Paul Constantin

Subdivisions of the arcopallium/posterior pallial amygdala complex are differentially involved in the control of fear behaviour in the Japanese quail

Growing evidence suggests that the arcopallium/posterior pallial amygdala plays a major role in the control of fear behaviour in birds. This brain region comprises several subdivisions, but no direct evidence is available about its functional parcellation. The aim of the present study was to investigate the relative involvement of two subdivisions of the arcopallium/posterior pallial amygdala complex in four classical tests of fear in quail: the presentation of a novel object, the hole-in-the-wall, open-field and tonic immobility tests. Bilateral electrolytic lesions damaging the posterior part of the arcopallium/posterior pallial amygdala resulted in an increase in fear behaviour in the open-field test, whereas quail with lesions damaging the anterior part of the arcopallium displayed a decrease in an overall fear score, compared to quail with bilateral nidopallium or sham lesions. The differential involvement of the anterior and posterior parts of the arcopallium/posterior pallial amygdala in fear behaviour is discussed in view of the known connections between the arcopallium/posterior pallial amygdala complex and brain regions considered to be limbic in nature.

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.

Effect of one week of stress on emotional reactivity and learning and memory performances in Japanese quail

Chronic stress is known to induce long term alterations of emotional behaviours as well as cognitive performances leading thereby to welfare or husbandry problems. These stress-induced consequences are observed following long periods of stress lasting from several weeks to several years. The current study examined whether a shorter period of stress (one week) produced similar impairing effects. Two-week old Japanese quail were either submitted to a series of aversive events over consecutive 8 days, at unpredictable times each day (treated animals) or left undisturbed (controls). Following the treatment period, animals were weighed and basal as well as aversive events-induced levels of plasma corticosterone were quantified. Quail were also tested for emotional reactivity in three tests (the tonic immobility test, the hole-in-the-wall and novel object tests) and for spatial reference memory. Although there was no difference in corticosterone levels between the two groups, the treated animals had lower body weight than controls. Behavioural investigations after the treatment period did not reveal any difference between the groups in the three emotional reactivity tests. In the spatial task, treated quail displayed enhanced behavioural flexibility as revealed by their higher performances during the reversal phase of the task. The alteration of growth suggests that a short period of repetitive exposures to unpredictable aversive events can be perceived by quail as stressful. Such a stress period can improve spatial learning performances in quail supporting the critical role played by the duration of the stress period on cognitive performance.

Changes in Heart Rate Variability during a tonic immobility test in quail

Tonic immobility (TI) is an unlearned fear response induced by a brief physical restraint and characterized by a marked autonomic nervous system involvement. This experiment aimed at studying the relative involvement of both autonomic sub-systems, the sympathetic and parasympathetic nervous systems, during TI, by analyzing Heart Rate Variability. Quail selected genetically for long (LTI) or short (STI) TI duration and quail from a control line (CTI) were used. The animals were surgically fitted with a telemetric device to record electrocardiograms before and during a TI test. Heart rate did not differ between lines at rest. The induction of TI, whether effective or not, induced an increase in HR characterized by a shift of the sympathovagal balance towards a higher sympathetic dominance. Parasympathetic activity was lower during effective than during non-effective inductions in CTI quail. During TI, the increase in sympathetic dominance was initially maintained and then declined, while relative parasympathetic activity remained low, especially in CTI and STI lines. The end of tonic immobility was characterized by a rise in overall autonomic activity in all lines and an increase in parasympathetic influence in CTI and STI quail. To conclude, the susceptibility to TI cannot be explained only by autonomic reflex changes. It is probably strongly related to the perception of the test by the quail. During TI, the differences between lines in autonomic responses probably reflect behavioural differences in the fear response.

Genetic selection for fast growth generates bone architecture characterised by enhanced periosteal expansion and limited consolidation of the cortices but a diminution in the early responses to mechanical loading

Bone strength is, in part, dependent on a mechanical input that regulates the (re)modelling of skeletal elements to an appropriate size and architecture to resist fracture during habitual use. The rate of longitudinal bone growth in juveniles can also affect fracture incidence in adulthood, suggesting an influence of growth rate on later bone quality. We have compared the effects of fast and slow growth on bone strength and architecture in the tibiotarsi of embryonic and juvenile birds. The loading-related biochemical responses (intracellular G6PD activity and NO release) to mechanical load were also determined. Further, we have analysed the proliferation and differentiation characteristics of primary tibiotarsal osteoblasts from fast and slow-growing strains. We found that bones from chicks with divergent growth rates display equal resistance to applied loads, but weight-correction revealed that the bones from juvenile fast growth birds are weaker, with reduced stiffness and lower resistance to fracture. Primary osteoblasts from slow-growing juvenile birds proliferated more rapidly and had lower alkaline phosphatase activity. Bones from fast-growing embryonic chicks display rapid radial expansion and incomplete osteonal infilling but, importantly, lack mechanical responsiveness. These findings are further evidence that the ability to respond to mechanical inputs is crucial to adapt skeletal architecture to generate a functionally appropriate bone structure and that fast embryonic and juvenile growth rates may predispose bone to particular architectures with increased fragility in the adult.

Emotional reactivity modulates autonomic responses to an acoustic challenge in quail.

Emotional reactivity modulates autonomic responses to an acoustic challenge in quail. Physio Behav 00(0) 000-000, 2006. This study investigated the relationship between emotional reactivity and behavioral and autonomic responses to an acoustic stimulus in quail. It was hypothesized that birds with high emotional reactivity would have higher motor inhibition combined with higher sympathetic activation than birds with low emotional reactivity. Two experiments were performed. The first looked for correlations between emotional reactivity, evaluated by a tonic immobility test, and motor and Heart Rate Variability in relation to an acoustic stimulus. The second experiment compared the motor and autonomic responses to the acoustic stimulus of quail selected on either long (LTI) or short (STI) duration of tonic immobility. The first experiment showed that the acoustic stimulation induced motor inhibition and cardiac activation. Correlations were found between tonic immobility duration and both autonomic activity before stimulation and sympathovagal balance after stimulation. In the second experiment, LTI quail showed strong sympathetic activation, whereas STI quail showed parasympathetic and sympathetic activation. The activation of the parasympathetic system induced by the noise in STI quail can be explained by the predominance of this system at rest in this line. In conclusion, both the basal autonomic activity and the autonomic responses differed according to the emotional reactivity, and changes in autonomic activity appear to be related to the genetic selection process.

Emotionality Modulates the Effect of Chronic Stress on Feeding Behaviour in Birds

Chronic stress is a long-lasting negative emotional state that induces negative consequences on animals’ psycho-physiological state. This study aimed at assessing whether unpredictable and repeated negative stimuli (URNS) influence feeding behaviour in quail. Sixty-four quail were exposed to URNS from day 17 to 40, while 64 quail were undisturbed. Two lines divergently selected on their inherent emotionality were used to assess the effect of genetic factors on the sensitivity to URNS. All quail were submitted to a sequential feeding procedure (using two diets of different energetic values) which placed them in a contrasting situation. Behavioural tests were performed to assess the emotional reactivity of the two lines. Results confirmed that differences exist between them and that their emotional reactivity was enhanced by URNS. Diet preferences, motivation and daily intake were also measured. URNS did not change the preferences for the hypercaloric diet compared to the hypocaloric diet in choice tests, but they reduced daily intakes in both lines. Motivations for each diet were differently affected by URNS: they decreased the motivation to eat the hypercaloric diet in quail selected for their low inherent fearfulness whereas they increased the motivation to eat the hypocaloric diet in quail selected for their high inherent fearfulness, which suggested a devaluation process in the former and a compensatory behaviour in the later. Growth was furthermore reduced and laying delayed by URNS in both lines. In conclusion, the exposure to URNS induced interesting changes in feeding behaviour added with an increase in emotional reactivity and an alteration of production parameters. This confirms that both lines of quail experienced a chronic stress state. However differences in feed motivation and emotional reactivity between lines under chronic stress suggested that they experienced different emotional state and use different ways to cope with it depending on their genetic background.

A higher inherent trait for fearfulness is associated with increased anxiety-like behaviours and diazepam sensitivity in Japanese quail

This study tested whether lines of Japanese quails divergently selected for a fear response, the tonic immobility, might constitute a reliable bird model for studying anxiety. Previous studies demonstrated that the selection modifies the general underlying emotionality of the birds rather than exerting its effect only on tonic immobility. The behavioural effects of intraperitoneal injections of diazepam, an anxiolytic drug, were assessed in two lines of quail selected either for their short (STI) or long (LTI) duration of tonic immobility. Effects of diazepam were examined in two tests used for measuring emotionality in birds, the open field and the tonic immobility tests. After being placed in the centre of the open field, birds with a high emotionality (LTI quails) stayed longer in the centre of the apparatus than STI quail. Diazepam had anxiolytic effect in LTI birds as it increased the time spent in the outer area. This effect of diazepam appears to be selective because the drug has no effect on other behaviours such as distress calls or escape attempts. The drug has also no effect on the tonic immobility response in any of the two lines. These findings reveal an anxiogenic trait of LTI birds in the open field test that can be modulated by the administration of an anxiolytic drug. Therefore quails selected for LTI and STI represent a valuable model to study the mechanisms underlying anxiety in birds.

Impact of high and low anxiety trait on object habituation and discrimination: Evidence from selected lines of Japanese quail

Compared to rodents, the relationship between anxiety and cognitive performances has been less studied in birds. Yet, birds are frequently exposed to stimulations that constitute a potential source of anxiety and can affect their adaptation to their living conditions. The present study was aimed at evaluating, in birds, the relationship between levels of anxiety and object habituation and discrimination with the use of Japanese quail lines divergently selected for a fear response, tonic immobility. Previous studies demonstrated that the selection programme has modified the general anxiety trait of the birds. The task consisted in 4 daily sessions of 8 successive presentations of the same object in the home cage of the quail in order to habituate each bird to the object. The observation that both quail with a high and a low anxiety trait progressively spent more time close to the object indicated that habituation occurred. Dishabituation was assessed during a single session of 8 presentations of a novel object. Only quail with a high anxiety trait exhibited significant discrimination. They spent significantly less time close to the novel object than to the habituated object. It is hypothesised that a high anxiety trait is associated with a more accurate processing of environmental cues or events resulting in better discriminative performances.

A trait for a high emotionality favors spatial memory to the detriment of cue-based memory in Japanese quail

Recent studies provided evidence that a personality trait such as a trait for a high or a low emotionality can either promotes or impairs learning and memory performances. This variability can be partly explained because this trait may have opposite effect on memory performances depending on the memory system involved. The present study investigated in Japanese quail the relationships between emotionality and two forms of memory, spatial- memory and cue-based memory. We showed that birds with a high emotionality trait (E+), compared with birds with a low emotionality trait (E-), reached slowly but more accurately the spatial location of a rewarded cup in an arena that contains 8 identical cups. Then a second cohort of Eu2009+u2009and E- quails was trained to solve a dual spatial/cued task in which they could use either their spatial or cue-based memory. Whereas Eu2009+u2009birds predominantly solved the task using their spatial memory, E- birds preferentially used their cue-based memory. These findings demonstrate that a trait for a high emotionality, can influence spatial memory performances but also contributes to favor the selection of this form of memory.