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Hamilton's rule provides the cornerstone for our understanding of the evolution of all forms of social behavior, from altruism to spite, across all organisms, from viruses to humans. In contrast to the standard prediction from Hamilton's rule, recent studies have suggested that altruistic helping can be favored even if it does not benefit relatives, as long as it decreases the environmentally induced variance of their reproductive success ("altruistic bet-hedging"). However, previous predictions both rely on an approximation and focus on variance-reducing helping behaviors. We derived a version of Hamilton's rule that fully captures environmental variability. This shows that decreasing (or increasing) the variance in the absolute reproductive success of relatives does not have a consistent effect-it can either favor or disfavor the evolution of helping. We then empirically quantified the effect of helping on the variance in reproductive success across 15 species of cooperatively breeding birds. We found that a) helping did not consistently decrease the variance of reproductive success and often increased it, and b) the mean benefits of helping across environments consistently outweighed other variability components of reproductive success. Altogether, our theoretical and empirical results suggest that the effects of helping on the variability components of reproductive success have not played a consistent or strong role in favoring helping.

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Comparative studies reliant on single personality surveys to rate wild primates are scarce yet remain critical for developing a holistic comparative understanding of personality. Differences in survey design, item exclusion, and factor selection impede cross-study comparisons. To address these challenges, we used consistently collected data to assess personality trait structures in wild rhesus (Macaca mulatta), bonnet (M. radiata), and long-tailed (M. fascicularis) macaques that varied in their degree of phylogenetic closeness, species-typical social styles, and anthropogenic exposure in urban or urban-rural environments. We administered 51-item personality surveys to familiar raters, and, after reliability and structure screenings, isolated 4-5 factor solutions among the species. Four consistent factors emerged: Confident, Sociable, Active, and Irritable/Equable. This latter factor had differential expression across species. Item composition of the Irritable/Equable factor was consistent with their anticipated differences in social styles, but confounded by cross-site anthropogenic variation. We also administered a 43-item survey confined to human-primate situations which paralleled our findings of social style variation, while also exhibiting variation that aligned with population differences in human density. Our findings indicate that macaque personality trait structures may be emergent outcomes of evolutionary and/or socioecological processes, but further research is needed to parse these processes' relative contributions.

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Domestic dogs have been shown to copy their caregiver's actions, including ones which are causally-irrelevant to a physical goal-a behaviour called "overimitation". In a new overimitation task with a non-food reward, this study investigated "causal misunderstanding"-falsely assuming causally-irrelevant actions to have functional relevancy-as an explanation for dog overimitation (N = 81). By providing dogs with prior experience of the task to learn about the consequences of its irrelevant box-stepping and relevant bucket-opening action to obtain a toy-ball, we tested whether and when dogs would copy their caregiver's irrelevant-action demonstrations. Dogs with and without prior experience were compared to a third (control) group of dogs, who had neither prior experience nor caregiver demonstrations of the task. Results revealed that the timing of overimitation, rather than its frequency, was closely related to dogs' prior experience: dogs with prior experience attended to their reward first, then interacted with the irrelevant box later ("post-goal overimitation"), while dogs without prior experience first interacted with the irrelevant box ("pre-goal overimitation"). Our results suggest that, when action consequences are understood, dogs are overimitating for a secondary social goal that is clearly distinct from the task goal of obtaining a physical reward.

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The study of social interactions lies at the core of several disciplines such as psychiatry, psychology and ethology, just to name a few. In this context, understanding the temporal patterns underlying interactive behaviors is of crucial importance. Here, we employed T-pattern detection and analysis to study social interactions in ten pairs of Wistar rats tested in an Open-Field environment. We found four different categories of interactive behaviors. One of them was of particular interest to us because it consisted of behavioral events that, taken individually, should not underlie an interaction of any kind; however, they were included in T-patterns, which is suggestive of a dyadic temporal coordination in the behavioral expression of two individuals. Within this category, we described for the first time a new subcategory of apparent interaction patterns characterized by events that one of the two rats repeats only if previously produced by the partner (i.e., behavioral mirroring). These findings are discussed in functional terms for rodents and in light of our current understanding of social interactions in humans.

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The commercial wasabi pastes commonly used for food preparation contain a homologous compound of chemosensory isothiocyanates (ITCs) that elicit an irritating sensation upon consumption. The impact of sniffing dietary alcoholic beverages on the sensation of wasabi spiciness has never been studied. While most sensory evaluation studies focus on individual food and beverages separately, there is a lack of research on the olfactory study of sniffing liquor while consuming wasabi. Here, a methodology is developed that combines the use of an animal behavioral study and a convolutional neural network to analyze the facial expressions of mice when they simultaneously sniff liquor and consume wasabi. The results demonstrate that the trained and validated deep learning model recognizes 29% of the images depicting co-treatment of wasabi and alcohol belonging to the class of the wasabi-negative liquor-positive group without the need for prior training materials filtering. Statistical analysis of mouse grimace scale scores obtained from the selected video frame images reveals a significant difference (P < 0.01) between the presence and absence of liquor. This finding suggests that dietary alcoholic beverages might have a diminishing effect on the wasabi-elicited reactions in mice. This combinatory methodology holds potential for individual ITC compound screening and sensory analyses of spirit components in the future. However, further study is required to investigate the underlying mechanism of alcohol-induced suppression of wasabi pungency.

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Social animals make behavioural decisions based on local habitat and conspecifics, as well as memorized past experience (i.e. 'familiarity') with habitat and conspecifics. Here, we develop a conceptual and empirical understanding of how spatial and social familiarity fit within the spatial-social interface-a novel framework integrating the spatial and social components of animal behaviour. We conducted a multi-scale analysis of the movements of GPS-collared plains bison (Bison bison, n = 66) residing in and around Yellowstone National Park, USA. We found that both spatial and social familiarity mediate how individuals respond to their spatial and social environments. For instance, individuals with high spatial familiarity rely on their own knowledge as opposed to their conspecifics, and individuals with high social familiarity rely more strongly on the movement of conspecifics to guide their own movement. We also found that fine-scale spatial and social phenotypes often scale up to broad-scale phenotypes. For instance, bison that select more strongly to align with their nearest neighbour have larger home ranges. By integrating spatial and social familiarity into the spatial-social interface, we demonstrate the utility of the interface for testing hypotheses, while also highlighting the pervasive importance of cognitive mechanisms in animal behaviour. This article is part of the theme issue 'The spatial-social interface: a theoretical and empirical integration'.

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Studying the spatial-social interface requires tools that distinguish between social and spatial drivers of interactions. Testing hypotheses about the factors determining animal interactions often involves comparing observed interactions with reference or 'null' models. One approach to accounting for spatial drivers of social interactions in reference models is randomizing animal movement paths to decouple spatial and social phenotypes while maintaining environmental effects on movements. Here, we update a reference model that detects social attraction above the effect of spatial constraints. We explore the use of our 'wrap-around' method and compare its performance to the previous approach using agent-based simulations. The wrap-around method provides reference models that are more similar to the original tracking data, while still distinguishing between social and spatial drivers. Furthermore, the wrap-around approach results in fewer false-positives than its predecessor, especially when animals do not return to one place each night but change movement foci, either locally or directionally. Finally, we show that interactions among GPS-tracked griffon vultures (Gyps fulvus) emerge from social attraction rather than from spatial constraints on their movements. We conclude by highlighting the biological situations in which the updated method might be most suitable for testing hypotheses about the underlying causes of social interactions. This article is part of the theme issue 'The spatial-social interface: a theoretical and empirical integration'.

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Animals simultaneously navigate spatial and social environments, and their decision-making with respect to those environments constitutes their spatial (e.g. habitat selection) and social (e.g. conspecific associations) phenotypes. The spatial-social interface is a recently introduced conceptual framework linking these components of spatial and social ecology. The spatial-social interface is inherently scale-dependent, yet it has not been integrated with the rich body of literature on ecological scale. Here, we develop a conceptual connection between the spatial-social interface and ecological scale. We propose three key innovations that incrementally build upon each other. First, the use-availability framework that underpins a large body of literature in behavioural ecology can be used in analogy to the phenotype-environment nomenclature and is transferable across the spatial and social realms. Second, both spatial and social phenotypes are hierarchical, with nested components that are linked via constraints-from the top down-or emergent properties-from the bottom up. Finally, in both the spatial and social realms, the definitions of environment and phenotype depend on the focal scale of inquiry. These conceptual innovations cast our understanding of the relationships between social and spatial dimensions of animal ecology in a new light, allowing a more holistic understanding and clearer hypothesis development for animal behaviour. This article is part of the theme issue 'The spatial-social interface: a theoretical and empirical integration'.

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Sensory Adaptation (SA) is a prominent aspect of how neurons respond to sensory signals, ubiquitous across species and modalities. However, SA depends on the activation state of the brain and the extent to which SA is expressed in awake, behaving animals during active sensation remains unclear. Here, we addressed this question by training head-fixed mice to detect an object using their whiskers and recording neuronal activity from barrel cortex whilst simultaneously imaging the whiskers in 3D. We found that neuronal responses decreased during the course of whisker-object touch sequences and that this was due to two factors. First, a motor effect, whereby, during a sequence of touches, later touches were mechanically weaker than early ones. Second, a sensory encoding effect, whereby neuronal tuning to touch became progressively less sensitive during the course of a touch sequence. The sensory encoding effect was whisker-specific. These results show that SA does occur during active whisker sensing and suggest that SA is fundamental to sensation during natural behaviour.

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Animals exhibiting mirror self-recognition (MSR) are considered self-aware; however, studies on their level of self-awareness remain inconclusive. Recent research has indicated the potential for cleaner fish (Labroides dimidiatus) to possess a sophisticated level of private self-awareness. However, as this study revealed only an aspect of private self-awareness, further investigation into other elements is essential to substantiate this hypothesis. Here, we show that cleaner fish, having attained MSR, construct a mental image of their bodies by investigating their ability to recall body size. A size-based hierarchy governs the outcomes of their confrontations. The mirror-naïve fish behaved aggressively when presented with photographs of two unfamiliar conspecifics that were 10% larger and 10% smaller than their body sizes. After passing the MSR test, they refrained from aggression toward the larger photographs but still behaved aggressively toward the smaller ones without re-examining their mirror images. These findings suggest that cleaner fish accurately recognize their body size based on mental images of their bodies formed through MSR. Additionally, mirror-experienced fish frequently revisited the mirror when presented with an intimidating larger photograph, implying the potential use of mirrors for assessing body size. Our study established cleaner fish as the first non-human animal to be demonstrated to possess private self-awareness.

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BACKGROUND: The natural light environment is far more complex than that experienced by animals under laboratory conditions. As a burrowing species, wild mice are able to self-modulate their light exposure, a concept known as light environment sampling behaviour. By contrast, under laboratory conditions mice have little opportunity to exhibit this behaviour. To address this issue, here we introduce a simple nestbox paradigm to allow mice to self-modulate their light environment. Dark nestboxes fitted with passive infrared sensors were used to monitor locomotor activity, circadian entrainment, decision making and light environment sampling behaviour. RESULTS: Under these conditions, mice significantly reduce their light exposure to an average of just 0.8 h across a 24 h period. In addition, mice show a distinct pattern of light environment sampling behaviour, with peaks at dawn and dusk under a ramped light dark cycle. Furthermore, we show that the timing of light environment sampling behaviour depends upon endogenous circadian rhythms and is abolished in mice lacking a circadian clock, indicating a feedback loop between light, the circadian clock and behaviour. CONCLUSIONS: Our results highlight the important role of behaviour in modifying the light signals available for circadian entrainment under natural conditions.

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Wildlife tagging provides critical insights into animal movement ecology, physiology, and behavior amid global ecosystem changes. However, the stress induced by capture, handling, and tagging can impact post-release locomotion and activity and, consequently, the interpretation of study results. Here, we analyze post-tagging effects on 1585 individuals of 42 terrestrial mammal species using collar-collected GPS and accelerometer data. Species-specific displacements and overall dynamic body acceleration, as a proxy for activity, were assessed over 20 days post-release to quantify disturbance intensity, recovery duration, and speed. Differences were evaluated, considering species-specific traits and the human footprint of the study region. Over 70% of the analyzed species exhibited significant behavioral changes following collaring events. Herbivores traveled farther with variable activity reactions, while omnivores and carnivores were initially less active and mobile. Recovery duration proved brief, with alterations diminishing within 4-7 tracking days for most species. Herbivores, particularly males, showed quicker displacement recovery (4 days) but slower activity recovery (7 days). Individuals in high human footprint areas displayed faster recovery, indicating adaptation to human disturbance. Our findings emphasize the necessity of extending tracking periods beyond 1 week and particular caution in remote study areas or herbivore-focused research, specifically in smaller mammals.

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BACKGROUND: Preterm birth is associated with brain injury and long-term behavioral abnormalities, for which there are limited prevention options. When born preterm, infants prematurely lose placental neurosteroid (allopregnanolone) support. This increases the risk of excitotoxic damage to the brain, which increases the risk of injury, causing long-term deficits in behavior, myelination, and alterations to neurotransmitter pathways. We propose that postnatal restoration of neurosteroid action through zuranolone therapy will reduce neurological impairments following preterm birth. METHODS: Guinea pig dams underwent survival cesarean section surgery to deliver pups prematurely (GA64) or at term (GA69). Between birth and term equivalence age, preterm pups received vehicle (15% ß-cyclodextrin) or the allopregnanolone analogue zuranolone (1 mg/kg/day). Behavioral analysis was performed at postnatal day (PND) 7 and 40, before tissue collection at PND 42. Immunostaining for myelin basic protein (MBP), as well as real-time polymerase chain reaction to characterize oligodendrocyte lineage and neurotransmitter pathways, was performed in frontal cortex tissues. RESULTS: Zuranolone treatment prevented the hyperactive phenotype in preterm-born offspring, most markedly in males. Additionally, preterm-related reductions in MBP were ameliorated. Several preterm-related alterations in mRNA expression of dopaminergic, glutamatergic, and GABAergic pathways were also restored back to that of a term control level. CONCLUSION: This is the first study to assess zuranolone treatment as a neuroprotective therapy following preterm birth. Zuranolone treatment improved behavioral outcomes and structural changes in the preterm offspring, which continued long term until at least a late childhood timepoint. Clinical studies are warranted for further exploring the neuroprotective possibilities of this treatment following preterm birth.

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The Ocean Networks Canada (ONC) cabled video-observatory at the Barkley Canyon Node (British Columbia, Canada) was recently the site of a Fish Acoustics and Attraction Experiment (FAAE), from May 21, 2022 to July 16, 2023, combining observations from High-Definition (HD) video, acoustic imaging sonar, and underwater sounds at a depth of 645 m, to examine the effects of light and bait on deep-sea fish and invertebrate behaviors. The unexpected presence of at least eight (six recurrent and two temporary) sub-adult male northern elephant seals (Mirounga angustirostris) was reported in 113 and 210 recordings out of 9737 HD and 2805 sonar videos at the site, respectively. Elephant seals were found at the site during seven distinct periods between June 22, 2022 and May 19, 2023. Ethograms provided insights into the seal's deep-sea resting and foraging strategies, including prey selection. We hypothesized that the ability of elephant seals to perform repeated visits to the same site over long periods (> 10 days) was due to the noise generated by the sonar, suggesting that they learned to use that anthropogenic source as an indicator of food location, also known as the "dinner bell" effect. One interpretation is that elephant seals are attracted to the FAAE site due to the availability of prey and use the infrastructure as a foraging and resting site, but then take advantage of fish disturbance caused by the camera lights to improve foraging success. Our video observations demonstrated that northern elephant seals primarily focused on actively swimming sablefish (Anoplopoma fimbria), ignoring stationary or drifting prey. Moreover, we found that elephant seals appear to produce (voluntary or involuntary) infrasonic sounds in a foraging context. This study highlights the utility of designing marine observatories with spatially and temporally cross-referenced data collection from instruments representing multiple modalities of observation.

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Both cats and dogs fetch, but the likely function(s) of this behavior for each species have not been compared. In this study, we assessed data from online surveys of cat and dog behavior (Fe-BARQ; C-BARQ) completed by cat (N = 8224) and dog owners (N = 73724). We assessed responses to the items "Plays 'fetch'; likes to retrieve thrown objects or toys" (Fe-BARQ) and "Will 'fetch' or attempt to fetch sticks, balls, or objects" (C-BARQ). Cats and dogs described as "sometimes," "usually" or "always" fetching were categorized as fetchers. Regression models were used to examine which animal-related (e.g., sex, age) and environmental factors best predicted fetching, and chi-square tests were used to explore the effect of breed on fetching behavior. Fetching was reported in 40.9% of cats and 77.8% of dogs. In cats, fetching was correlated with play and activity. In dogs, fetching was correlated with overall trainability. In both cats and dogs, being female, older, living with (other) dogs, and having health problems decreased the likelihood of fetching. Breed effects were observed in both species, with fetching more prominent in cat breeds originating in the Far East (e.g., Burmese, Siamese, and Tonkinese) and in dog breeds from the Retriever, UK Rural, Poodle, Pointer and Spaniel clades. We discuss the results in the context of domestication history of both cats and dogs and posit several hypotheses about why fetching behavior is observed in both.

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Rumination behavior in cattle can provide valuable information for monitoring health status and animal welfare, but continuous monitoring is essential to detect changes in rumination behavior. In a previous study validating the use of a respiration rate sensor equipped with a triaxial accelerometer, the regurgitation process was also clearly visible in the pressure and accelerometer data. The aim of the present study, therefore, was to measure the individual lengths of rumination cycles and to validate whether the sensor data showed the same number of regurgitations as those counted visually (video or direct observation). For this purpose, 19 Holstein Friesian cows equipped with a respiration rate sensor were observed for two years, with a focus on rumination behavior. The results showed a mean duration of 59.27 ± 9.01 s (mean ± SD) per rumination cycle and good agreement (sensitivity: 99.1-100%, specificity: 87.8-95%) between the two methods (sensor and visual observations). However, the frequency of data streaming (continuously or every 30 s) from the sensor to the data storage system strongly influenced the classification performance. In the future, an algorithm and a data cache will be integrated into the sensor to provide rumination time as an additional output.

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Accurate detection and tracking of animals across diverse environments are crucial for studying brain and behavior. Recently, computer vision techniques have become essential for high-throughput behavioral studies; however, localizing animals in complex conditions remains challenging due to intra-class visual variability and environmental diversity. These challenges hinder studies in naturalistic settings, such as when animals are partially concealed within nests. Moreover, current tools are laborious and time-consuming, requiring extensive, setup-specific annotation and training procedures. To address these challenges, we introduce the 'Detect-Any-Mouse-Model' (DAMM), an object detector for localizing mice in complex environments with minimal training. Our approach involved collecting and annotating a diverse dataset of single- and multi-housed mice in complex setups. We trained a Mask R-CNN, a popular object detector in animal studies, to perform instance segmentation and validated DAMM's performance on a collection of downstream datasets using zero-shot and few-shot inference. DAMM excels in zero-shot inference, detecting mice and even rats, in entirely unseen scenarios and further improves with minimal training. Using the SORT algorithm, we demonstrate robust tracking, competitive with keypoint-estimation-based methods. Notably, to advance and simplify behavioral studies, we release our code, model weights, and data, along with a user-friendly Python API and a Google Colab implementation.

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The reinforcing nature of social interactions is necessary for the maintenance of appropriate social behavior. However, the neural substrates underlying social reward processing and how they might differ based on the sex and internal state of the animal remains unknown. It is also unclear whether these neural substrates are shared with those involved in nonsocial rewarding processing. We developed a fully automated, two choice (social-sucrose) operant assay in which mice choose between social and nonsocial rewards to directly compare the reward-related behaviors associated with two competing stimuli. We performed cellular resolution calcium imaging of medial prefrontal cortex (mPFC) neurons in male and female mice across varying states of water restriction and social isolation. We found that mPFC neurons maintain largely non-overlapping, flexible representations of social and nonsocial reward that vary with internal state in a sex-dependent manner. Additionally, optogenetic manipulation of mPFC activity during the reward period of the assay disrupted reward-seeking behavior across male and female mice. Thus, using a two choice operant assay, we have identified sex-dependent, non-overlapping neural representations of social and nonsocial reward in the mPFC that vary with internal state and that are essential for appropriate reward-seeking behavior.

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BACKGROUND: Reaching and grasping (R&G) in rats is commonly used as an outcome measure to investigate the effectiveness of rehabilitation or treatment strategies to recover forelimb function post spinal cord injury. Kinematic analysis has been limited to the wrist and digit movements. Kinematic profiles of the more proximal body segments that play an equally crucial role in successfully executing the task remain unexplored. Additionally, understanding of different forelimb muscle activity, their interactions, and their correlation with the kinematics of R&G movement is scarce. NEW METHOD: In this work, novel methodologies to comprehensively assess and quantify the 3D kinematics of the proximal and distal forelimb joints along with associated muscle activity during R&G movements in adult rats are developed and discussed. RESULTS: Our data show that different phases of R&G identified using the novel kinematic and EMG-based approach correlate with the well-established descriptors of R&G stages derived from the Whishaw scoring system. Additionally, the developed methodology allows describing the temporal activity of individual muscles and associated mechanical and physiological properties during different phases of the motor task. COMPARISON WITH EXISTING METHOD(S): R&G phases and their sub-components are identified and quantified using the developed kinematic and EMG-based approach. Importantly, the identified R&G phases closely match the well-established qualitative descriptors of the R&G task proposed by Whishaw and colleagues. CONCLUSIONS: The present work provides an in-depth objective analysis of kinematics and EMG activity of R&G behavior, paving the way to a standardized approach to assessing this critical rodent motor function in future studies.

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