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In a previous study, we found that Positive Reinforcement Training reduced cortisol of wolves and dogs; however, this effect varied across trainer–animal dyads. Here we investigate whether and how the trainers’ use of speech may contribute to this effect. Dogs’ great interest in high-pitched, intense speech (also known as Dog Directed Speech) has already been reported, but whether and how wolves respond similarly/differently to voice characteristics has never been studied before. We analyzed 270 training sessions, conducted by five trainers, with nine mixed-breed dogs and nine wolves, all human-socialized. Through Generalized Linear Mixed Models, we analyzed the effects of (a) three speech categories (nice, neutral, reprehensive) and laugh; and (b) acoustic characteristics of trainers’ voices on animals’ responses (correct responses, latency, orientation, time at less than 1 m, non-training behaviors, tail position/movements, cortisol variation). In both subspecies, tail wagging occurred more often in sessions with longer durations of nice speech, and less often in sessions with reprehensive speech. For dogs, the duration of reprehensive speech within a session was also negatively related to correct responses. For wolves, retreat time was associated with more reprehensive speech, whereas duration of nice speech was positively associated with time spent within one meter from the trainer. In addition, most dog behavioral responses were associated with higher average intonations within sessions, while wolf responses were correlated with lower intonations within sessions. We did not find any effects of the variables considered on cortisol variation. Our study highlights the relevance of voice tone and speech in a training context on animals’ performances and emotional reactions.

Hybridization is an important mechanism of evolution. While hybrids often express inferior traits and are selected against, hybridization can promote phenotypic variation and produce trait combinations distinct from the parentals, generating novel adaptive potential. Among other traits, hybridization can impact behavior and cognition and may reinforce species boundaries when hybrids show decreased cognitive abilities. However, the hypothesized role of hybridization in the diversification of cognitive phenotypes remains enigmatic. To test this idea, we compare the performance of female guppies (iPoecilia reticulata/i), Endler's guppies (iPoecilia wingei/i), and their experimental hybrids in color association and reversal learning. In addition, we introduce a new approach to compare multidimensional cognitive phenotypes. We found that hybrids showed intermediate learning abilities in both tasks compared with the parentals. Moreover, hybrids had slightly higher phenotypic dispersion, new trait combinations occurred in some hybrid individuals, and the mean phenotype of one hybrid group deviated away from the axis of variation of the parentals. Our method should hence be useful in further exploring how hybridization and other evolutionary processes impact behavioral and cognitive traits. Our results suggest that hybridization may promote cognitive variation and generate new trait combinations, even when learning performance at the group level is intermediate between parentals.

Despite ongoing advances in sexual selection theory, the evolution of mating decisions remains enigmatic. Cognitive processes often require simultaneous processing of multiple sources of information from environmental and social cues. However, little experimental data exist on how cognitive ability affects such fitness-associated aspects of behaviour. Using advanced tracking techniques, we studied mating behaviours of guppies artificially selected for divergence in relative brain size, with known differences in cognitive ability, when predation threat and sex ratio was varied. In females, we found a general increase in copulation behaviour in when the sex ratio was female biased, but only large-brained females responded with greater willingness to copulate under a low predation threat. In males, we found that small-brained individuals courted more intensively and displayed more aggressive behaviours than large-brained individuals. However, there were no differences in female response to males with different brain size. These results provide further evidence of a role for female brain size in optimal decision-making in a mating context. In addition, our results indicate that brain size may affect mating display skill in male guppies. We suggest that it is important to consider the association between brain size, cognitive ability and sexual behaviour when studying how morphological and behavioural traits evolve in wild populations.

Cognitive ageing is the general process when certain mental skills gradually deteriorate with age. Across species, there is a pattern of a slower brain structure degradation rate in large-brained species. Hence, having a larger brain might buffer the impact of cognitive ageing and positively affect survival at older age. However, few studies have investigated the link between relative brain size and cognitive ageing at the intraspecific level. In particular, experimental data on how brain size affects brain function also into higher age is largely missing. We used 288 female guppies (Poecilia reticulata), artificially selected for large and small relative brain size, to investigate variation in colour discrimination and behavioural flexibility, at 4–6, 12 and 24 months of age. These ages are particularly interesting since they cover the life span from sexual maturation until maximal life length under natural conditions. We found no evidence for a slower cognitive ageing rate in large-brained females in neither initial colour discrimination nor reversal learning. Behavioural flexibility was predicted by large relative brain size in the youngest group, but the effect of brain size disappeared with increasing age. This result suggests that cognitive ageing rate is faster in large-brained female guppies, potentially due to the faster ageing and shorter lifespan in the large-brained selection lines. It also means that cognition levels align across different brain sizes with older age. We conclude that there are cognitive consequences of ageing that vary with relative brain size in advanced learning abilities, whereas fundamental aspects of learning can be maintained throughout the ecologically relevant life span.

Photography is ubiquitous worldwide. We analyzed why people take, share, and use personal photographs, independent of their specific cultural background. These behaviors are still poorly understood. Experimental research on them is scarce. Smartphone technology and social media have pushed the success of photography, but cannot explain it, as not all smartphone features are widely used just because they are available. We analyzed properties of human nature that have made taking and using photographs functional behaviors. We did this based on the four levels, which Nikolaas Tinbergen suggested for analyzing why animals behave in a particular way. Including findings from multiple disciplines, we developed a novel conceptual framework—the “Mental Utilization Hypothesis of Photography.” It suggests that people adopt photography because it matches with core human mental mechanisms mainly from the social domain, and people use photography as a cognitive, primarily social coping strategy. Our framework comprises a range of testable predictions, provides a new theoretical basis for future empirical investigations into photography, and has practical implications. We conclude that photography has become a human universal, which is based on context-sensitive mental predispositions and differentiates itself in the social and societal environment.

Predation risk during early ontogeny can impact developmental trajectories and permanently alter adult phenotypes. Such phenotypic plasticity often leads to adaptive changes in traits involved in anti-predator responses. While plastic changes in cognition may increase survival, it remains unclear whether early predation experience shapes cognitive investment and drives developmental plasticity in cognitive abilities. Here, we show that predation risk during early ontogeny induces developmental plasticity in two cognitive domains. We reared female guppies (Poecilia reticulata) with and without predator cues and tested their adult cognitive abilities. We found that females reared under simulated predation took longer to learn a simple association task, yet outperformed animals reared without predation threat in a reversal learning task testing cognitive flexibility. These results show that predation pressure during ontogeny shapes adult cognitive abilities, which we argue is likely to be adaptive. Our study highlights the important role of predator-mediated developmental plasticity on cognitive investment in natural populations and the general role of plasticity in cognitive performance.

Understanding the basis of behavior requires dissecting the complex waves of gene expression that underlie how the brain processes stimuli and produces an appropriate response. In order to determine the dynamic nature of the neurogenomic network underlying mate choice, we use transcriptome sequencing to capture the female neurogenomic response in two brain regions involved in sensory processing and decision-making under different mating and social contexts. We use differential coexpression (DC) analysis to evaluate how gene networks in the brain are rewired when a female evaluates attractive and nonattractive males, greatly extending current single-gene approaches to assess changes in the broader gene regulatory network. We find the brain experiences a remarkable amount of network rewiring in the different mating and social contexts we tested. Further analysis indicates the network differences across contexts are associated with behaviorally relevant functions and pathways, particularly learning, memory and other cognitive functions. Finally, we identify the loci that display social context-dependent connections, revealing the basis of how relevant neurological and metabolic pathways are differentially recruited in distinct social contexts. More broadly, our findings contribute to our understanding of the genetics of mating and social behavior by identifying gene drivers behind behavioral neural processes, illustrating the utility of DC analysis in neurosciences and behavior.

Reversal learning assays are commonly used across a wide range of taxa to investigate associative learning and behavioural flexibility. In serial reversal learning, the reward contingency in a binary discrimination is reversed multiple times. Performance during serial reversal learning varies greatly at the interspecific level, as some animals adopt a rule-based strategy that enables them to switch quickly between reward contingencies. A larger relative brain size, generating enhanced learning ability and increased behavioural flexibility, has been proposed to be an important factor underlying this variation. Here, we experimentally tested this hypothesis at the intraspecific level. We used guppies (Poecilia reticulata) artificially selected for small and large relative brain size, with matching differences in neuron number, in a serial reversal learning assay. We tested 96 individuals over 10 serial reversals and found that learning performance and memory were predicted by brain size, whereas differences in efficient learning strategies were not. We conclude that variation in brain size and neuron number is important for variation in learning performance and memory, but these differences are not great enough to cause the larger differences in efficient learning strategies observed at higher taxonomic levels., Summary: Relative brain size affects differences in advanced learning ability in small- and large-brained female guppies; however, regardless of brain size, they do not learn a generalized rule from earlier experience.

Collective movement is achieved when individuals adopt local rules to interact with their neighbours. How the brain processes information about neighbours’ positions and movements may affect how individuals interact in groups. As brain size can determine such information processing it should impact collective animal movement. Here we investigate whether brain size affects the structure and organisation of newly forming fish shoals by quantifying the collective movement of guppies (Poecilia reticulata) from large- and small-brained selection lines, with known differences in learning and memory. We used automated tracking software to determine shoaling behaviour of single-sex groups of eight or two fish and found no evidence that brain size affected the speed, group size, or spatial and directional organisation of fish shoals. Our results suggest that brain size does not play an important role in how fish interact with each other in these types of moving groups of unfamiliar individuals. Based on these results, we propose that shoal dynamics are likely to be governed by relatively basic cognitive processes that do not differ in these brain size selected lines of guppies.

Competition over mates is a powerful force shaping trait evolution. For instance, better cognitive abilities may be beneficial in male-male competition and thus be selected for by intrasexual selection. Alternatively, investment in physical attributes favoring male performance in competition for mates may lower the resources available for brain development, and more intense male mate competition would coincide with smaller brains. To date, only indirect evidence for such relationships exists, and most studies are heavily biased toward primates and other homoeothermic vertebrates. We tested the association between male brain size (relative to body size) and male-male competition across N = 30 species of Chinese anurans. Three indicators of the intensity of male mate competition—operational sex ratio (OSR), spawningsite density, and male forelimb muscle mass—were positively associated with relative brain size, whereas the absolute spawning group size was not. The relationship with the OSR and male forelimb muscle mass was stronger for the male than for the female brains. Taken together, our findings suggest that the increased cognitive abilities of larger brains are beneficial in male-male competition. This study adds taxonomic breadth to the mounting evidence for a prominent role of sexual selection in vertebrate brain evolution.

Mind attribution to nonhuman agents is claimed to be the base of ascribing moral rights to them. It is sufficiently common to be considered a human universal, and the way it is done is related with a range of personal and animal-related factors. The aim of the present study was to identify the folk representations that underlie the attribution of emotions and complex mental states to two local primate species and a range of wild and domestic animals. For this, we conducted a series of semi-structured interviews to a convenience sample of 23 women and men from the community of Conhuas, Campeche, situated near the Biosphere Reserve of Calakmul in southern Mexico. Data from the interviews were analyzed via a qualitative content analysis. We found that the nature of mental states ascribed to animals is based on a set of triggering factors that include behavioral expressions, animal characteristics, and circumstances in which these behaviors occur that altogether point to an underlying mental state. We discuss these results considering the potential cognitive processes involved in these attributions.

Species can evolve diverse strategies to survive periods of uncertainty. Animals may either invest in energy storage, allowing them to decrease foraging costs, such as locomotion or risk of predation, or they may invest in better cognitive abilities helping them to flexibly adapt their behavior to meet novel challenges. Here, we test this idea of a fat-brain trade-off in 38 species of Chinese anurans by relating the coefficient of variation of body mass (CVbodymass ; as an indicator of how much animals invest into storage over the season) to brain anatomical features. After correcting for shared ancestry and body mass, we found a negative relationship between relative brain size and CVbodymass . This indicates that anurans seem to trade-off physiological and cognitive buffering during energy shortages. As similar patterns have been reported in arboreal mammals and primates our findings suggest that the fat-brain trade-off, where animals either invest into physiological or cognitive strategies to survive harsh conditions, may be a general pattern across vertebrates.

The evolution of collective behaviour has been proposed to have important effects on individual cognitive abilities. Yet, in what way they are related remains enigmatic. In this context, the ‘distributed cognition’ hypothesis suggests that reliance on other group members relaxes selection for individual cognitive abilities. Here, we test how cognitive processes respond to evolutionary changes in collective motion using replicate lines of guppies (Poecilia reticulata) artificially selected for the degree of schooling behaviour (group polarization) with >15% difference in schooling propensity. We assessed associative learning in females of these selection lines in a series of cognitive assays: colour associative learning, reversal-learning, social associative learning, and individual and collective spatial associative learning. We found that control females were faster than polarization selected females at fulfilling a learning criterion only in the colour associative learning assay, but they were also less likely to reach a learning criterion in the individual spatial associative learning assay. Hence, although testing several cognitive domains, we found weak support for the distributed cognition hypothesis. We propose that any cognitive implications of selection for collective behaviour lie outside of the cognitive abilities included in food-motivated associative learning for visual and spatial cues.

The relationship between brain size and ageing is a paradox. The cognitive benefits of large brains should protect from extrinsic mortality and thus indirectly select for slower ageing. However, the substantial energetic cost of neural tissue may also impact the energetic budget of large-brained organisms, causing less investment in somatic maintenance and thereby faster ageing. While the positive association between brain size and survival in the wild is well established, no studies exist on the direct effects of brain size on ageing. Here we test how brain size influences intrinsic ageing in guppy ( Poecilia reticulata) brain size selection lines with 12% difference in relative brain size. Measuring survival under benign conditions, we find that large-brained animals live 22% shorter than small-brained animals and the effect is similar in both males and females. Our results suggest a trade-off between investment into brain size and somatic maintenance. This implies that the link between brain size and ageing is contingent on the mechanism of mortality, and selection for positive correlations between brain size and ageing should occur mainly under cognition-driven survival benefits from increased brain size. We show that accelerated ageing can be a cost of evolving a larger brain.

Despite the common assumption that the brain is mainly malleable to surrounding conditions during ontogeny, plastic neural changes can occur also in adulthood. One of the driving forces responsible for alterations in brain morphology is increasing environmental complexity that may demand for enhanced cognitive abilities (e.g. attention, memory and learning). However, studies looking at the relationship between brain morphology and learning are scarce. Here, we tested the effects of both learning and environmental enrichment on neural plasticity in guppies (Poecilia reticulata), by means of either a reversal-learning test or a spatial-learning test. Given considerable evidence supporting environmentally-induced plastic alterations, two separate control groups that were not subjected to any cognitive test were included to account for potential changes induced by the experimental setup alone. We did not find any effect of learning on any of our brain measurements. However, we found strong evidence for an environmental effect, where fish given access to the spatial-learning environment had larger relative brain size and optic tectum size in relation to those exposed to the reversal-learning environment. Our results demonstrate the plasticity of the adult brain to respond adaptively mainly to environmental conditions, providing support for the environmental enhancement theory.

Neurons are the basic computational units of the brain, but brain size is the predominant surrogate measure of brain functional capacity in comparative and cognitive neuroscience. This approach is based on the assumption that larger brains harbor higher numbers of neurons and their connections, and therefore have a higher information‐processing capacity. However, recent studies have shown that brain mass may be less strongly correlated with neuron counts than previously thought. Till now, no experimental test has been conducted to examine the relationship between evolutionary changes in brain size and the number of brain neurons. Here, we provide such a test by comparing neuron number in artificial selection lines of female guppies (Poecilia reticulata) with >15% difference in relative brain mass and numerous previously demonstrated cognitive differences. Using the isotropic fractionator, we demonstrate that large‐brained females have a higher overall number of neurons than small‐brained females, but similar neuronal densities. Importantly, this difference holds also for the telencephalon, a key region for cognition. Our study provides the first direct experimental evidence that selection for brain mass leads to matching changes in number of neurons and shows that brain size evolution is intimately linked to the evolution of neuron number and cognition.

We published a study recently testing the link between brain size and behavioural plasticity using brain size selected guppy (Poecilia reticulata) lines (2019, Journal of Evolutionary Biology, 32, 218-226). Only large-brained fish showed habituation to a new, but actually harmless environment perceived as risky, by increasing movement activity over the 20-day observation period. We concluded that "Our results suggest that brain size likely explains some of the variation in behavioural plasticity found at the intraspecific level". In a commentary published in the same journal, Haave-Audet et al. challenged the main message of our study, stating that (a) relative brain size is not a suitable proxy for cognitive ability and (b) habituation measured by us is likely not adaptive and costly. In our response, we first show that a decade's work has proven repeatedly that relative brain size is indeed positively linked to cognitive performance in our model system. Second, we discuss how switching from stressed to unstressed behaviour in stressful situations without real risk is likely adaptive. Finally, we point out that the main cost of behavioural plasticity in our case is the development and maintenance of the neural system needed for information processing, and not the expression of plasticity. We hope that our discussion with Haave-Audet et al. helps clarifying some central issues in this emerging research field.

Understanding how animal personality (consistent between-individual behavioural differences) arises has become a central topic in behavioural sciences. This endeavour is complicated by the fact that not only the mean behaviour of individuals (behavioural type) but also the strength of their reaction to environmental change (behavioural plasticity) varies consistently. Personality and cognitive abilities are linked, and we suggest that behavioural plasticity could also be explained by differences in brain size (a proxy for cognitive abilities), since accurate decisions are likely essential to make behavioural plasticity beneficial. We test this idea in guppies (Poecilia reticulata), artificially selected for large and small brain size, which show clear cognitive differences between selection lines. To test whether those lines differed in behavioural plasticity, we reared them in groups in structurally enriched environments and then placed adults individually into empty tanks, where we presented them daily with visual predator cues and monitored their behaviour for 20 days with video-aided motion tracking. We found that individuals differed consistently in activity and risk-taking, as well as in behavioural plasticity. In activity, only the large-brained lines demonstrated habituation (increased activity) to the new environment, whereas in risk-taking, we found sensitization (decreased risk-taking) in both brain size lines. We conclude that brain size, potentially via increasing cognitive abilities, may increase behavioural plasticity, which in turn can improve habituation to novel environments. However, the effects seem to be behaviour-specific. Our results suggest that brain size likely explains some of the variation in behavioural plasticity found at the intraspecific level.

BackgroundBrain size is expected to evolve by a balance between cognitive benefits and energetic costs. Several influential hypotheses have suggested that large brains may be especially beneficial in social contexts. Group living and competition may pose unique cognitive challenges to individuals and favor the evolution of increased cognitive ability. Evidence comes from comparative studies on the link between social complexity and brain morphology, but the strength of empirical support has recently been challenged. In addition, the behavioral mechanisms that would link cognitive ability to sociality are rarely studied. Here we take an alternative approach and investigate experimentally how brain size can relate to the social competence of individuals within species, a problem that so far has remained unresolved. We use the unique guppy brain size selection line model system to evaluate whether large brains are advantageous by allowing individuals to better assess their performance in a social contest situation. Based on theoretical literature, we predict that contest duration should depend on the brain size of the loser, as it is the capitulation of the losing individual that ends the fight.ResultsFirst, we show that studying the movement of competitors during contests allows for precise estimation of the dominance timeline in guppies, even when overt aggression is typically one-sided and delayed. Second, we staged contests between pairs of male that had been artificially selected for large and small relative brain size, with demonstrated differences in cognitive ability. We show that dominance was established much earlier in contests with large-brained losers, whereas the brain size of the winner had no effect. Following our prediction, large-brained individuals gave up more quickly when they were going to lose.ConclusionsThese results suggest that large-brained individuals assess their performance in contests better and that social competence indeed can depend on brain size. Conflict resolution may therefore be an important behavioral mechanism behind macro-evolutionary patterns between sociality and brain size. Since conflict is ubiquitous among group-living animals, the possible effects of the social environment on the evolution of cognition may be more broadly applicable than previously thought.

Food preferences may be driven by a species’ ecology. Closely related species such as dogs and wolves may have evolved preferences for different foods owing to their differing foraging styles. Wolves have been shown to be more persistent in problem-solving experiments and more risk-prone in a foraging task. A possible element affecting these (and other) results is a potential wolf-dog difference in food preferences. To address this possibility, we tested similarly raised and kept dogs and wolves in two different food choice tasks, a classic two-choice task and a multiple-choice paradigm. We predicted that if dogs have adapted to a more opportunistic, scavenging foraging style, they would show a weaker preference for meat over starch rich foods (such as kibble) and be less affected by hunger than wolves. Alternatively, given the recentness of the new niche dogs have created, we predicted no substantial differences between dogs’ and wolves’ food preferences. We found that our subjects did not differ in their preference for meat over kibble in either paradigm. However, wolves’ (but not dogs’) choice patterns were affected by satiation, with wolves being less “selective” when hungry. Furthermore, when fed before testing, wolves were more selective than dogs. These differences were more noticeable in the multiple-choice paradigm than the two-choice task, suggesting that the former, novel paradigm may be more sensitive and better capable of evaluating food preferences in a diverse range of species. Overall, we found that the distinct differences in wolves’ and dogs’ ecology and foraging styles do not appear to have affected their food preferences and thus, differences in food preferences are unlikely to have influenced results of previous experiments demonstrating wolf-dog differences in cognitive skills.

Understanding what drives animal decisions is fundamental in evolutionary biology, and mate choice decisions are arguably some of the most important decisions in any individual's life. As cognitive ability can impact decision-making, elucidating the link between mate choice and cognitive ability is necessary to fully understand mate choice. To experimentally study this link, we used guppies (Poecilia reticulata) artificially selected for divergence in relative brain size and with previously demonstrated differences in cognitive ability. A previous test in our female guppy selection lines demonstrated the impact of brain size and cognitive ability on information processing during female mate choice decisions. Here we evaluated the effect of brain size and cognitive ability on male mate choice decisions. Specifically, we investigated the preferences of large-brained, small-brained, and non-selected guppy males for female body size, a key indicator of female fecundity in this species. For this, male preferences were quantified in dichotomous choice tests when presented to dyads of females with small, medium and large body size differences. All types of males showed preference for larger females but no effect of brain size was found in the ability to discriminate between differently sized females. However, we found that non-selected and large-brained males, but not small-brained males, showed context-dependent preferences for larger females depending on the difference in female size. Our results have two important implications. First, they provide further evidence that male mate choice occurs also in a species in which secondary sexual ornamentation occurs only in males. Second, they show that brain size and cognitive ability have important effects on individual variation in mating preferences and sexually selected traits.

It has become increasingly clear that a larger brain can confer cognitive benefits. Yet not all of the numerous aspects of cognition seem to be affected by brain size. Recent evidence suggests that some more basic forms of cognition, for instance colour vision, are not influenced by brain size. We therefore hypothesize that a larger brain is especially beneficial for distinct and gradually more complex aspects of cognition. To test this hypothesis, we assessed the performance of brain size selected female guppies ( Poecilia reticulata ) in two distinct aspects of cognition that differ in cognitive complexity. In a standard reversal-learning test we first investigated basic learning ability with a colour discrimination test, then reversed the reward contingency to specifically test for cognitive flexibility. We found that large-brained females outperformed small-brained females in the reversed-learning part of the test but not in the colour discrimination part of the test. Large-brained individuals are hence cognitively more flexible, which probably yields fitness benefits, as they may adapt more quickly to social and/or ecological cognitive challenges. Our results also suggest that a larger brain becomes especially advantageous with increasing cognitive complexity. These findings corroborate the significance of brain size for cognitive evolution.

Predation is a near ubiquitous factor of nature and a powerful selective force on prey. Moreover, it has recently emerged as an important driver in the evolution of brain anatomy, though population comparisons show ambiguous results with considerable unexplained variation. Here, we test the reproducibility of reduced predation on evolutionary trajectories of brain evolution. We make use of an introduction experiment, whereby guppies ( Poecilia reticulata ) from a single high predation stream were introduced to four low predation streams. After 8–9 years of natural selection in the wild and two generations of common garden conditions in the laboratory, we quantified brain anatomy. Relative brain region sizes did not differ between populations. However, we found a general increase and striking variation in relative brain size of introduced populations, which varied from no change to a 12.5% increase in relative brain weight, relative to the ancestral high predation population. We interpret this as evidence for non-parallel evolution, which implies a weak or inconsistent association of relative brain size with fitness in low predation sites. The evolution of brain anatomy appears sensitive to unknown environmental factors, or contingent on either chance events or historical legacies of environmental change.

Self-control, that is, overcoming impulsivity towards immediate gratification in favour of a greater but delayed reward, is seen as a valuable skill when making future-oriented decisions. Experimental studies in nonhuman primates revealed that individuals of some species are willing to tolerate delays of up to several minutes in order to gain food of a higher quantity or quality. Recently, birds (carrion crows, Corvus corone, common ravens, Corvus corax, Goffin cockatoos, Cacatua goffiniana) performed comparably to primates in an exchange task, contradicting previous notions that birds may lack any impulse control. However, performance differed strikingly with the currency of exchange: individuals of all three species performed better when asked to wait for a higher food quality, rather than quantity. Here, we built on this work and tested whether the apparent difference in levels of self-control expressed in quality versus quantity tasks reflects cognitive constraints or is merely due to methodological limitations. In addition to the exchange paradigm, we applied another established delay maintenance methodology: the accumulation task. In this latter task, food items accumulated to a maximum of four pieces, whereas in the exchange task, an initial item could be exchanged for a reward item after a certain time delay elapsed. In both tasks, birds (seven crows, five ravens) were asked to wait in order to optimize either the quality or the quantity of food. We found that corvids were willing to delay gratification when it led to a food reward of higher quality, but not when waiting was rewarded with a higher quantity, independent of the experimental paradigm. This study is the first to test crows and ravens with two different paradigms, the accumulation and the exchange of food, within the same experiment, allowing for fair comparisons between methods and species.

Highlights • We compared the results of corvids in experiments with familiar/unfamiliar humans. • We investigated behavioural reactions towards familiar and unfamiliar humans. • Corvids performed significantly better in experiments with familiar humans. • Corvids did not show more neophobia towards unfamiliar humans. • Hence, familiarity positively affected the experimental performance of corvids., When humans and animals interact with one another over an extended time span they familiarise and may develop a relationship, which can exert an influence on both partners. For example, the behaviour of an animal in experiments may be affected by its relationship to the human experimenter. However, few studies have systematically examined the impact of human–animal relationships on experimental results. In the present study we investigated if familiarity with a human experimenter influences the performance of Common ravens (Corvus corax) and Carrion crows (Corvus corone corone) in interactive tasks. Birds were tested in two interactive cognitive tasks (exchange, object choice) by several experimenters representing different levels of familiarity (long and short-term). Our findings show that the birds participated more often in both tasks and were more successful in the exchange task when working with long-term experimenters than when working with short-term experimenters. Behavioural observations indicate that anxiety did not inhibit experimental performance but that the birds’ motivation to work differed between the two kinds of experimenters, familiar and less familiar. We conclude that human–animal relationships (i.e. familiarity) may affect the experimental performance of corvids in interactive cognitive tasks.

It has been suggested that affiliated social relations may facilitate information transfer between individuals. We here tested this rarely examined hypothesis with juvenile and adult jackdaws (Corvus monedula) in three stimulus enhancement tasks, both in a non-food context (experiment 1) and in a food context (experiments 2 and 3). We first show that siblings and pair partners maintain stronger bonded social relations than do non-siblings and non-pair partners. We therefore tested individuals in sibling and non-sibling dyads and, later in ontogeny, in pair and non-pair dyads. Jackdaws either did not learn from any other conspecific (experiment 1), or they learned from non-affiliated individuals (non-siblings, non-pair partners in experiments 2 and 3). This may be related to two main characteristics of jackdaws' affiliated relationships. First, affiliates share food at a high rate and may rely on their knowledgeable partners to secure food rather than learning from them. Second, affiliates spend most time in close spatial proximity to each other which increases the probability that they simultaneously experience occurrences in their environment. Hence, spatially more distant individuals, which are more likely to be non-affiliated, face different foraging situations and may therefore provide more relevant information which may lead to selective social learning.

It has been suggested that social dynamics affect social learning but empirical support for this idea is scarce. Here we show that affiliate relationships among kin indeed enhance the performance of common ravens, Corvus corax, in a social learning task. Via daily behavioural protocols we first monitored social dynamics in our group of captive young ravens. Siblings spent significantly more time in close proximity to each other than did nonsiblings. We subsequently tested birds on a stimulus enhancement task in model-observer dyads composed of both siblings and nonsiblings. During demonstration the observer could watch the model manipulating one particular object (target object) in an adjacent room. After removing the model, the observer was confronted with five different objects including the former target object. Observers from sibling dyads handled the target object for significantly longer periods of time as compared with the other four available objects, whereas observers from nonsibling dyads did not show a preference for the target object. Also, siblings matched the model's decision to cache or not to cache objects significantly more often than did nonsiblings. Hence, siblings were likely to attend to both, the behaviour of the model (caching or noncaching) and object-specific details. Our results support the hypothesis that affiliate relations between individuals affect the transmission of information and may lead to directed social learning even when spatial proximity has been experimentally controlled for.

Co-orientation with others by using their gaze direction is considered to be adaptive for detecting food or predators or monitoring social interactions. Like the great apes, common ravens are capable of following human experimenters' gaze direction not only into distant space but also behind visual barriers. We investigated the ontogenetic development of these abilities by confronting birds with a human foster parent looking up (experiment 1) and behind visual barriers (experiment 3) and their modification by habituation (experiments 2 and 4). We tested a group of 12 hand-reared ravens during their first 10 months of life. Ravens responded to others' look-ups soon after fledging but could track their gaze behind a visual barrier only 4 months later, at the age they usually become independent from their parents. Furthermore, ravens quickly ceased responding to repeated look-ups by the model, but did not habituate to repeated gaze cues directed behind a barrier. Our findings support the idea that the two modes of gaze following reflect different cognitive levels in ravens and, possibly, have different functions.

Recent studies suggest the existence of primate-like cognitive abilities in corvids. Although the learning abilities of corvids in comparison to other species have been investigated before, little is known on how corvids perform on simple discrimination tasks if tested in experimental settings comparable to those that have been used for studying complex cognitive abilities. In this study, we tested a captive group of 12 ravens (Corvus corax) on four discrimination problems and their reversals. In contrast to other studies investigating learning abilities, our ravens were not food deprived and participation in experiments was voluntary. This preliminary study showed that all ravens successfully solved feature and position discriminations and several of the ravens could solve new tasks in a few trials, making very few mistakes.

Lay Summary Does a large brain make you smarter? If you are a guppy male searching for a female in a maze, it does. The association between brain size and smartness is a debated issue, largely due to the lack of experimental data. We compared guppies artificially bred for large and small brains and found that large-brained males learned the route through a spatial maze faster. These results thus support a link between brain size and smartness., Brain size varies dramatically among vertebrates, and selection for increased cognitive abilities is thought to be the key force underlying the evolution of a large brain. Indeed, numerous comparative studies suggest positive relationships between cognitively demanding aspects of behavior and brain size controlled for body size. However, experimental evidence for the link between relative brain size and cognitive ability is surprisingly scarce and to date stems from a single study on brain size selected guppies (Poecilia reticulata), where large-brained females were shown to outperform small-brained females in a numerical learning assay. Because the results were inconclusive for males in that study, we here use a more ecologically relevant test of male cognitive ability to investigate whether or not a relatively larger brain increases cognitive ability also in males. We compared mate search ability of these artificially selected large- and small-brained males in a maze and found that large-brained males were faster at learning to find a female in a maze. Large-brained males decreased the time spent navigating the maze faster than small-brained males and were nearly twice as fast through the maze after 2 weeks of training. Our results support that relatively larger brains are better also for males in some contexts, which further substantiates that variation in vertebrate brain size is generated through the balance between energetic costs and cognitive benefits.

Our ability to make logical inferences is considered as one of the cornerstones of human intelligence, fuelling investigations of reasoning abilities in non-human animals. Yet, the evidence to date is equivocal, with apes as the prime candidates to possess these skills. For instance, in a two-choice task, apes can identify the location of hidden food if it is indicated by a rattling noise caused by the shaking of a baited container. More importantly, they also use the absence of noise during the shaking of the empty container to infer that this container is not baited. However, since the inaugural report of apes solving this task, to the best of our knowledge, no comparable evidence could be found in any other tested species such as monkeys and dogs. Here, we report the first successful and instantaneous solution of the shaking task through logical inference by a non-ape species, the African grey parrot. Surprisingly, the performance of the birds was sensitive to the shaking movement: they were successful with containers shaken horizontally, but not with vertical shaking resembling parrot head-bobbing. Thus, grey parrots seem to possess ape-like cross-modal reasoning skills, but their reliance on these abilities is influenced by low-level interferences.

Other-regarding preferences are a critical feature of human cooperation but to what extent non-human animals exhibit these preferences is a matter of intense discussion. We tested whether jackdaws show prosocial behaviour (providing benefits to others at no cost to themselves) and altruism (providing benefits to others while incurring costs) with both sibling and non-sibling recipients. In the prosocial condition, a box was baited on both the actor's and the recipient's side (1/1 option), whereas another box provided food only for the actor (1/0 option). In the altruistic condition, the boxes contained food for either the actor (1/0 option) or the recipient (0/1 option). The proportion of selfish (1/0 option) and cooperative (1/1 and 0/1 option, respectively) actors' choices was significantly affected by the recipients' behaviour. If recipients approached the boxes first and positioned themselves next to the box baited on their side, trying to access the food reward (recipient-first trials), actors were significantly more cooperative than when the actors approached the boxes first and made their choice prior to the recipients' arrival (actor-first trials). Further, in recipient-first trials actors were more cooperative towards recipients of the opposite sex, an effect that was even more pronounced in the altruistic condition. Hence, at no cost to the actors, all recipients could significantly influence the actors' behaviour, whereas at high costs this could be achieved even more so by recipients of different sex. Local/stimulus enhancement is discussed as the most likely cognitive mechanism to account for these effects.

International audience; An animal's emotional responses are the result of its cognitive appraisal of a situation. This appraisal is notably influenced by the possibility of an individual to exert control over an aversive event. Although the fact that environment controllability decreases emotional responses in animals is well established, far less is known about its potential trans-generational effects. As the levels of avian yolk hormones can vary according to the mother's environment, we hypothesized that housing environment of mothers would modulate the quality of her eggs and in turn her offspring's behaviour. Two groups of female japanese quail were constituted: a group that had access to a place to hide in their home-cage (Hd, n = 20) and a group that had nowhere to hide (NoHd, n = 20) when stressed. Both groups were submitted to daily human disturbances for a twenty-day-period. Hd females produced eggs with both less testosterone and androstenedione than did NoHd females. The emotional and social reactivity of Hd female's offspring were lower and their growth wes slower than those of NoHd females' offspring. Our results show that a minor difference in housing environment had substantial effects on eggs and offspring. The presence of a shelter probably helped quail to cope with daily human disturbances, producing less reactive offspring. This transgenerational effect caused by an opportunity to hide could lead to applications in care of laboratory animals, conservation biology and animal welfare.

Background: Among birds, corvids and parrots are prime candidates for advanced cognitive abilities. Still, hardly anything is known about cognitive similarities and dissimilarities between them. Recently, exclusion has gained increasing interest in comparative cognition. To select the correct option in an exclusion task, one option has to be rejected (or excluded) and the correct option may be inferred, which raises the possibility that causal understanding is involved. However, little is yet known about its evolutionary history, as only few species, and mainly mammals, have been studied. Methodology/Principal Findings: We tested ravens and keas in a choice task requiring the search for food in two differently shaped tubes. We provided the birds with partial information about the content of one of the two tubes and asked whether they could use this information to infer the location of the hidden food and adjust their searching behaviour accordingly. Additionally, this setup allowed us to investigate whether the birds would appreciate the impact of the shape of the tubes on the visibility of food. The keas chose the baited tube more often than the ravens. However, the ravens applied the more efficient strategy, choosing by exclusion more frequently than the keas. An additional experiment confirmed this, indicating that ravens and keas either differ in their cognitive skills or that they apply them differently. Conclusion: To our knowledge, this is the first study to demonstrate that corvids and parrots may perform differently in cognitive tasks, highlighting the potential impact of different selection pressures on the cognitive evolution of these largebrained birds.