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Physiological bases of prosocial behaviors

Periodic Reporting for period 1 - PHYSPROSOC (Physiological bases of prosocial behaviors)

Reporting period: 2023-07-01 to 2025-06-30

Social life combines cooperation and competition, and both factors shape the evolution of social cognition. The complexity of group relationships requires rapid adjustment to changing circumstances through behavioural and physiological flexibility. While primates have been the traditional model for studying these processes, corvids also display advanced socio-cognitive skills, including the formation of long-term affiliative bonds, maintenance of structured dominance hierarchies, and use of both cooperative and competitive strategies. These traits made them a suitable model for investigating the developmental and physiological mechanisms that underlie prosocial behaviour.

Behavioural studies in corvids have described many social strategies, yet the real-time physiological dynamics supporting these behaviours remained poorly understood, particularly in naturalistic group settings. The neurovisceral integration framework links behavioural flexibility to the brain’s ability to regulate the autonomic nervous system. Physiological measures such as heart rate, heart rate variability, and body temperature provide a non-invasive means to track this regulation, but their value as indicators of social bonding, cognitive flexibility, and stress coping in birds had not been systematically tested.

The project addressed this gap by combining continuous physiological monitoring with detailed behavioural and cognitive observations in free-moving crows, supplemented by limited comparative work in ring doves. Custom-designed wireless biosensors were used to record heart rate, heart rate variability, and body temperature during daily social interactions and in targeted behavioural tests. This approach captured the autonomic signatures of social bonding, cooperation, and alliance formation, and allowed longitudinal tracking of these patterns during development.

The work established direct links between individual differences in physiology and variation in prosocial tendencies, social learning and recognition. It demonstrated that early-life experiences and individual predispositions were reflected in measurable physiological patterns. Integrating behavioural ethology, physiology, and neuroscience provided a methodological framework for investigating complex social behaviour in a group setting.

Beyond its scientific contribution, the study showed that continuous, little-invasive physiological recording can inform animal welfare assessments and help to evaluate the impact of social isolation on development. These findings have wider relevance for understanding the biological foundations of social behaviour and the consequences of disrupted social contact, a topic that has gained urgency in light of recent global events.
The work was carried out on a newly established social group of carrion crows and existing captive colony of ring doves. The crows were collected from the field as early nestlings and hand raised in two separate groups. This approach ensured full control over their socialisation history and allowed systematic tracking of behavioural and physiological development from the earliest stages.

All experimental procedures were successfully implemented. The implantation of wireless biosensors functioned reliably, enabling continuous recording of heart rate and body temperature over extended periods. In crows, data collection was repeated at three key developmental stages: the early juvenile stage (around three months after hatching, shortly after independence from caregivers), the late juvenile stage (around six months, before the first breeding season), and the subadult stage (around one year, approaching full sexual maturity). This longitudinal approach captured individual changes in physiology and behaviour across their developmental trajectory.

Behavioural observations were conducted both in spontaneous social settings within the aviary and in a series of structured behavioural tasks as described in the proposal. These tasks included the attention task, the social learning task, the audio recognition task of conspecific and heterospecific individuals, and the temporal separation task. All four tasks were conducted at each developmental stage, allowing direct comparisons of behavioural performance and associated physiological patterns over time. This repeated-measures design provided detailed insight into how prosocial behaviour, cognitive performance, and autonomic regulation developed in individuals.

An additional experiment investigated recognition of social valence using conditioned auditory stimuli from the human who raised the crows. During the hand-rearing phase, two distinct human-produced sounds were paired with different types of interaction: one consistently associated with gentle social preening, and the other with a mildly aversive handling procedure. In later testing, these conditioned cues were presented together with additional familiar and unfamiliar human sound cues, as well as selected vocalisations from conspecifics and heterospecifics. Testing took place in a controlled context while heart rate and body temperature were recorded continuously. This design allowed assessment of whether crows differentiated between positive and negative social associations, and how familiarity and valence influenced autonomic and behavioural responses. Repeating the experiment at multiple developmental stages provided insight into how learned social valence and recognition of familiar versus unfamiliar cues changed during maturation.

In ring doves, the method was applied in pairs and in mixed sex small social groups within aviaries. Physiological data were successfully collected in spontaneous social contexts, confirming the feasibility of the approach in this species.
The project advanced the study of animal social cognition by providing the first longitudinal dataset linking continuous physiological recordings of heart rate and body temperature with both spontaneous and experimentally elicited behaviour in hand-raised crows. The integration of physiological monitoring with repeated behavioural testing across three defined developmental stages offered a more detailed picture of how social behaviour, cognitive performance, and autonomic regulation change from early juvenile life to sexual maturity.

The successful implantation and long-term functioning of custom wireless biosensors in free-moving crows was a significant methodological achievement. Continuous, high-quality physiological data were obtained over extended periods without compromising animal welfare, both in naturalistic social settings and in controlled experiments. This approach went beyond previous studies, which mostly relied on short-term or isolated measurements, and demonstrated the feasibility of following autonomic function in socially interacting animals over months and through key developmental stages.

Preliminary behavioural results showed marked individual variation in social learning and social attention, with performance depending on affiliation levels and dominance status within the group. Birds with higher affiliation scores and central positions in the dominance hierarchy engaged more readily in social learning tasks and showed stronger attention to conspecific cues. These patterns were similar in the physiological data, with more socially integrated individuals hinting at distinct autonomic profiles compared to low-affiliation or subordinate birds. The match between behavioural and physiological patterns has to be further analyzed but however provides potential for the link between social integration, cognitive engagement, and autonomic regulation.

The additional preliminary results on conditioned human auditory cues showed that crows retained distinct autonomic and behavioural responses to positive and mildly aversive social experiences, and that these responses were shaped by both the familiarity of the stimulus and its valence. These findings could extend the understanding of social valence recognition and highlight how early-life experiences leave lasting effects on the physiological sensitivity to social signals.

The methodology and results can have value beyond basic science. The approach offers a powerful tool for assessing social integration, monitoring welfare, and detecting the physiological effects of social isolation or stress in both captive and wild populations. It can be applied in conservation programmes, animal management, and comparative studies across species. The connection between developmental social experience and long-term physiological regulation also creates a connection to human research on social bonding and emotional regulation, supporting collaboration between behavioural ecology, neuroscience, and psychology.

Future research I aim to pursue is to combine this approach with hormonal analyses, particularly oxytocin and corticosterone measurements, to capture the endocrine dimension of social regulation. Integrating these measures with continuous autonomic monitoring would provide a more complete understanding of how social relationships shape behaviour and physiology over time. Strengthening interdisciplinary work and improving fabrication of specialised biosensor would help ensure scientific advances and innovation in the field of behavioral physiology.
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