Biology subjects

Clayton, N. S.

Publications and source records attributed to Clayton, N. S..

10 recordsLinked to original sources

bioRxiv2026

Development of cognition in corvids

Corvids - members of the crow family - exhibit some of the most sophisticated cognitive abilities outside the primate lineage, yet the developmental origins of many of these abilities remain poorly understood. Here, we present a systematic review of the past 20 years (from 2005) of empirical research on corvid cognitive development, synthesising evidence across core/ foundational, social and physical cognitive domains. Using a structured search strategy and detailed coding framework, we identified 47 relevant studies spanning 16 corvid species. We evaluate and discuss developmental trajectories, species/ taxa-level variation and methodological robustness across studies. For within and between-taxa comparisons, we particularly focus on the best represented abilities in the coded sample: 1) object permanence and caching; 2) tool-use/ manufacture; 3) object manipulation and play; and 4) gaze following. Corvid developmental patterns show both parallels and divergences from those documented in primates and other taxa. However, the existing corvid evidence base is constrained by small samples, captive biases, limited longitudinal data and under-representation of key cognitive abilities, such as executive function, causal reasoning, self-control, metacognition, spatial memory and social learning. We outline critical gaps and future directions, emphasising the need for comparative, longitudinal and ecologically grounded approaches, including the science of magic and Theory of Mind, to better understand how early-life cognition shapes later behaviour, cognition and fitness in this model avian family.

animal behavior and cognition↗

bioRxiv2024

Reasoning by exclusion in food-caching Eurasian jays (Garrulus glandarius)

Two related methods have been widely used to test animals capacity to reason by exclusion: Calls (2004) 2-cups task, where subjects choose between two cups in which one reward was hidden and the empty cup is revealed; and Premack & Premacks task (1994) where one reward goes in each of the two cups and one reward is removed. In both cases, success is identifying the cup with the reward. It has been suggested that among corvids, the demands associated with caching may foster exclusion abilities by reinforcing the experience of relevant contraries (e.g., empty vs. full), or by supporting domain general capacities that support caching and exclusion inferences. However, Shaw et al. (2013) found that Eurasian jays, proficient cachers, failed a version of Calls 2-cups task. To further test the caching hypothesis, we amended the procedure using invisible displacement to hide rewards, removing aspects of Shaw et al.s procedure that may have masked competence of Eurasian jays. We tested seven Eurasian jays, who showed spontaneous success on both Calls 1-reward/show-empty (80% correct) and Premack and Premacks 2-reward/remove-1 (85% correct). These results align with the hypothesis that corvids exclusion reasoning is related to caching behaviour.

animal behavior and cognition↗

bioRxiv2024

Can we trace the social affiliation of rooks (Corvusfrugilegus) through their vocal signature?

Inter-individual recognition is crucial for stable social relationships and it is frequently mediated through vocal signatures. In socially complex species, recognition may additionally require additional levels corresponding to other layers of social organisation such as the pair, family, social group or colony. Additional vocal signatures may encode these different levels of social organisations for recognition. We investigated this hypothesis in the calls of the rook (Corvus frugilegus), a highly social corvid. Rooks form large breeding colonies where multiple pairs nest in clusters. We recorded the calls of five colonies located in France and in Great Britain, including both wild and captive colonies. To exclude variations due to different call types, we focused on the loud nest call produced exclusively by nesting females during the breeding season. We compared the acoustic distance of calls from each individual and between individuals at various levels of nest proximity, i.e. from the same nest cluster, from different nest clusters, from colonies within the same country, and from colonies in different countries. The only vocal signatures we found were at the individual level, but not at the nest cluster or colony level. This suggests a lack of vocal convergence in this species, at least for the nest call, which may be important for pair recognition in large colonies. Further studies should now evaluate if types of calls other than the nest call better carry vocal signatures as markers of different layers of sociality in this species, or if vocal divergence is a more general vocal phenomenon. In that case, applying new methods of monitoring vocal signatures in wild individuals should help understand the cognitive, social and environmental mechanisms underlying this vocal singularisation. 1. Significance statementInter-individual recognition is crucial for social relationships in animals, and is often mediated by individual-specific acoustic characteristics in vocalisations, called a vocal signature. High levels of social organisations, such as a social group of familiar conspecifics or a breeding colony, may likewise be signalled by vocal signatures shared by multiple individuals. We used machine-learning techniques to investigate vocal signatures at multiple social levels in the nest call of brooding female rooks, a corvid species that breeds colonially but lives year-round in social groups. We find evidence of a strong individual vocal signature, but no common vocal signature even in females that nest close together, or in the same colony. A strong individual vocal signature may be a potent tool to monitor populations in this species with minimal disturbance and minimal material, especially as corvids are frequently targeted by human-fauna conflicts in continental Europe.

animal behavior and cognition↗

bioRxiv2023

Social influences on delayed gratification in New Caledonian crows and Eurasian jays

Self-control underlies goal-directed behaviour in humans and other animals. Delayed gratification - a measure of self-control - requires the ability to tolerate a delay and/or invest more effort to obtain a reward of higher value over one of lower value, such as food or mates. Social context, in particular, the presence of competitors, may influence delayed gratification. We adapted the rotating-tray paradigm, where subjects need to forgo an immediate, lower-quality (i.e. less preferred) reward for a delayed, higher-quality (i.e. more preferred) one, to test social influences on delayed gratification in two corvid species: New Caledonian crows and Eurasian jays. We compared choices for immediate vs. delayed rewards while alone, in the presence of a competitive conspecific and in the presence of a non-competitive conspecific. We found that species differed: jays were more likely to select the immediate, less preferred reward than the crows. We also found that jays were more likely to select the immediate, less preferred reward when a competitor or non-competitor was present than when alone, or when a competitor was present compared to a non- competitor, while the crows selected the delayed, highly preferred reward irrespective of social presence. We discuss our findings in relation to species differences in socio-ecological factors (adult sociality and food caching) and wider implications of social influences on self-control.

animal behavior and cognition↗

bioRxiv2021

Individual repeatability, species differences, and the influence of socio-ecological factors on neophobia in 10 corvid species

Behavioural responses to novelty, including fear and subsequent avoidance of novel stimuli, behaviours referred to as neophobia, determine how animals interact with their environment. Neophobia aids in navigating risk and impacts on adaptability and survival. There is variation within and between individuals and species, however, lack of large-scale, comparative studies critically limits investigation of the socio-ecological drivers of neophobia. In this study, we tested responses to novel objects and food (alongside familiar food) versus a baseline (familiar food alone) in 10 corvid species (241 subjects) across 10 labs worldwide. There were species differences in the latency to touch familiar food in the novel object and food conditions relative to the baseline. Three of seven socio-ecological factors influenced object neophobia: 1) use of urban habitat (vs not), 2) territorial pair vs family group sociality and 3) large vs small flock size (whereas range, caching, hunting live animals, and genus did not); while only flock size influenced food neophobia. We found that, overall, individuals were temporally and contextually repeatable (i.e. consistent) in their novelty responses in all conditions, indicating neophobia is a stable behavioural trait. With this study, we have established a network of corvid researchers, demonstrating potential for further collaboration to explore the evolution of cognition in corvids and other bird species. These novel findings enable us, for the first time in corvids, to identify the socio-ecological correlates of neophobia and grant insight into specific elements that drive higher neophobic responses in this avian family group.

animal behavior and cognition↗

bioRxiv2021

Little evidence that Eurasian jays (Garrulus glandarius) protect their caches by responding to cues about a conspecific's desire and visual perspective

Corvids appear to be capable of adjusting their behaviour according to anothers perspective, knowledge and desire. For example, Eurasian jays have been found to employ a variety of cache protection strategies to minimise cache loss by responding to cues about the visual perspective or current desire of an observing conspecific. However, it is not known whether these jays (or any other corvid) can integrate multiple cues about different mental states and perform the optimal response accordingly. Across five experiments, we found little evidence that our Eurasian jays responded to either the visual perspective or current desire of another agent. In Experiments 1 and 2 we investigated whether Eurasian jays can limit the risk of cache loss by responding simultaneously to cues about the desire and perspective of a potential conspecific pilferer. Building on established paradigms, we used opaque and clear barriers to manipulate the observers visual access to cache locations, and specific satiety to manipulate the observers desire towards different types of food. Across both experiments the jays caching pattern provided no evidence that they could integrate information about the observers desire and perspective. Moreover, the results were also inconsistent with the previously reported effects that jays protect their caches by responding to either the visual access or specific satiety of the observer independently. To gain further insight into these unexpected results, we conducted three more experiments. In Experiments 3 and 4, we attempted to replicate the previous finding that Eurasian jays prefer to cache behind an opaque barrier over a clear barrier when observed by a conspecific than when caching in private. In Experiment 5, we further investigated the previous finding that jays preferentially cache a type of food that had been eaten to satiety by a conspecific over a food that had not been eaten by the conspecific. Experiments 3, 4 and 5 found no significant effects in the direction of the previously reported effects, questioning their robustness. We conclude by discussing the implications of our study for the field of corvid cognition and highlight several key issues that affect the reliability of comparative cognition research.

animal behavior and cognition↗

bioRxiv2020

Testing two competing hypotheses for Eurasian jays' caching for the future: planning versus compensatory caching

Previous research reported that corvids preferentially cache food in a location where no food will be available or cache more of a specific food in a location where this food will not be available. Here, we consider possible explanations for these prospective caching behaviours and directly compare two competing hypotheses. The Compensatory Caching Hypothesis suggests that birds learn to cache more of a particular food in places where that food was less frequently available in the past. In contrast, the Future Planning Hypothesis suggests that birds recall what-when-where features of specific past events to predict the future availability of food. We designed a protocol in which the two hypotheses predict different caching patterns across different caching locations such that the two explanations can be disambiguated. We formalised the hypotheses in a Bayesian model comparison and tested this protocol in two experiments with one of the previously tested species, namely Eurasian jays. Consistently across the two experiments, the observed caching pattern did not support either hypothesis; rather it was best explained by a uniform distribution of caches over the different caching locations. Future research is needed to gain more insight into the cognitive mechanism underpinning corvids caching for the future.

animal behavior and cognition↗

bioRxiv2019

How flexible is tool use in Eurasian jays (Garrulus glandarius)?

Eurasian jays (Garrulus glandarius) do not habitually use tools, yet they can be trained to solve object-dropping tasks, i.e. to insert a tool into an apparatus to release a food reward. Previous research suggests the these jays can learn a preference toward functional tools - objects allowing to obtain a food reward placed inside an apparatus - according to their density (Cheke et al., 2011). However, it is not yet known whether they can also select functional tools (tool selectivity) according to other physical properties such as size and shape, and use different kinds of tools to solve a similar task. Here we conducted three object-dropping experiments aimed at exploring these abilities in Eurasian jays. In Experiment 1, jays tended to select large stones as tools irrespective of the diameter of the apparatus. However, jays progressively developed a preference for the small tool, which was functional with both the wide and the narrow apparatuses. In Experiment 2, only vertically-oriented long stones could fit into the narrow apparatus, whereas both long and round stones were functional with the wide apparatus. Jays showed a preference for the long stone and, with the narrow apparatus, tended to achieve the correct manipulation after one or more unsuccessful attempts. In Experiment 3, jays were able to use sticks and adopt a novel technique on the same object-dropping apparatus, thus providing the first evidence that Eurasian jays can use sticks as tools. Taken together, these results indicate that Eurasian jays may have limited tool selectivity abilities but nonetheless can use different kinds of tools to solve similar tasks.

animal behavior and cognition↗

bioRxiv2019

Assembly of novel, nuclear dimers of the PI3-Kinase regulatory subunits underpins the pro-proliferative activity of the Cdc42-activated tyrosine kinase, ACK

The tyrosine kinase ACK is an oncogene associated with poor prognosis in human cancers. ACK promotes proliferation, in part, by contributing to the activation of Akt, the major PI3-Kinase effector. We show that ACK also regulates PI3-Kinase directly, via interactions with the PI3-Kinase regulatory subunits. ACK interacts with all five regulatory subunit isoforms and directly phosphorylates p85, p85{beta}, p55 and p50 on Tyr607 (or equivalent). Phosphorylation of p85{beta} at this residue promotes cell proliferation but, counterintuitively, ACK does not stimulate PI3-Kinase catalytic activity. We show that ACK stabilizes p85 levels by promoting an interaction between the p85 nSH2 domain and pTyr607, protecting p85 from ubiquitination. We demonstrate that ACK interacts with p85 exclusively in nuclear-enriched cell fractions where the increased levels of the regulatory subunits, together with the nSH2-pTyr607 interaction, promote formation of dimeric p85. We postulate that these novel dimers undertake nuclear functions that contribute to Cdc42-ACK driven oncogenesis. We propose that ACK shapes PI3-Kinase signalling by dampening the PIP3 response, whilst continuing to drive cell proliferation through Akt activation and hereto unexplored but crucial functions of nuclear dimeric p85. These new regulatory subunit dimers represent a previously undescribed mode of regulation for PI3-Kinase and potentially reveal additional avenues for therapeutic intervention.

molecular biology↗

bioRxiv2019

Decision-making flexibility in New Caledonian crows, young children and adult humans in a multi-dimensional tool-use task

The ability to make profitable decisions in natural foraging contexts may be influenced by an additional requirement of tool-use, due to increased levels of relational complexity and additional work-effort imposed by tool-use, compared with simply choosing between an immediate and delayed food item. We examined the flexibility for making the most profitable decisions in a multi-dimensional tool-use task, involving different apparatuses, tools and rewards of varying quality, in 3-5-year-old children, adult humans and tool-making New Caledonian crows (Corvus moneduloides). We also compared our results to previous studies on habitually tool-making orangutans (Pongo abelii) and non-tool-making Goffins cockatoos (Cacatua goffiniana). Adult humans, cockatoos and crows, though not children and orangutans, did not select a tool when it was not necessary, which was the more profitable choice in this situation. Adult humans, orangutans and cockatoos, though not crows and children, were able to refrain from selecting non-functional tools. By contrast, the birds, though not primates tested, struggled to attend to multiple variables - where two apparatuses, two tools and two reward qualities were presented simultaneously - without extended experience. These findings indicate: (1) in a similar manner to humans and orangutans, New Caledonian crows and Goffins cockatoos can flexibly make profitable decisions in some decision-making tool-use tasks, though the birds may struggle when tasks become more complex; (2) children and orangutans may have a bias to use tools in situations where adults and other tool-making species do not.

animal behavior and cognition↗