturnix coturnix japonica). For this, behavioural variables recorded during ethological tests evaluating emotional and social reactivity were analysed using a principal component analysis. This analysis revealed significant differences between the behavioural profile of stimulated chicks and that of non-exposed chicks. Indeed, chicks exposed to NS expressed more intense emotional responses in fearful situations, but less neophobia when exposed to a novel environment or object, whereas chicks exposed to AS appeared more sensitive to social isolation. Our results show that the acoustic environment of embryos can influence the way young birds subsequently interact with their social and physical environment after hatching, and face challenges in changing living conditions.

Prenatal maternal stress (PMS) influences many facets of offspring's phenotype including morphology, behaviour and cognitive abilities. Recent research suggested that PMS also induced epigenetic modifications. In the present study, we analysed, in the Japanese quail, the effects of PMS on the emotional reactivity and cognitive abilities of the F1 offspring. We also investigated in the hippocampus, the paraventricular hypothalamic nucleus and subnuclei of the arcopallium/amygdala the level of two histone post-translational modifications, H3K4me2 and H3K27me3, known to be impacted by stress. We found that PMS does not affect F1 quail's learning abilities but increases their emotional reactivity. Moreover, we demonstrated that PMS induced an increased density of H3K27me3 positive cells, in the hippocampus, paraventricular hypothalamic nucleus and dorsal nucleus of the amygdala, but not variations of H3K4me2. As these brain regions are involved in the control of vertebrates' emotional responses, the effect of PMS on the epigenetic mark H3K27me3 could possibly be a mechanism involved in the behavioural effects we observed in F1 quail.

Range expansions are key processes shaping the distribution of species; their ecological and evolutionary dynamics have become especially relevant today, as human influence reshapes ecosystems worldwide. Many attempts to explain and predict range expansions assume, explicitly or implicitly, so‐called ‘pulled' expansion dynamics, in which the low‐density edge populations provide most of the ‘fuel' for the species advance. Some expansions, however, exhibit very different dynamics, with high‐density populations behind the front ‘pushing' the expansion forward. These two types of expansions are predicted to have different effects on e.g. genetic diversity and habitat quality sensitivity. However, empirical studies are lacking due to the challenge of generating reliably pushed versus pulled expansions in the laboratory, or discriminating them in the field. We here propose that manipulating the degree of connectivity among populations may prove a more generalizable way to create pushed expansions. We demonstrate this with individual‐based simulations as well as replicated experimental range expansions (using the parasitoid wasp Trichogramma brassicae as model). By analyzing expansion velocities and neutral genetic diversity, we showed that reducing connectivity led to pushed dynamics. Low connectivity alone, i.e. without density‐dependent dispersal, can only lead to ‘weakly pushed' expansions, where invasion speed conforms to pushed expectations, but the decline in genetic diversity does not. In empirical expansions however, low connectivity may in some cases also lead to adjustments to the dispersal‐density function, recreating ‘classical' pushed expansions. In the current context of habitat loss and fragmentation, we need to better account for this relationship between connectivity and expansion regimes to successfully predict the ecological and evolutionary consequences of range expansions.