The domestic chick (Gallus gallus domesticus) has been widely used as an animal model to investigate spatial orientation and the neural mechanisms underlying this function. In all vertebrate species the hippocampus plays an essential role in spatial orientation. Since the hippocampus is a bilateral structure, it is important to investigate the specific role of the left and the right hippocampi in spatial processing. Although, the domestic chick has been often used as animal model to assess cognitive lateralization, the involvement of the left and the right hippocampal formation in spatial orientation has been poorly investigated in this model. Behavioral studies using monocular eye occlusion have shown that in chicks the left eye-system (right hemisphere) is involved in the elaboration of spatial relational information, while the right eye-system (left hemisphere) processes local information. However, while visual lateralization in chicks had been traditionally considered to be induced by embryonic light exposure, recent studies suggest the presence of structural and behavioural asymmetries also in dark-incubated chicks. Thus, the main aim of this thesis was to test the lateralization of hippocampal functions in dark incubated chicks, both in spatial and non-spatial tasks. In the first study dark-incubated chicks were trained to orient in a large circular arena using spatial relational information provided by free-standing objects. Once chicks reached a learning criterion they were tested binocularly or under a monocular eye-occlusion condition. This study provided the first demonstration that domestic chicks are able to orient by relational spatial information provided by free-standing objects, in binocular vision conditions. However, if either one of the two eyes was occluded, chicks failed the orientation task. These results show that at least in dark-incubated chicks binocular integration is needed to solve this spatial orientation task. We also investigated if chicks have a preference to orient by local or spatial information provided by free-standing objects and if this ability is influenced by eye occlusion. Chicks preferred to use local over spatial cues to orient, both in binocular and monocular conditions (independently of which eye was occluded). These results indicate that local cues are processed by both eye-systems and do not require access to information from both eyes, contrary to relational spatial cues. Using the same setup, in the second study we directly investigated the involvement of chicks’ left and right hippocampal formation during orientation by free-standing objects. For this purpose we performed an immunohistochemical staining of the immediate early gene product c-Fos (a neural activity marker). Two independent groups of dark-incubated chicks were trained to find food in the large circular arena and the level of hippocampal activation was compared between the two groups. One group was trained to orient exclusively by local cues, while the other was orienting by spatial relational information provided by free-standing objects. This revealed selective activation of the right hippocampus during orientation by spatial relational information in dark-incubated chicks. While monocular occlusion has often been used to test lateralization of spatial functions in chicks, it is still unclear whether this manipulation affects hippocampal activation. The aim of the third study was to clarify this issue, by exposing dark-incubated chicks to a novel environment in conditions of monocular occlusion or binocular vision. Activation of the hippocampal formation was once again measured by c-Fos expression. Exposure to a novel environment is known to trigger hippocampal activation in different animals, including domestic chicks. As expected, exposure to the novel environment activated the hippocampus in binocular vision conditions. However, if either one of the eyes was occluded, the hippocampal c-Fos expression did not rise above what observed in the baseline condition (chicks maintained in a familiar environment). Thus, successful hippocampal response to a novel environment requires input from both eyes. Our results also suggest that monocular occlusion equally affects the left and the right hippocampus. Overall, access to information from both eyes plays a crucial role for the acquisition of a spatial map of a novel environment, in line with the behavioral results of the first study. Moreover, a task independent lateralization effect, with higher c-Fos expression in the left compared to the right hippocampus, could be observed in all the experimental conditions. This confirms the presence of neuroanatomical lateralization in dark-incubated chicks. The last study investigated whether chicks’ hippocampus would also respond to novel social stimuli, in line with the activation observed in this structure after exposure to a novel environment. Only few studies have directly investigated the involvement of birds’ hippocampal formation in social functions. Here, the hippocampal activation was compared between chicks exposed to an unfamiliar conspecific vs. chicks exposed to a familiar one. We found that the ventral and dorsomedial portion of the right hippocampus of dark-incubated chicks responds to an unfamiliar individual. This provides the first demonstration of hippocampal sensitivity to social novelty in birds. Overall the studies performed in this thesis indicate a selective lateralized involvement of domestic chicks’ hippocampal formation not only in spatial, but also in social functions.

Lateralization of hippocampal functions in domestic chicks (Gallus gallus domesticus) / Morandi Raikova, Anastasia. - (2021 Nov 12), pp. 1-138. [10.15168/11572_321134]

Lateralization of hippocampal functions in domestic chicks (Gallus gallus domesticus)

Morandi Raikova, Anastasia
2021-11-12

Abstract

The domestic chick (Gallus gallus domesticus) has been widely used as an animal model to investigate spatial orientation and the neural mechanisms underlying this function. In all vertebrate species the hippocampus plays an essential role in spatial orientation. Since the hippocampus is a bilateral structure, it is important to investigate the specific role of the left and the right hippocampi in spatial processing. Although, the domestic chick has been often used as animal model to assess cognitive lateralization, the involvement of the left and the right hippocampal formation in spatial orientation has been poorly investigated in this model. Behavioral studies using monocular eye occlusion have shown that in chicks the left eye-system (right hemisphere) is involved in the elaboration of spatial relational information, while the right eye-system (left hemisphere) processes local information. However, while visual lateralization in chicks had been traditionally considered to be induced by embryonic light exposure, recent studies suggest the presence of structural and behavioural asymmetries also in dark-incubated chicks. Thus, the main aim of this thesis was to test the lateralization of hippocampal functions in dark incubated chicks, both in spatial and non-spatial tasks. In the first study dark-incubated chicks were trained to orient in a large circular arena using spatial relational information provided by free-standing objects. Once chicks reached a learning criterion they were tested binocularly or under a monocular eye-occlusion condition. This study provided the first demonstration that domestic chicks are able to orient by relational spatial information provided by free-standing objects, in binocular vision conditions. However, if either one of the two eyes was occluded, chicks failed the orientation task. These results show that at least in dark-incubated chicks binocular integration is needed to solve this spatial orientation task. We also investigated if chicks have a preference to orient by local or spatial information provided by free-standing objects and if this ability is influenced by eye occlusion. Chicks preferred to use local over spatial cues to orient, both in binocular and monocular conditions (independently of which eye was occluded). These results indicate that local cues are processed by both eye-systems and do not require access to information from both eyes, contrary to relational spatial cues. Using the same setup, in the second study we directly investigated the involvement of chicks’ left and right hippocampal formation during orientation by free-standing objects. For this purpose we performed an immunohistochemical staining of the immediate early gene product c-Fos (a neural activity marker). Two independent groups of dark-incubated chicks were trained to find food in the large circular arena and the level of hippocampal activation was compared between the two groups. One group was trained to orient exclusively by local cues, while the other was orienting by spatial relational information provided by free-standing objects. This revealed selective activation of the right hippocampus during orientation by spatial relational information in dark-incubated chicks. While monocular occlusion has often been used to test lateralization of spatial functions in chicks, it is still unclear whether this manipulation affects hippocampal activation. The aim of the third study was to clarify this issue, by exposing dark-incubated chicks to a novel environment in conditions of monocular occlusion or binocular vision. Activation of the hippocampal formation was once again measured by c-Fos expression. Exposure to a novel environment is known to trigger hippocampal activation in different animals, including domestic chicks. As expected, exposure to the novel environment activated the hippocampus in binocular vision conditions. However, if either one of the eyes was occluded, the hippocampal c-Fos expression did not rise above what observed in the baseline condition (chicks maintained in a familiar environment). Thus, successful hippocampal response to a novel environment requires input from both eyes. Our results also suggest that monocular occlusion equally affects the left and the right hippocampus. Overall, access to information from both eyes plays a crucial role for the acquisition of a spatial map of a novel environment, in line with the behavioral results of the first study. Moreover, a task independent lateralization effect, with higher c-Fos expression in the left compared to the right hippocampus, could be observed in all the experimental conditions. This confirms the presence of neuroanatomical lateralization in dark-incubated chicks. The last study investigated whether chicks’ hippocampus would also respond to novel social stimuli, in line with the activation observed in this structure after exposure to a novel environment. Only few studies have directly investigated the involvement of birds’ hippocampal formation in social functions. Here, the hippocampal activation was compared between chicks exposed to an unfamiliar conspecific vs. chicks exposed to a familiar one. We found that the ventral and dorsomedial portion of the right hippocampus of dark-incubated chicks responds to an unfamiliar individual. This provides the first demonstration of hippocampal sensitivity to social novelty in birds. Overall the studies performed in this thesis indicate a selective lateralized involvement of domestic chicks’ hippocampal formation not only in spatial, but also in social functions.
12-nov-2021
XXXIII
2019-2020
Università degli Studi di Trento
Cognitive and Brain Sciences
Mayer, Uwe
no
Inglese
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