How outing conditions relate to the motivation of movement-restricted cattle to access an outdoor exercise yard.

Assessing animal motivation to access a given resource is one method available to evaluate what to provide in the living environments of captive animals. Providing increased opportunities for movement can be seen as an important source of enrichment, but we need to know the point of view of the animal. The objective of our study was to test a novel combination of behaviours in order to assess the motivation of cows to access an outdoor exercise paddock. Three trials were conducted, each enrolling 15-16 tie-stall-housed cows as a model for movement-restricted animals. Cows were provided with access to an outdoor exercise yard 5 days/week for the duration of the trial, each trial presenting different conditions such as paddock size, duration of access and animal handling. We recorded the trips' durations and cows' behaviours during the trips going to (go-out) and coming back (go-in) from the paddock. LMr comparisons on PCA were used to assess cow motivation profiles. The same two dimensions of speed and stop quality emerged from the PCA in all three trials, showing the method's robustness. Additionally, three motivation profiles were established, representing how the cows' motivation was affected by the conditions prevailing in each trial.

Fos expression in the medial preoptic area and nucleus accumbens of female Japanese quail (Coturnix japonica) after maternal induction and interaction with chicks.

The neural system underlying maternal caregiving has often been studied using laboratory rodents and a few other mammalian species. This research shows that the medial preoptic area (mPOA) integrates sensory cues from the young that, along with hormonal and other environmental signals, control maternal acceptance of neonates. The mPOA then activates the mesolimbic system to drive maternal motivation and caregiving activities. How components of this neural system respond to maternal experience and exposure to young in non-mammals has rarely been examined. To gain more insight into this question, virgin female Japanese quail (Coturnix japonica) were induced to be maternal through four days of continuous exposure to chicks (Maternal), or were not exposed to chicks (Non-Maternal). Chicks were removed overnight from the Maternal group and half the females from each group were then exposed to chicks for 90 minutes (Exposed), or not exposed to chicks (Non-Exposed), before euthanasia. The number of Fos-immunoreactive (Fos-ir) cells was examined as a marker of neuronal activation. As expected, repeated exposure to chicks induced caregiving behavior in the Maternal females, which persisted after the overnight separation, suggesting the formation of a maternal memory. In contrast, Non-Maternal females were aggressive and rejected the chicks when exposed to them. Exposed females, whether or not they were given prior experience with chicks (i.e., regardless if they accepted or rejected chicks during the exposure before euthanasia), had more Fos-ir cells in the mPOA compared to Non-Exposed females. In the nucleus accumbens (NAC), the number of Fos-ir cells was high in all Maternal females whether or not they were Exposed to chicks again before euthanasia. In the lateral bed nucleus of the stria terminalis, a site involved in general stress responding, groups did not differ in the number of Fos-ir cells. These data indicate a conserved role for the mPOA and NAC in maternal caregiving across vertebrates, with the mPOA acutely responding to the salience rather than valence of offspring cues, and the NAC showing longer-term changes in activity after a positive maternal experience even without a recent exposure to young.

Chicks from stressed females elicit overprotective behaviour in adoptive mother quail.

Beyond the genetic links between mother and young, prenatal effects experienced by offspring can modulate their mother's behaviour and stress. Here we investigated the consequences of stress-induced prenatal maternal effects on quail chicks' behaviour and maternal behaviour. We evaluated how adoptive quail mothers (unstressed) raised chicks of females that had either been stressed (PS) or were non-stressed (NPS) during laying. We demonstrated previously that our social stress procedure during laying increases levels of egg yolk steroid and chicks' emotional reactivity during the postnatal period. We found that, compared to NPS mothers, PS mothers remained closer to their chicks, were aggressed and neglected them less, emitted more maternal vocalisations and called their chicks more when they were separated. Moreover, PS chicks made more requests on their mother. Our results show that mothers responded to chicks' behavioural changes caused by prenatal stress by adapting their maternal behaviour. As exemplified here, for the first time in birds, prenatal environment can affect postnatal maternal behaviour by changing their offspring's behaviour.

Providing an Outdoor Exercise Area Affects Tie-Stall Cow Reactivity and Human-Cow Relations.

Confinement and restriction of movement are a reality for most dairy cows. Providing outdoor access is one method to increase movement opportunities. However, leading cows to an outdoor exercise area increases their exposure to manipulations different from those of an indoor housing system. These situations have the potential to induce fear reactions, which can lead to injuries for the cow and danger or economic losses for the farmer. Our aim was to evaluate the development of the human-cow relationship and general reactivity of cows after a 12-week period of outdoor access provision in winter, summer and fall. A total of 16 cows in the winter, 16 in the summer, and 15 in the fall were enrolled in the study and either allocated to the treatment (Out) or stayed in the tiestall (NonOut). A human reactivity test and suddenness test were performed before and after the 12-week treatment period. In winter and to a lesser extent in fall, Out cows had a better human reaction score compared to NonOut cows, suggesting that cows with outdoor access during the winter associated human approaches with positive events. Conversely, no difference in the human reaction score was found between treatments during the summer. For summer and fall, Out cows did, however, show a decrease in their reaction score to the suddenness test compared to NonOut cows. The results of the human reactivity test in the summer suggested that cows with outdoor access did not associate the manipulation with a positive event. Interestingly, this result is not due to the cows being more frightened, since the suddenness test suggested that the Out cows were less fearful than NonOut cows. The way in which cows were led to the outdoor area could explain the differences in cow responses. Here, summer cows faced greater movement restrictions during trips to the outdoor area than in the winter, which may have been negatively perceived by the cows. We conclude that, besides the provision of outdoor access, the manner in which cows are handled during these events may have significant impacts on their reactions and could facilitate future handling.

Mother-chick interactions are affected by chicks' sex and brood composition in Japanese quail.

Sex is a trait which leads mothers to invest asymmetrically in their young, especially in species with a strong dimorphism. Authors usually consider that members of the largest sex require the most investment by mothers. In other cases, mothers care preferentially for members of the sex that increase their fitness, e.g. males, in numerous species. In quail, a precocial bird, mother seems to take more care of their males which are more sensitive to their separation. This study investigates the influence of chicks' sex and brood's sexual composition on the maternal behavior of quail. We evaluated three batches of mothers, composed of (a) unisex broods of males (MM), (b) unisex broods of females (FF), and (c) mixed sex broods (Mx). We recorded mother-chick interactions during 11 days of mothering. We found that MM mothers spent more time warming their chicks and FF mothers were more active. Moreover, females rejected more aggressively and earlier their female chicks than their male chicks. Finally, Mx chicks, whatever their sex, interacted more with their mothers. Our results highlight a greater quantity and quality of maternal care towards male. Mothers invest more in male chicks. We discussed results in terms of each sex's needs.

High social motivation induces deficits in maternal behaviour but not plasticity of the subventricular zone in Japanese quail (Coturnix japonica).

Maternal behaviour develops differently depending on the characteristics of an individual, such as age or emotional reactivity. Social motivation, defined as the propensity to establish social contact, has received little attention in relation to maternal behaviour in birds. In addition, the transition to motherhood is a time of plasticity in the brain of the new mother in mammals. However, it remains to be determined how maternal brain plasticity is affected in avian species. The present study investigated how a the social motivation of a mother alters maternal behaviour and brain plasticity of the Japanese quail (Coturnix japonica). Adult females from lines selected for high and low social motivation were exposed to chicks for 11 days. After maternal care testing, and at matched time points in controls, the brains of females were perfused for assessment of doublecortin-immunoreactive staining, a marker of neurogenesis, in the subventricular zone (SVZ), a neurogenic niche. The results obtained showed that high socially motivated female quail spent significantly less time performing maternal behaviour when exposed to chicks compared to low socially motivated females. Moreover, the warming of chicks by high socially motivated females involved less covering postures and mothers were more rejecting of chicks. Interestingly, the plasticity indicators in the SVZ did not differ between low and high socially motivated females and were not associated with differences in maternal caregiving when using doublecortin-immunoreactive staining. Thus, high social motivation in this avian species does not favour maternal behaviour and this level of motivation to the mother is not related to changes in neuroplasticity in the SVZ of the female quail.

Maternal care affects chicks' development differently according to sex in quail.

Maternal behavior is known to influence the behavioral development of young. Recently, it was demonstrated that maternal behavior also differed according to sex of chicks and brood sex composition. So, here, we explored if these factors influenced behavioral development of quail chicks when they were brooded, and what characteristics of chicks and foster females could best explain this development. We studied three sets of chick pairs brooded by foster females: unisex male, unisex female, and mixed broods. We found that both emotivity profile and sociality depended on the sex: females were more reactive and less social than males. Females' emotivity profile was correlated with brood composition and foster female activity during maternal care. In males, only sociality was correlated with foster females' scores of aggressive rejection. Our results evidence that male and female chicks respond differentially to maternal behavior. This is discussed in terms of ecological and physiological constraints on development according to sex.

Brood size can influence maternal behaviour and chick's development in precocial birds.

Mothers have a crucial influence on offspring development. Variations of maternal behaviour can be due to numerous parameters, for instance costs are related to the size of a brood/litter, which in turn can influence the level of mothers' investment in each offspring. Here we investigated the influence of brood size on the behaviour of Japanese quail mothers and chicks during the mothering period and on offspring development. We compared two types of broods: small broods of three chicks (N=9) and large broods of six chicks (N=9). Behavioural tests assessed chicks' social and emotional traits. Mothers of large broods emitted more maternal vocalisations at the beginning of the mothering period, but at the end they assumed more non-covering postures and trampled chicks more than mothers of small broods. Chicks in large broods huddled up more whereas chicks in small broods rested alone more frequently. Moreover, the social motivation of chicks in large broods was higher than that of chicks in small broods, although their emotional reactivity levels were similar. Our results evidence the importance of brood size for maintaining family cohesion and the influence of brood size on chicks' interactions with their siblings. We evaluated the influence of mothers and siblings on chicks' behavioural development.

Maternal diet influences offspring feeding behavior and fearfulness in the precocial chicken.

BACKGROUND: In chicken, oils in the maternal diet confer a specific scent to the yolk. Embryos are known to perceive and memorize chemosensory signals of the surrounding environment; however, the potential impact of the maternal diet has not previously been investigated. In the present study, we hypothesized that chicken embryos memorize the chemical signals of the maternal diet and that this perceptual learning may orient subsequent feeding behavior of the hatchlings. METHODOLOGY/PRINCIPAL FINDINGS: Laying hens were fed standard food enriched with 2% menhaden oil (MH group) or 2% soybean oil (controls). The scent of menhaden was significantly more detected in MH egg yolks than in control yolks by a human panel. We analyzed the development and behavior of offspring towards different types of food, bearing or not bearing the menhaden scent. When chicks were exposed to a 3-min choice test between the familiar food bearing the menhaden scent and the familiar food without menhaden, no effect of treatment was observed. In a 3-min choice test with unfamiliar food (mashed cereals) MH chicks showed a clear positive orientation toward the unfamiliar food bearing the menhaden scent. By contrast, control chicks showed a preference for the non-odorized unfamiliar food. MH chicks expressed higher emotional reactivity level than control chicks as expressed by food neophobia and longer immobility in a restraint test. CONCLUSION/SIGNIFICANCE: Chicks exposed in ovo to menhaden oil via the maternal diet preferentially oriented their feeding behavior towards food containing menhaden oil, but only when the food was unfamiliar. We propose that oil in the maternal diet engenders maternal effects and contributes to the development of behavioral phenotype in the offspring. In ovo chemosensory learning may have evolved to prepare precocial offspring for their environment. This suggests a common principle of embryonic chemosensory learning across vertebrate taxa.