Brazilian tegus (Tupinambis merianae) were instrumented with telemetry encoder implants that measured and broadcast heart rate (HR), breathing rate (fR), and deep body temperature (Tb) and were then allowed to freely roam in outdoor enclosures mimicking their natural environment for a full year (2004) in order to monitor the circadian and circannual patterns in behaviour and cardio-respiratory physiology. The year was divided up into 5 seasons based upon the physiology and behaviour of the tegus: early activity season (Sept.-Nov.), late activity season (Dec.-Feb.), entrance into hibernation (March-April), hibernation (May-July), and arousal from hibernation (August). The activity seasons were characterized by warm weather with frequent rainfall which slowly decreased in temperature and precipitation as tegus started entering hibernation so that the end of the dormant season was marked by dry, cold weather. Tegus in the early activity season demonstrated high activity associated with breeding demands, displayed elevated HR and fR, and were able to maintain a large temperature differential (4-7°C) between deep body temperature (Tb) and their respective burrow (Tburrow) during sleep. As the season progressed into late activity season, average Tb remained constant but average HR and fR progressively declined indicating nightly torpor. Periods of inactivity during the active seasons were rare and associated with inclement weather. Tegus entered hibernation through bouts of inactivity that progressively increased in frequency and duration. During this time, Tb was regulated but declined at different rates in regards to daytime and nighttime values. Heart rate through the entrance into hibernation and hibernation periods frequently demonstrated arrhythmias that increased in duration but decreased with frequency as hibernation progressed. Through hibernation, Tb continued to decline for the first month but HR and fR were constant, demonstrating a temperature independent suppression of metabolism. Through the hibernation season tegus sporadically aroused and emerged from their burrows to warm up and after a short basking period would return to the burrows and swiftly resume hibernation. While hibernating, heart rate was characteristically regular but breathing was sporadic orepisodic. Arousals became more frequent towards the end of hibernation so that when they entered arousal from hibernation season most tegus were emerging daily. At this time daily maximum deep body temperature (Tbmax) swiftly returned to active season values but nighttime daily minimum deep body temperature (Tbmirt) values only showed a gradual increase through August, indicating different body temperature set points (Tbset) for active and sleep states. Changes in heart rate and breathing rate during the year showed greatest correlation with changes in photoperiod, although throughout hibernation HR and fR also showed tight correlation with Tb.

Brazilian tegus (Tupinambis merianae) were instrumented with telemetry encoder implants that measured and broadcast heart rate (HR), breathing rate (fR), and deep body temperature (Tb) and were then allowed to freely roam in outdoor enclosures mimicking their natural environment for a full year (2004) in order to monitor the circadian and circannual patterns in behaviour and cardio-respiratory physiology. The year was divided up into 5 seasons based upon the physiology and behaviour of the tegus: early activity season (Sept.-Nov.), late activity season (Dec.-Feb.), entrance into hibernation (March-April), hibernation (May-July), and arousal from hibernation (August). The activity seasons were characterized by warm weather with frequent rainfall which slowly decreased in temperature and precipitation as tegus started entering hibernation so that the end of the dormant season was marked by dry, cold weather. Tegus in the early activity season demonstrated high activity associated with breeding demands, displayed elevated HR and fR, and were able to maintain a large temperature differential (4-7°C) between deep body temperature (Tb) and their respective burrow (Tburrow) during sleep. As the season progressed into late activity season, average Tb remained constant but average HR and fR progressively declined indicating nightly torpor. Periods of inactivity during the active seasons were rare and associated with inclement weather. Tegus entered hibernation through bouts of inactivity that progressively increased in frequency and duration. During this time, Tb was regulated but declined at different rates in regards to daytime and nighttime values. Heart rate through the entrance into hibernation and hibernation periods frequently demonstrated arrhythmias that increased in duration but decreased with frequency as hibernation progressed. Through hibernation, Tb continued to decline for the first month but HR and fR were constant, demonstrating a temperature independent suppression of metabolism. Through the hibernation season tegus sporadically aroused and emerged from their burrows to warm up and after a short basking period would return to the burrows and swiftly resume hibernation. While hibernating, heart rate was characteristically regular but breathing was sporadic orepisodic. Arousals became more frequent towards the end of hibernation so that when they entered arousal from hibernation season most tegus were emerging daily. At this time daily maximum deep body temperature (Tbmax) swiftly returned to active season values but nighttime daily minimum deep body temperature (Tbmirt) values only showed a gradual increase through August, indicating different body temperature set points (Tbset) for active and sleep states. Changes in heart rate and breathing rate during the year showed greatest correlation with changes in photoperiod, although throughout hibernation HR and fR also showed tight correlation with Tb.

Brazilian tegus (Tupinambis merianae) were instrumented with telemetry encoder implants that measured and broadcast heart rate (HR), breathing rate (fR), and deep body temperature (Tb) and were then allowed to freely roam in outdoor enclosures mimicking their natural environment for a full year (2004) in order to monitor the circadian and circannual patterns in behaviour and cardio-respiratory physiology. The year was divided up into 5 seasons based upon the physiology and behaviour of the tegus: early activity season (Sept.-Nov.), late activity season (Dec.-Feb.), entrance into hibernation (March-April), hibernation (May-July), and arousal from hibernation (August). The activity seasons were characterized by warm weather with frequent rainfall which slowly decreased in temperature and precipitation as tegus started entering hibernation so that the end of the dormant season was marked by dry, cold weather. Tegus in the early activity season demonstrated high activity associated with breeding demands, displayed elevated HR and fR, and were able to maintain a large temperature differential (4-7°C) between deep body temperature (Tb) and their respective burrow (Tburrow) during sleep. As the season progressed into late activity season, average Tb remained constant but average HR and fR progressively declined indicating nightly torpor. Periods of inactivity during the active seasons were rare and associated with inclement weather. Tegus entered hibernation through bouts of inactivity that progressively increased in frequency and duration. During this time, Tb was regulated but declined at different rates in regards to daytime and nighttime values. Heart rate through the entrance into hibernation and hibernation periods frequently demonstrated arrhythmias that increased in duration but decreased with frequency as hibernation progressed. Through hibernation, Tb continued to decline for the first month but HR and fR were constant, demonstrating a temperature independent suppression of metabolism. Through the hibernation season tegus sporadically aroused and emerged from their burrows to warm up and after a short basking period would return to the burrows and swiftly resume hibernation. While hibernating, heart rate was characteristically regular but breathing was sporadic orepisodic. Arousals became more frequent towards the end of hibernation so that when they entered arousal from hibernation season most tegus were emerging daily. At this time daily maximum deep body temperature (Tbmax) swiftly returned to active season values but nighttime daily minimum deep body temperature (Tbmirt) values only showed a gradual increase through August, indicating different body temperature set points (Tbset) for active and sleep states. Changes in heart rate and breathing rate during the year showed greatest correlation with changes in photoperiod, although throughout hibernation HR and fR also showed tight correlation with Tb. / Science, Faculty of / Zoology, Department of / Graduate

La régulation endogène de la physiologie saisonnière, comme les rythmes circannuels et l’histoire photopériodique vécue pendant la gestation, permet aux individus d’anticiper à l’arrivée des saisons. L’objectif de cette thèse a été l’étude de l’effet de ces deux phénomènes à long terme sur les voies neuroendocrines contrôlant la physiologie saisonnière. Ces études montrent que les rythmes circannuels sont initiés au niveau de l’expression de la thyréostimuline dans la pars tuberalis et celle-ci est transmisse à l’hypothalamus où elle contrôle les niveaux d’hormone thyroïdienne et de neuropeptides. De façon similaire, la photopériode vécue pendant la gestation semble programmer l’expression des déiodinases dans l’hypothalamus des rongeurs adultes. Dans l’ensemble, ces résultats montrent la conservation du rôle de la pars tuberalis en tant que site central pour l’intégration des signaux impliqués dans le contrôle de la physiologie saisonnière. / Internal long-term timing strategies, like circannual rhythms and photoperiodic history, allow changes in physiology in anticipation to the seasons. In this project I investigated the effects of both types of history dependent timing on the neuro endocrine pathways for regulation of seasonal physiology. My work revealed that internal circannual timing is initiated through the control of thyroid stimulating hormone at the pars tuberalis and is integrated into the regulation of hypothalamic thyroid hormone levels and neuropeptides.Similarly, history-dependent expression in hypothalamic deiodinases was observed in adult rodents based on their gestational photoperiodic experience. Overall, my results highlight the pars tuberalis as a conserved central site in mammals for the integration of multiple seasonal cues, which via differential control of thyroid hormone levels in the hypothalamus dictates the timing in seasonal physiology.

O uso de répteis como modelos experimentais é limitado, principalmente devido às particularidades fisiológicas da classe, como as oscilações diárias e sazonais em seu comportamento. O objetivo deste estudo foi avaliar a atividade locomotora e nociceptiva de teiús (Salvator merianae) e iguanas-verdes (Iguana iguana) submetidos a testes comportamentais, ao longo do dia e do ano. Foram utilizados seis exemplares de cada espécie, mantidos em sala com temperatura ambiental (24 a 30°C) e fotoperíodo (12h:12h) controlados. Esses foram avaliados ao longo do dia (0:00h, 6:00h, 12:00h, 18:00h) e ao longo do ano (análise mensal). A análise locomotora foi realizada através do teste de campo aberto (teiús e iguanas), no qual o animal foi colocado no centro de uma arena circular por 15 minutos, e recebeu um ponto por cada ultrapassagem pelas subdivisões da mesma; e pelo teste de natação forçada (iguanas), no qual o tempo de atividade foi mensurado em piscina sem saída por 2 minutos. A resposta nociceptiva foi avaliada por meio de mensuração do período de latência do membro em resposta a estímulo nocivo térmico (25 segundos; 245 ± 7 mW/cm2) na superfície plantar do membro do animal. Não foram detectadas oscilações ao longo do ano no teste de campo aberto em nenhuma das espécies. Ao longo do dia, oscilações foram detectadas de Fevereiro a Dezembro em teiús; e em Abril, Maio, Junho e Outubro em iguanas. O tempo de atividade das iguanas foi menor em Janeiro e às 0:00h. Maiores latências de retirada do membro foram observadas nos meses de Maio e Agosto e às 6:00h em ambas espécies. Conclue-se que teiús e iguanas-verdes em ambiente controlado apresentam oscilações significativas em comportamento observado em teste de campo aberto, teste de natação forçada (somente iguanas) e teste plantar, que aparentemente não seguem padrões anuais claros, sendo provavelmente influenciado por fatores múltiplos ainda não compreendidos para as espécies. / The use of reptiles as experimental models is limited due to their physiological particularities, such as daily and annual fluctuations in behavior. The aim of this study was to evaluate locomotor and nociceptive activities of tegus (Salvator merianae) and green iguanas (Iguana iguana) throughout the day and the year, when undergoing behavioral tests. Six animals from each species, kept under controlled room temperature (24 to 30°C) and photoperiod (12h:12h), were used. They were evaluated throughout the day (0:00h, 6:00h, 12:00h and 18:00) and the year (monthly). Locomotor activity was measured by the open field test, in which the animal was placed in the center of a round arena for 15 minutes and the number of crossings through the subdivisions of the arena was counted, and by the forced swim test (iguanas), in which the activity period was timed after animals were placed in a pool with no scape for 2 minutes. Nociceptive activity was measured as the latency to limb withdrawal reflex in response to a noxious thermal stimulus (25 seconds, 245 ± 7 mW/cm2) in the plantar surface of the animal\'s limb. No differences were detected in locomotor activity in the open field test throughout the year in both species. Throughout the day, fluctuations were detected from February to December in tegus, and in April, May, June and October in iguanas. Activity period in iguanas were shorter in January and at 0:00h. Longer latencies to withdrawal reflex were detected in May and August and at 6:00h in both species. In conclusion, tegus and green iguanas kept in controlled environment have fluctuations in behavior presented in the open field test, the forced swim (only iguanas) test and the plantar test, which apparently do not follow a clear seasonal pattern and are probably influenced by multiple factors that are still unknown for both species.

O uso de répteis como modelos experimentais é limitado, principalmente devido às particularidades fisiológicas da classe, como as oscilações diárias e sazonais em seu comportamento. O objetivo deste estudo foi avaliar a atividade locomotora e nociceptiva de teiús (Salvator merianae) e iguanas-verdes (Iguana iguana) submetidos a testes comportamentais, ao longo do dia e do ano. Foram utilizados seis exemplares de cada espécie, mantidos em sala com temperatura ambiental (24 a 30°C) e fotoperíodo (12h:12h) controlados. Esses foram avaliados ao longo do dia (0:00h, 6:00h, 12:00h, 18:00h) e ao longo do ano (análise mensal). A análise locomotora foi realizada através do teste de campo aberto (teiús e iguanas), no qual o animal foi colocado no centro de uma arena circular por 15 minutos, e recebeu um ponto por cada ultrapassagem pelas subdivisões da mesma; e pelo teste de natação forçada (iguanas), no qual o tempo de atividade foi mensurado em piscina sem saída por 2 minutos. A resposta nociceptiva foi avaliada por meio de mensuração do período de latência do membro em resposta a estímulo nocivo térmico (25 segundos; 245 ± 7 mW/cm2) na superfície plantar do membro do animal. Não foram detectadas oscilações ao longo do ano no teste de campo aberto em nenhuma das espécies. Ao longo do dia, oscilações foram detectadas de Fevereiro a Dezembro em teiús; e em Abril, Maio, Junho e Outubro em iguanas. O tempo de atividade das iguanas foi menor em Janeiro e às 0:00h. Maiores latências de retirada do membro foram observadas nos meses de Maio e Agosto e às 6:00h em ambas espécies. Conclue-se que teiús e iguanas-verdes em ambiente controlado apresentam oscilações significativas em comportamento observado em teste de campo aberto, teste de natação forçada (somente iguanas) e teste plantar, que aparentemente não seguem padrões anuais claros, sendo provavelmente influenciado por fatores múltiplos ainda não compreendidos para as espécies. / The use of reptiles as experimental models is limited due to their physiological particularities, such as daily and annual fluctuations in behavior. The aim of this study was to evaluate locomotor and nociceptive activities of tegus (Salvator merianae) and green iguanas (Iguana iguana) throughout the day and the year, when undergoing behavioral tests. Six animals from each species, kept under controlled room temperature (24 to 30°C) and photoperiod (12h:12h), were used. They were evaluated throughout the day (0:00h, 6:00h, 12:00h and 18:00) and the year (monthly). Locomotor activity was measured by the open field test, in which the animal was placed in the center of a round arena for 15 minutes and the number of crossings through the subdivisions of the arena was counted, and by the forced swim test (iguanas), in which the activity period was timed after animals were placed in a pool with no scape for 2 minutes. Nociceptive activity was measured as the latency to limb withdrawal reflex in response to a noxious thermal stimulus (25 seconds, 245 ± 7 mW/cm2) in the plantar surface of the animal\'s limb. No differences were detected in locomotor activity in the open field test throughout the year in both species. Throughout the day, fluctuations were detected from February to December in tegus, and in April, May, June and October in iguanas. Activity period in iguanas were shorter in January and at 0:00h. Longer latencies to withdrawal reflex were detected in May and August and at 6:00h in both species. In conclusion, tegus and green iguanas kept in controlled environment have fluctuations in behavior presented in the open field test, the forced swim (only iguanas) test and the plantar test, which apparently do not follow a clear seasonal pattern and are probably influenced by multiple factors that are still unknown for both species.

L’intégrité fonctionnelle des organismes vivants, Homme y compris dépend des rythmes biologiques. La perturbation de ces rythmes due aux conditions de vie du monde moderne (travail posté, jet-lag,…) ou des circonstances naturelles (vieillissement), favorise l’installation de pathologies spécifiques (troubles du sommeil, l’obésité, le diabète,…). Afin de retarder l’apparition de ces troubles, la conception des projets expérimentaux sur des modèles d’animaux diurnes et des modèles d’animaux vivants dans des biotopes particuliers où l’Homme est présent (zone désertique) est nécessaire. Le but de ma thèse est d’essayer de comprendre les mécanismes neurophysiologiques d’adaptation aux conditions environnementales extrêmes chez le dromadaire (adapté à la chaleur) et le hamster d’Europe (adapté au froid). Une première partie révèle qu’en plus de la photopériode, la température ambiante est un véritable synchroniseur de l’horloge biologique du moins en absence du cycle lumière-obscurité. Dans une seconde partie, notre objectif était de vérifier si les variations saisonniers de la température ambiante sont capables comme les changements photopériodiques d’induire des modifications saisonnières sur le rythme de mélatonine et/ou de la température corporelle. Nous avons commencé par la vérification de l’existence de l’hétérothermie adaptative chez le dromadaire surtout que ce phénomène a été contesté récemment. Nos résultats démontrent que le dromadaire privé d’eau de boisson et placé sous des températures ambiantes élevées, adopte une régulation complexe de thermorégulation caractérisée par une alternance quotidienne de phase de poïkilothermie et d’homéothermie. Cet état d’hétérothermie a également été observé chez des dromadaires parfaitement hydratés, sujet d’une restriction alimentaire. Nos résultats concluent donc que l’hétérothermie adaptative chez le dromadaire est une combinaison de plusieurs facteurs qui interagissent le long du cycle lumière-obscurité, à savoir la température ambiante, la privation hydrique et la prise alimentaire. Finalement, nous avons démontré chez le hamster d’Europe que les neurones des noyaux arqués sont capables d’intégrer le signal photopériodique et cela indépendamment de la présence de la mélatonine. L’existence de ce mécanisme particulier d’intégration de la photopériode chez d’autres mammifères y compris l’Homme doit maintenant être recherchée. Nos résultats ouvrent la voie à la mise en évidence de son intérêt pratique dans le contrôle des rythmes biologiques en particulier dans celui des rythmes circannuels. / The functional integrity of living organisms, including human, depends on the biological rhythms. Disruption of these rhythms due to the living conditions of the modern world (shift work, jet lag ...) or natural circumstances (aging), leads various abnormalities (sleep disorders, obesity, diabetes ...). In order to understand pathophysiology of these abnormalities and adaptation in extreme environment, we need to design experiments on diurnal animals that cohabitate with human in specific biotopes. The aim of my thesis is to understand the neurophysiological mechanisms of adaptation to extreme environmental conditions in the camel (adapted to heat) and the European hamster (adapted to cold). Earlier we found that in addition to photoperiod, ambient temperature is a real synchronizer of the biological clock, at least in the absence of light-dark cycle. In the second part of my project, we investigated if seasonal variations of ambient temperature are capable to changes the rhythm of melatonin secretion and/or body temperature like photoperiod. Since heterothermy in camel challenged recently, we started our study by confirming the existence of adaptive heterothermy in camels. Our results demonstrate that dehydrated camels during exposure to daily heat show adaptive heterothermy. This mechanism is more complex because it is characterized by a daily alternation of two periods of poikilothermy and homeothermy. This adaptive heterothermy was also observed when camels are hydrated and food deprived. Based on our results, we can conclude that adaptive heterothermy in the Arabian camel is a combination of three factors interacting throughout the light-dark cycle: heat stress, water restriction, and level of food intake. Finally, we have demonstrated in European hamster that neurons of the arcuate nucleus are able to integrate photoperiodic signal, independent of melatonin. The existence of this particular mechanism of integration of photoperiod in other mammals including humans must be investigated. Our results promote to study the role of this new mechanism of integration of photoperiod in the control of biological rhythms in particular the circannual rhythms