Effect of mothering on the spatial exploratory behavior of quail chicks.

Previous maternal deprivation experiments demonstrated that absence of maternal care impacts the behavioral development of young animals. Here we assessed the influence of the presence of a mothering hen on the spatial exploration of Japanese quail chicks, after the mothering period. Brooded and nonbrooded chicks were placed in a novel environment containing feeding troughs. The distribution of chicks and their inter-individual distances were measured during repeated tests. Brooded chicks exhibited a higher ability to disperse, thereby progressively exploiting larger surfaces and gaining access to food more easily. The fact that exploration by nonbrooded chicks was delayed suggests a deficit in their exploratory motivation and/or spatial skills. We hypothesize that brooded chicks experienced the constraint to follow the mothering hen, and to adapt to frequent reconfigurations of their environment. The lack of this variability in the environment of nonbrooded chicks could have reduced adaptability of their spatial behavior.

Higher inherent fearfulness potentiates the effects of chronic stress in the Japanese quail.

There is considerable variability in the susceptibility of individuals to the adverse effects of chronic stress. In humans and other mammals, individual traits such as high anxiety are proposed as a vulnerability factor for the development of stress-related disorders. In the present study, we tested whether a similar behavioural trait in birds, higher emotional reactivity, also favours the occurrence of chronic stress-related behavioural and physiological dysfunction. For this, lines of Japanese quail divergently selected for a typical fear response in birds, the duration of tonic immobility, were subjected to unpredictable aversive stimulation over 2 weeks. Previous studies demonstrate that the selection program modifies the general underlying emotionality of the birds rather than exerting its effect only on tonic immobility. Interestingly, only birds selected for their higher emotionality exhibited significantly enhanced latency to first step and decreased locomotor activity in the open-field test after exposure to chronic stress compared to non-stressed control birds. This effect of chronic stress was selective for the tested dimension of bird emotional reactivity because there was no observed effect on the tonic immobility response. Moreover, chronically stressed birds selected for their higher emotionality exhibited significantly decreased basal corticosterone levels, a physiological marker of stress. These findings show that chronic stress is associated with changes in emotional reactivity and related physiological markers in birds. They also highlight emotional reactivity as an important predisposing factor for the occurrence of the adverse effects of chronic stress in birds.

Foraging opportunity: a crucial criterion for horse welfare?

This study aimed at determining the effect of the increase of foraging opportunities on the behaviour and welfare of breeding mares housed in individual boxes but allowed outside 6 h a day in a bare paddock. One hundred Arab breeding mares were divided into two groups of 50 according to the treatment and allowed outside in two bare paddocks at the same density (115 mare/ha) where water and shelter were provided. The treatment consisted in providing the opportunity to forage on hay. Twenty-minute animal focal samplings and scan samplings were used to determine the time budget of the mares during the period from 0900 to 1500 h and study their social behaviour. A total of 300 focal sampling (6000 min), 3300 individual scan sampling (6000 min) and 62 group observations (1240 min) corresponding to the 100 mares were recorded. Non-parametric tests were used to analyse data. Results showed that experimental mares spent more time feeding (65.12% ± 2.40% v. 29.75% ± 2.45%, P < 0.01) and less time in locomotion (11.70% ± 1.31% v. 23.56% ± 1.34%, P < 0.01), stand resting (11.76% ± 2.57% v. 27.52% ± 2.62%, P < 0.01) and alert standing (5.23% ± 1.2% v. 14.71% ± 1.23%, P < 0.01). There was more bonding among experimental mares than control ones (26 v. 14, P < 0.05). Experimental mares showed more positive social interactions (P < 0.01) and less aggression (P < 0.01). These results suggest that giving densely housed mares foraging opportunities improves their welfare.

Are 6-month-old foals sensitive to dam's influence?

A recent study has shown that gently handling dams in front of their few days old foals may strongly influence the development of human-foal relationships. In the present study, we test whether 6-month-old foals remain sensitive to their dams' influence. The study was performed on 16 foal-mare dyads, with half of the mares receiving positive contacts from the experimenter in presence of their 6-month-old foals (n = 8) whereas the other mares were not handled (n = 8). All foals were tested 15 and 30-35 days later under various conditions (reaction to a motionless human, approach test, saddle-pad tolerance test). We observe a positive effect of mare' handling on foals' reactions to humans but with a high interindividual variability, suggesting a higher effect of the foals' own behavioral characteristics at this age than at earlier stages.

Influence of various early human-foal interferences on subsequent human-foal relationship.

Whereas the way animals perceive human contact has been particularly examined in pet animals, a small amount of investigations has been done in domestic ungulates. It was nevertheless assumed that, as pet animals, non-aggressive forms of tactile contact were as well rewarding or positive for these species, even though the features of intraspecific relationships in pet animals and domestic ungulates may be to some extent different. We test here the hypothesis that horses may not consider physical handling by humans as a positive event. When comparing different early human-foal interactions, we found that early exposure to a motionless human enhanced slightly foals reactions to humans whereas forced stroking or handling in early life did not improve later human-foal relation. Foals that were assisted during their first suckling (e.g., brought to the dam's teat) even tended to avoid human approach at 2 weeks, and physical contact at 1 month of age. We argue that interspecies differences may exist in how tactile stimulation is perceived. It may be important for the establishment of a bond that a young animal is active in the process and able, through its behavioral responses, to help define what is positive for it. This way of investigation may have important general implications in how we consider the development of social relations, both within and between species.

Mutual influence of the maternal hen's food calling and feeding behavior on the behavior of her chicks.

It is widely accepted that brooding hens attract their chicks to food by food calling, but until now, the concurrent behavior of hens and chicks has never been investigated in depth. This study provides a detailed analysis of both the behavior of brooded chicks and the distance to their mother in relation to her feeding sequences, and whether they contained food calling. Our results revealed that brooding hens utter food calls while pecking, especially when their chicks are not feeding and/or have been at some distance for several seconds. Chicks' response to their mother's feeding activities was more pronounced in the presence than in the absence of food call. Chicks responded to this call by approaching their mother and increasing their pecking; their response became more efficient as they grew older. We thus consider food calls as an arousal vocalization that directs the chicks' attention to a food item chosen by a hen.

Experience modulates emission of food calls in broody hens.

The aim of these experiments was to determine if previous experience of chicks' response to food calling influences subsequent propension of maternal hens to utter food calls. Seventeen broody hens were tested three times a day without their 3- or 4-day-old chicks. Hens were tested in two situations: chicks were returned either after each test or at the end of all the day's tests. As palatability influences food calling in maternal hens, experiments were conducted first with a highly preferred food item and then with the hens' usual feed. The chicks' capacity to respond regularly to their mother influences the hens' capacity to emit food calls. In fact, although the hens did not lose their maternal state, they uttered fewer food calls when their chicks were removed all day. These results suggest that the chicks' behaviour following food calling could be a social reinforcement for broody hens.

Behaviour and hormone concentrations in nest deprived and renesting hens.

1. Nest and egg deprivation is a procedure traditionally used to disrupt incubation behaviour expression in commercial flocks. The aim of this study was to establish how nest deprivation affects the subsequent changes in behaviour and hormone secretion and whether readiness to renest is related to hormone concentration before and during nest deprivation. 2. Incubating broiler breeder hens were deprived of their nest either in their familiar environment, by blocking the nest entrance in the home cage or in an unfamiliar environment, by transfer in a different cage. After 3 days of nest deprivation, next access was allowed and readiness to incubate tested. 3. Both methods of nest deprivation resulted in the expression of similar behaviours associated with the disruption of incubation, and similar increases in plasma LH and oestradiol and decreases in plasma prolactin. The percentage (approximately 70%) of hens which renested after either method of nest deprivation did not differ significantly. 4. Readiness to renest was not related to the concentrations of plasma prolactin measured before or during the 3 days of nest deprivation. However, hens which would later renest could be identified by using behavioural criteria. They emitted more avoidance trills before, and sat on the wire floor for longer periods during, the nest deprivation period than the others.

Influence of stimuli from chicks on behavior and concentrations of plasma prolactin and luteinizing hormone in incubating hens.

Removal of eggs from the nests of incubating birds or substitution of eggs for chicks disrupts incubation behavior and induces changes in the secretion of prolactin and luteinizing hormone (LH). The aim of the present study was to determine how different stimuli, such as physical contact with eggs and tactile, visual, and/or auditory cues from chicks, interact to control the transition between incubation and brooding and to induce changes in prolactin and LH plasma concentrations. Physical contact with chicks, in the presence or absence of eggs, induced brooding behavior and an immediate fall in plasma prolactin concentration and a gradual increase in LH concentration. Vocalizations, particularly clucking and food calls, increased rapidly while incubation and nest attachment disappeared slowly. No change in plasma prolactin or LH concentration was observed in incubating hens which could hear and see or only hear chicks. These incubating hens showed no interest in chicks and continued to incubate persistently. To conclude, tactile stimuli, alone or in combination with visual and/or auditory stimuli from newly hatched chicks, are the only cues that induce the transition from incubation to brooding and the associated decrease in prolactin secretion and increase in LH secretion.

Effects of environmental and social factors on incubation behavior, endocrinological parameters, and production traits in turkey hens (Meleagris gallopavo).

Hens raised in three different environments were assessed for changes in egg production performance, the rate of incubation behavior expression, and plasma levels of luteinizing hormone (LH) and prolactin for 20 wk following the laying of the first egg. The environments were individual (IFP) or collective floor pens (CFP) and individual battery cages (Cp and Cnp). The hens from three experimental groups (IFP, CFP, and Cp) were transferred from a short (6 h) to a long (14 h) photoperiod, whereas the ones from the remnant (Cnp) were left under a short one. Increase of the photoperiod induced significant increases (P < 0.05) in levels of prolactin and LH after 1 d, and resulted in the onset of egg laying in a delay of 14 d in all groups. However, the overall egg laying performance was highest for the IFP hens. The CFP and IFP hens laid 98 and 24% of their eggs inside the nest boxes, respectively. The hens raised in battery cages did not express incubation behavior, whereas 50 and 33%, respectively, of the CFP and IFP hens did. During the 1st wk of egg laying, levels of prolactin increased for all photostimulated hens but to a greater extent for CFP hens. Higher increases in levels of prolactin were associated with the expression of incubation behavior; however, prolactin levels of nonincubating laying hens were also higher under the CFP treatment. It appears that the rate of expression of incubation behavior, as well as changes in the plasma levels of prolactin and LH throughout an egg production period, are dependent upon rearing conditions in turkey hens.

The effect of ovariectomy on concentrations of plasma prolactin and LH and parental behavior in the domestic fowl.

The role of ovarian hormones in the expression of parental behavior and in the regulation of LH secretion was investigated in incubating commercial meat-type hens. After ovariectomy, incubating hens continued to incubate eggs normally and brooded day-old chicks given to replace eggs, in a manner similar to sham-ovariectomized control hens. The concentration of plasma LH increased significantly in incubating hens after ovariectomy while the concentration of plasma prolactin remained high. Plasma LH remained depressed in sham-ovariectomized incubating control hens. The increase in plasma LH in incubating hens after ovariectomy (3.92 +/- 0.7 ng/ml) was less than that following the ovariectomy of nonincubating, nonlaying hens (5.3 +/- 1.2 ng/ml). The two groups of hens differed in that plasma prolactin concentrations were high (527 +/- 7.4 ng/ml) in the incubating hens and low (70 +/- 9 ng/ml) in the nonincubating hens. Nest deprivation resulted in an increase in plasma LH in both ovariectomized and sham-ovariectomized incubating hens with a significantly larger increase occurring in the ovariectomized hens (8.5 +/- 1.41 ng/ml compared to 2.48 +/- 0.65 ng/ml). Nest deprivation resulted in a similar rapid decrease in plasma prolactin in both ovariectomized and sham-ovariectomized hens. Replacement of eggs with day-old chicks in ovariectomized or sham-ovariectomized incubating hens resulted in a rapid decrease in plasma prolactin and after 6 days, in an increase in plasma LH in the ovariectomized but not sham-operated hens. It is concluded that once incubation behavior is established, ovarian hormones are not required for its maintenance or the readiness to brood day-old chicks. Ovarian hormones do, however, suppress LH release during incubation while the high concentration of plasma prolactin supplements this suppression.

Influence of the presence of chicks on the ability to resume incubation behavior in domestic hens (Gallus domesticus).

In this study, hormonal and behavioral changes associated with nest deprivation in the absence or in the presence of chicks (replaced every day or not) were investigated in incubating hens. Prolactin levels decreased, whereas LH and E2 levels increased following nest deprivation. Surprisingly, the presence of chicks had no effect on prolactin changes. However, the presence of chicks significantly limited increases in plasma LH and estradiol. Furthermore, this inhibitory effect was greater when day-old chicks were replaced every day. On the other hand, the presence of chicks induced the emergence of specific maternal behavior, while, surprisingly, one-third of the nest-deprived hens without chicks continued to incubate. After a 3-day deprivation period, readiness to renest was maintained for a percentage of hens. Thus more than 2/3 of hens deprived of the nest for 3 days in the presence of chicks returned to their nests during the first day of being allowed to do so, versus less than 1/3 of hens without chicks among those that previously disrupted incubation behavior. Plasma prolactin concentrations of these renesting hens increased rapidly and reached again levels characteristic of incubating hens. We conclude that, under our experimental conditions, presence of chicks maintains readiness to incubate without maintaining high levels of plasma prolactin.

Respective effects of chicks and nest on behavior and hormonal concentrations of incubating domestic hens.

Removal of incubating hens from their nestboxes or substitution of chicks for eggs disrupt incubation; prolactin levels decrease whereas luteinizing hormone and gonadal steroid concentrations tend to rise. The present experiment was undertaken to determine the relative influence of removing the nest (nest-deprived hens), adding chicks (maternal hens), or both (maternal nest-deprived hens), on the behavior and hormone concentrations of incubating hens. The results confirm that nest removal, as well as adding chicks, stops incubation. No differences were found between maternal and maternal nest-deprived hens' behavioral responses or hormonal concentrations. These results do not support the hypothesis that the drop of plasma prolactin observed in maternal hens is the consequence of the nest abandonment. In contrast, nest-deprived hens presented lower concentrations of prolactin and higher concentrations of estradiol than maternal and maternal nest-deprived hens. Moreover, the nest-deprived hens presented lower concentrations of prolactin and higher concentrations of LH and estradiol than the hens given chicks. We assume that physical contact with chicks, during brooding bouts, slows down the decrease of prolactin secretion and inhibits LH and estradiol release at the end of incubation.

Incubation and maternal behaviour in domestic hens: influence of the presence of chicks on circulating luteinising hormone, prolactin and oestradiol and on behaviour.

1. The consequences of the adoption of chicks and their subsequent removal on behaviour and plasma hormone concentrations of incubating hens were investigated. Birds were divided into two group: in group A, incubating hens were given chicks for 11 d; in group B chicks were left with the hens for 3 d only. 2. Incubating hens given chicks immediately showed maternal responses. The introduction of chicks induced a gradual nest desertion. Their removal stopped nest desertion temporarily on day 4 in group B hens. 3. Plasma prolactin concentrations fell one day after introduction of chicks and continued to decline for about one week in group A hens, although there was no further significant decrease in group B hens. Circulating prolactin tended to decrease with time in both groups. 4. Plasma luteinising hormone (LH) concentrations increased concurrently with the decrease of prolactin. The increase was more abrupt in group B hens. 5. Plasma oestradiol concentrations decreased slightly on the day chicks were introduced. The decline was arrested by removal of chicks in group B; in group A the tendency was reversed about 10 days after chick introduction. 6. Irrespective of group, before chick removal hens which deserted their nest rapidly had less contact with chicks and lower prolactin concentrations.

Tendency to display spontaneous incubation does not affect maternal responsiveness in the domestic hen.

Induction of maternal behavior was performed in 63-week-old hens of the JV15 (or "Vedette") strain which had previously exhibited brief incubating phases during the preceding egg-laying period to discover if their maternal behavior differed from that induced in hens who had no previous record of incubating. The degree of maternal behavior exhibited showed large variation among individuals. However, hens with a history of one or more spontaneous, brief incubation bouts and hens without such histories showed similar responsiveness toward chicks and similar levels of general activity. In most of the subjects, the establishment of maternal behavior was accompanied by a significant long-term fall in the plasma levels of luteinizing hormone whereas the plasma levels of prolactin appeared to be unaffected.

Influence of food restriction and of the presence of chicks on the reproductive system of the domestic hen.

An experiment was conducted with 4 groups of hens: a control group of laying hens, a group subjected to food deprivation for 7 d, a group subjected to food deprivation for 7 d, then re-fed for 10 d and a group subjected to food deprivation for 7 d then re-fed for 10 d with two chicks per hen introduced during the last 7 d of refeeding. Food deprivation provoked the involution of the oviduct and ovarian regression, as well as a decrease in the plasma concentrations of luteinising hormone (LH) and sex steroids (progesterone, testosterone and oestradiol). After 3 d of re-feeding, there was a significant increase in the plasma concentrations of LH and steroid hormones. This phenomenon was even more marked after 10 d of re-feeding; most of the hens of the third group which were not given chicks, were at the point of lay. The presence of chicks resulted in the expression of maternal behaviour and suppressed a rapid return to laying. This was especially marked in hens showing typical maternal behaviour traits, for which the morphological and endocrine measurements indicated a decrease in the activity of the hypothalamo-hypophyseal-gonadal axis.

Induction of maternal behavior in incubating and non-incubating hens: influence of hormones.

Maternal responses and variations in plasma levels of prolactin and testosterone have been studied in incubating and in non-incubating, non-laying hens during forced adoption experiments. The results demonstrate the ability of incubating hens to display complete maternal behavior as early as the 10th day of incubation after being exposed to stimulation by chicks during one night. Maternal responses also emerged in non-laying hens but more gradually. In both groups, a decline in plasma testosterone occurred after the introduction of the chicks and, in the incubating hens, prolactin levels fell as they abandoned their nests.

Responses to successive test of induction of maternal behaviour in hens.

This experiment was undertaken to find out if the kinetics of the behaviours which emerge gradually during a first experiment of maternal induction was modified by a second and then a third similar experiment, or not. In addition, during the second experiment the behaviour of experienced hens was compared to that of naive hens of the same strain undergoing their first induction. Three important facts appeared : (i) In each series, maternal behaviours (accepting contact of chicks, clucking) emerged gradually (significant day effect). (ii) Repeating these experiments revealed on the whole neither increase nor decline of hens' responsiveness towards chicks (negligible series effect). No significant difference between naive hens and hens having already had a first induction experience was found. (iii) There was an important difference between hens concerning their delay in adopting, and a good stability of individual profiles was found between series (significant hen effect).

[Induced maternal behavior in the domestic hen. Influence of partial or total separation on the maintenance of maternal responsiveness]. / Comportement maternel induit chez la poule domestique. Influence d'une séparation partielle ou totale sur le maintien de la réactivité maternelle.

The influence of direct physical contact on maintaining maternal responsiveness in induced adult hens was investigated using a separation procedure. Partial separation from chicks causes a significant decline of the clucking rate in hens, this response however does not disappear as in the case of total separation. The possibility for physical contact between hen and chicks contributes largely, but not uniquely, to maintain maternal responsiveness in the domestic hen.

Induction of maternal behavior and some hormonal and physiological correlates in the domestic hen.

The manifestation of maternal behavior and the variations of plasmatic levels of gonadic hormones (testosterone, delta 4-androstenedione, and 17 beta-estradiol) have been studied in two groups of domestic hens (layers and nonlayers) during a forced adoption experiment. Maternal behavior appears later in actively laying hens than in nonlayers. The former hens show a higher level of androgens (T and delta 4). The appearance of typical maternal behavior coincides with a pause in laying and a decrease in all the plasmatic steroid levels measured.