Interaction between animal personality and animal cognition

The study of animal personality has attracted considerable attention, as it has revealed a number of similarities in personality between humans and several nonhuman species. At the same time the adaptive value and evolutionary maintenance of different personalities are the subject of debate. Since Pavlov＇s work on dogs, students of comparative cognition have been aware that animals display vast individual differences on cognitive tasks, and that these differences may not be entirely accounted for differences in cognitive abilities. Here, we argue that personality is an important source of variation that may affect cognitive performance and we hypothesise mutual influences between personality and cognition across an individual＇s lifespan. In particular, we suggest that： 1） personality profiles may be markers of different cognitive styles; 2） success or failure in cognitive tasks could affect different personalities differently; 3） ontogenetic changes of personality profiles could be reflected in changes in cognitive performance. The study of such interplay has implications in animal welfare as well as in neuroscience and in translational medicine [Current Zoology 57 （4）： 491-498, 2011].

Behavioral plasticity, behavioral syndromes and animal personality in crustacean decapods： An imperfect map is better than no map

Despite their key role as model organisms in many behavioral studies, crustacean decapods have been only slightly touched upon by the recent surge of scientific interest in animal personality. Only seven articles investigated the issue in a handful of species among hermit crabs, crabs, and crayfish. Obviously, a limited number of publications does not mean that personality is rare in decapods. On the contrary, few studies might be the result of a form of reluctance by behavioral ecologists to deal with such a phenomenon in these and other invertebrates. This reluctance contrasts with the enthusiasm shown in tackling the beha- vioral plasticity issue. Here we discuss the possible theoretical and methodological difficulties raised by applying the animal per-sonality perspective to decapods and analyze implications of personality studies for their ecology, conservation, and welfare. By highlighting gaps in knowledge and directions of future research, our intention is to increase scientific emphasis on the issue.

Covariation between personalities and individual differences in coping with stress:Converging evidence and hypotheses

In the past decade there has been a profusion of studies highlighting covariation between individual differences in stress physiology and behavioural profiles, here called personalities. Such individual differences in ways of coping with stress are relevant both in biomedicine, since different personalities may experience a different stress and disease vulnerability, and in behavioural ecology, since their adaptive value and evolutionary maintenance are the subject of debate. However, the precise way in which individual stress differences and personalities are linked is unclear. Here we provide an updated overview of this covariation across different species and taxa, consider its functional significance and present working hypotheses for how behavioural and physiological responses to stress might be causally linked, affecting life-history traits such as dispersal and life-span [Current Zoology 56 （6）： 728-740, 2010].

The glutamate metabotropic receptor 5(GRM5)gene is associated with beef cattle home range and movement tortuosity

Background:The grazing behaviour of herbivores and their grazing personalities might in part be determined genetically,but there are few studies in beef cattle illustrating this.In this study,we investigated for first time the genetic variation within a candidate‘grazing gene',the glutamate metabotropic receptor 5 gene(GRM5),and tested associations between variation in that gene and variation in grazing personality behaviours(GP-behaviours)displayed by free-ranging cows during winter grazing in the steep and rugged rangelands of New Zealand.Mature beef cows(n=303,from 3 to 10 years of age)were tracked with global positioning system(GPS)and,with 5-minutes(min)relocation frequency,various GP-behaviours were calculated.These included horizontal and vertical distances travelled,mean elevation,elevation range,elevation gain,slope,home range and movement tortuosity,variously calculated using daily relocation trajectories with repeated measurements(i.e.,7 to 24 days(d))and satellite-derived digital elevation models(DEM).The different GP-behaviours were fitted into mixed models to ascertain their associations with variant sequences and genotypes of GRM5.Results:We discovered three GRM5 variants(A,B and C)and identified the six possible genotypes in the cattle studied.The mixed models revealed that A was significantly associated with elevation range,home range and movement tortuosity.Similarly,GRM5 genotypes were associated(P<0.05)to home range and movement tortuosity,while trends suggesting association(P<0.1)were also revealed for elevation range and horizontal distance travelled.Most GP-behaviour models were improved by correcting for cow age-class as a fixed factor.The analysis of GP-behaviours averaged per cow age-class suggests that grazing personality is fully established as beef cows reached 4 years of age.Home range and movement tortuosity were not only associated with GRM5 variation,but also negatively correlated with each other(r=-0.27,P<0.001).Conclusions:There seems to be a genetically determined trade-off between home range and movement tortuosity that may be useful in beef cattle breeding programmes aiming to improve the grazing distribution and utilisation of steep and rugged rangelands.

Odor cues rather than personality affect tadpole deposition in a neotropical poison frog

Animals constantly need to evaluate available external and internal information to make appropriate decisions.Identifying,assessing,and acting on relevant cues in contexts such as mate choice,intra-sexual competition,and parental care is particularly important for optimizing individual reproductive success.Several factors can infuence decision-making,such as external environmental cues and the animal’s own internal state,yet,we have limited knowledge on how animals integrate available information.Here,we used an entire island population(57 males,53 females,and 1,109 tadpoles)of the neotropical brilliant-thighed poison frog Allobates femoralis to investigate how 2 factors(olfactory cues and personality traits)infuence the ability of males to fnd and use new resources for tadpole deposition.We experimentally manipulated the location of tadpole deposition sites and their associated olfactory cues,and repeatedly measured exploration and boldness in adult males.We further reconstructed tadpole deposition choices via inferred parent–offspring relationships of adult frogs and tadpoles deposited in our experimental pools using molecular parentage analysis.We found that the discovery and use of new rearing sites were heavily infuenced by olfactory cues;however,we did not fnd an effect of the measured behavioral traits on resource discovery and use.We conclude that in highly dynamic environments such as tropical rainforests,reliable external cues likely take priority over personality traits,helping individuals to discover and make use of reproductive resources.

Social experience promotes the exploratory behaviour in Java Sparrows

Personality widely exists in diverse animal taxa.Such inter-individual variance in behaviour is supposed to be influenced by social context.However,it remains unknown whether the experience of social life has any carryover effects on the subsequent expression of personality.Here,we examined exploratory behaviour in caged Java Sparrows(Lonchura oryzivora)using exploration assays.Birds were assigned to live in either a solitary or a social context for four weeks.We compared the expressions of exploration before and after the treatments,and found that birds showed higher exploration tendencies after than before social life,while the isolated birds were consistent in their exploratory behaviours.Different living experience led to differences in the exploration activities for birds without significant differences in exploration before.Our results indicate that social experience can make birds more proactive.

Temperature increase impacts personality traits in aquatic non-native species： Implications for biological invasion under climate change

Climate change, such as elevated temperatures, may facilitate biological invasion. Aquatic animal systems are more vulnerable to biological invasion when the temperature rises. Animal personality also plays an important role in the possibility of successful establishment of invasive species. However, it is still unclear how personality traits of invasive species will respond to global warming. This study focuses on juveniles of red swamp crawfish Procambarus clarkii, an invasive poikilothermic species with wide distribution throughout the world, and explores how slight temperature increases influence three personality traits （ac- tivity, boldness and aggressiveness） ofP. clarkii. For each personality, individual variation is to be presented in condition of the same temperature. Individual personality values can significantly increase with the subtle rising of water temperature. Significant correlations among activity, boldness and aggressiveness are found at any temperature stage, and such relationships are main- tained at a similar level in the face of different temperatures. It is most likely that significant expressions of personality traits may be an ecological compensation strategy to offset increased metabolic costs when faced with small temperature increases. Aggres- sion syndromes are formed due to stable linkages between personality traits, in order to acquire allopatric resources efficiently and establish a new population in this species

Nonlinear maternal effects on personality in a rodent species with fluctuating densities

Consistent among individual variation in behavior,or animal personality,is present in a wide variety of species.This behavioral variation is maintained by both genetic and environmental factors.Parental effects are a special case of environmental variation and are expected to evolve in populations experiencing large fluctuations in their environment.They represent a non-genetic pathway by which parents can transmit information to their offspring,by modulating their personality.While it is expected that parental effects contribute to the observed personality variation,this has rarely been studied in wild populations.We used the multimammate mouse Mastomys natalensis as a model system to investigate the potential effects of maternal personality on offspring behavior.We did this by repeatedly recording the behavior of individually housed juveniles which were born and raised in the lab from wild caught females.A linear correlation,between mother and offspring in behavior,would be expected when the personality is only affected by additive genetic variation,while a more complex relationship would suggests the presence of maternal effects.We found that the personality of the mother predicted the behavior of their offspring in a non-linear pattern.Exploration behavior of mother and offspring was positively correlated,but only for slow and average exploring mothers,while this correlation became negative for fast exploring mothers.This may suggests that early maternal effects could affect personality in juvenile M.natalensis,potentially due to density-dependent and negative frequency-dependent mechanisms,and therefore contribute to the maintenance of personality variation.

A critical evaluation of subjective ratings： Unacquainted observers can reliably assess certain personality traits

Methods to measure consistent individual differences in behavior （i.e. animal personality） fall into two categories, subjective ratings and behavioral codings. Ratings are seldom used despite being potentially more efficient than codings. One potential limitation for the use of ratings is that it is assumed that long-term observers or experts in the field are required to score individuals. This can be problematic in many cases, especially for long-term ecological studies where there is high turnover in personnel. We tested whether raters who were unacquainted with subjects could produce reliable and valid personality assessments of yellow-bellied marmots Marmota flaviventris. Two raters, previously unacquainted with individuals and marmot be- havior, scored 130 subjects on fifteen different adjectives in both open-field （OF） and mirror image stimulation （MIS） trials. Eight OF and nine MIS adjectives were reliable as indicated by both a high degree of intra-observer and inter-observer reliability. Additionally, some ratings were externally valid, correlating with behavioral codings. Our data suggest that activity/exploration and sociability can be a reliable and valid measurement of personality traits in studies where raters were unacquainted with subjects. These traits are observable with the personality tests we used; otherwise researchers using unacquainted raters should be cautious in the tests they employ [Current Zoology 60 （2）： 162-169, 2014].

Responses to threat in a freshwater invader： longitudinal data reveal personality, habituation, and robustness to changing water temperatures in the ＂killer shrimp＂ Dikerogammarus villosus （Crustacea： Amphipoda）

Freshwater biodiversity and ecosystem integrity are under threat from biological invasions. The ＂killer shrimp＂ Dikerogammarus villosus is a highly predatory amphipod that has spread readily across Central Europe and recently the UK and its arrival has been associated with the significant loss of resident species. Despite this, studies of its behavioral ecology are sparse, even though its be- havior may contribute to its invasion success. For the first time, we investigated antipredator ＂fleeing＂ behavior in D. villosus and how this changed with water temperature. Three key patterns emerged from our analysis. First, within a particular temperature condition there are moderate but consistent among-individual differences in behavior. These are driven by a combination of mean level among-individual differences and within-individual relative consistency in behavior, and pro- vide the key marker for animal personalities. Second, the fleeing responses were not influenced by temperature and third, regardless of temperature, all individuals appeared to habituate to a repeated nondangerous stimulus, indicating a capacity for individual learning. We suggest that the antipreda- tor behavior of D. villosus contributes to its rapid spread and that consistent among-individual differ- ences in behavior may promote biological invasions across heterogeneous conditions. Robustness to changing water temperatures may also be potentially advantageous, particularly in an era of glo- bal climate change, where average temperatures could be elevated and less predictable.

Animal personalities and their implications for complex signaling

Animal personalities have been a major focus of behavioral ecology over the past decade. Consistent individual dif ferences in behavior have been found across taxa, and have been shown to influence a range of ecological processes. The role of personalities in sexual selection has been considered, and examples exist that show selection for personality traits with both assortative and disassortative mating patterns between personality types. One overlooked aspect of the personality and sexual se lection literature is the potential for personalitysignaling interactions, specifically with complex signaling. Complex signaling is a diverse topic in itself, and in short, consists of multiple signals within one or more modalities that interact to elicit a receiver response. Research into complex signaling has been thorough, although at times studies discover complex signaling systems that fail to fit into one of the existing hypotheses in the literature. Here, we argue that personalities may interact with complex signal ing, which should be considered by researchers of both personality and sexual selection and communication. We describe several ways in which personalitycomplex signaling interactions could affect both the signaler and receiver, and the way in which they may drive personalityspecific signals as well as receiver preferences. Finally, we discuss how considering personality in com plex signaling studies may inform theory as well as improve the ability of researchers to accurately describe its function.

Is the speed of adjusting to environmental change condition dependent?An experiment with house mice(Mus musculus)

Environmental conditions change constantly either by anthropogenic perturbation or naturally across space and time.Often,a change in behavior is the frst response to changing conditions.Behavioral fexibility can potentially improve an organism’s chances to survive and reproduce.Currently,we lack an understanding on the time-scale such behavioral adjustments need,how they actually affect reproduction and survival and whether behavioral adjustments are suffcient in keeping up with changing conditions.We used house mice(Mus musculus)to test whether personality and life-history traits can adjust to an experimentally induced food-switch fexibly in adulthood or by intergenerational plasticity,that is,adjustments only becoming visible in the offspring generation.Mice lived in 6 experimental populations of semi-natural environments either on high or standard quality food for 4 generations.We showed previously that high-quality food induced better conditions and a less risk-prone personality.Here,we tested whether the speed and/or magnitude of adjustment shows condition-dependency and whether adjustments incur ftness effects.Life-history but not personality traits reacted fexibly to a food-switch,primarily by a direct reduction of reproduction and sloweddown growth.Offspring whose parents received a food-switch developed a more active stress-coping personality and gained weight at a slower rate compared with their respective controls.Furthermore,the modulation of most traits was condition-dependent,with animals previously fed with high-quality food showing stronger responses.Our study highlights that life-history and personality traits adjust at different speed toward environmental change,thus,highlighting the importance of the environment and the mode of response for evolutionary models.

Expanding through the Emerald Isle:exploration and spatial orientation of non-native bank voles in Ireland

Whether introduced into a completely novel habitat or slowly expanding their current range,the degree to which animals can effciently explore and navigate new environments can be key to survival,ultimately determining population establishment and colonization success.We tested whether spatial orientation and exploratory behavior are associated with non-native spread in free-living bank voles(Myodes glareolus,N=43)from a population accidentally introduced to Ireland a century ago.We measured spatial orientation and navigation in a radial arm maze,and behaviors associated to exploratory tendencies and risk-taking in repeated open-feld tests,at the expansion edge and in the source population.Bank voles at the expansion edge re-visited unrewarded arms of the maze more,waited longer before leaving it,took longer to start exploring both the radial arm maze and the open feld,and were more risk-averse compared to conspecifcs in the source population.Taken together,results suggest that for this small mammal under heavy predation pressure,a careful and thorough exploration strategy might be favored when expanding into novel environments.

Behavioral syndrome in a native and an invasive hymenoptera species

Recent studies have focused on the role of behavior in biological invasions. Individuals may differ consistently in time for several behavioral traits （personality） which covary （behavioral syndrome） resulting in different behavioral types, some of them favoring invasion. Social hymenopterans have a strong potential to be invaders and their success depends primarily on the foundresses＇ ability to found viable colonies. They are expected to be active, explorative and bold for optimally establishing their nest. In Europe, 2 homet species coexist： the native Vespa crabro and the invasive Vespa velutina. These 2 species may compete for nesting sites and we suggest that the initial success of V. velutina has been favored by its behavior in outperforming V. crabro for the traits involved in nest initiation. Here, we （i） defined the personality of V. crabro and V：. velutina, （ii） tested for the existence of behavioral syndrome in these species, and （iii） compared their performances using an open-field test. Our results show that V. crabro foundresses behave consistently but not V. velutina; this lack of consistency being mainly due to reduced variance among individuals. This result questions the possibility of detecting consistent behavioral differences in species having recently undergone a strong bottleneck. Both species exhibit the same correlations between activity, boldness and exploration and V. velutina clearly outperforms 14. crabro for all traits. Our results suggest that activity, boldness, and exploration are implicated in both hornet nest initiation and invasion process which contributed to explain why social hymenopterans are so successful at colonization.

Natal dispersal in a social landscape： Considering individual behavioral phenotypes and social environment in dispersal ecology

Natal dispersal, the movement of an organism from its birthplace to the site of first reproduction, is fundamental to many ecological and evolutionary processes. Mechanistically, individual dispersal decisions can depend on both individual phe- notype and environmental cues. In particular, many established evolutionary theories of dispersal highlight the importance of the social environment. More recent research in behavioral ecology has focused on the importance of individual behavioral pheno- types. We reviewed the literature on individual behavioral phenotypes and dispersal and suggest that how individual behavioral phenotypes interact with the immediate social environment experienced by individuals in influencing dispersal is still poorly un- derstood, despite growing interest. We found that very few studies had examined the interaction of individual behavioral pheno- types and social factors, and behavioral phenotypes related to social tendencies were less commonly measured than were beha- vioral phenotypes related to exploration or response to risk. Further, and unsurprisingly, studies on social behavioral phenotypes and dispersal behaviors during the transience stage of dispersal were underrepresented compared to the departure or settlement stages. Future studies in this area should aim to＂ a） make explicit links between behavioral traits and their proposed effects on dispersal decisions throughout multiple stages of dispersal, b） integrate more continuous dispersal variables, and c） consider the effects of the spatial distribution and phenotypes of conspecifics （i.e., the social landscape） encountered by individual dispersers

The ecological consequences of temperament in spiders

Ecological and evolutionary studies on spiders have been featured prominently throughout the contemporary behavio- ral syndromes movement. Here we review the behavioral syndromes literature devoted to spiders, and identify some ways in which behavioral syndromes can impact the function of spiders in ecological communities. We further highlight three general themes within the behavioral syndromes literature for which spiders have served as front running model systems： （1） how trait correlations beget performance trade-offs, （2） the influence that behavioral trait variants have on interspecific interactions and （3） mechanisms that aid in maintaining behavioral variation within-and among-populations. Research on behavioral syndromes con-tinues to grow at an impressive rate, and we feel the success of behavioral syndromes studies in spiders bodes well for their con-tinued prominence.

Phenotypic plasticity of labile traits in the wild

Individual-based studies allow quantification ofphenotypic plasticity in behavioural, life-history and other labile traits The study of phenotypic plasticity in the wild can shed new light on the ultimate objectives （1） whether plasticity itself can evolve or is constrained by its genetic architecture, and （2） whether plasticity is associated to other traits, including fitness （selection）. I describe the main statistical approach for how repeated records of individuals and a description of the environment （E） allow quantification of variation in plasticity across individuals （IxE） and genotypes （GxE） in wild populations. Based on a literature review of life-history and behavioural studies on plasticity in the wild, I discuss the present state of the two objectives listed above. Few studies have quantified GxE of labile traits in wild populations, and it is likely that power to detect statistically sig- nificant GxE is lacking. Apart from the issue of whether it is heritable, plasticity tends to correlate with average trait expression （not fully supported by the few genetic estimates available） and may thus be evolutionary constrained in this way. Individ- ual-specific estimates of plasticity tend to be related to other traits of the individual （including fitness）, but these analyses may be anti-conservative because they predominantly concern stats-on-stats. Despite the increased interest in plasticity in wild popula- tions, the putative lack of power to detect GxE in such populations hinders achieving general insights. I discuss possible steps to invigorate the field by moving away from simply testing for presence of GxE to analyses that ＇scale up＇ to population level proce-sses and by the development of new behavioural theory to identify quantitative genetic parameters which can be estimated

Does exploratory behavior or activity in a wild mouse explain susceptibility to virus infection？

Exploration and activity are often described as trade-offs between the fitness benefits of gathering information and resources, and the potential costs of increasing exposure to predators and para- sites. More exploratory individuals are predicted to have higher rates of parasitism, but this relationship has rarely been examined for virus infections in wild populations. Here, we used the multimammate mouse Mastomys natalensis to investigate the relationship between exploration, activity, and infection with Morogoro virus （MORV）. We characterized individual exploratory behav- ior （open field and novel object tests） and activity （trap diversity）, and quantified the relationship between these traits and infection status using linear regression. We found that M. natalensis expresses consistent individual differences, or personality types, in exploratory behavior （repeat- ability of 0.30, 95% CI： 0.21-0.36）. In addition, we found a significant contrasting effect of age on ex- ploration and activity where juveniles display higher exploration levels than adults, but lower field- activity. There was however no statistical evidence for a behavioral syndrome between these 2 traits. Contrary to our expectations, we found no correlation between MORV infection status and exploratory behavior or activity, which suggests that these behaviors may not increase exposure probability to MORV infection. This would further imply that variation in viral infection between individuals is not affected by between-individual variation in exploration and activity.

Boldness and predator evasion in naturally hybridizing swordtails （Teleostei： Xiphophorus）

Correlations among behavioral traits can generate trade-offs and constrain phenotypic evolution. Interspecific hybridization has the potential to alter behavioral trait correlations, but the effect of hybridization on suites of behavioral traits has received relatively little attention. We evaluated how natural hybridization changes the relationship between boldness （time of emergence and proportion of time out of shelter） and response to a simulated predator threat in swordtails （Teleostei： Xiphophorus）. In poeciliid fishes, bold individuals have increased survival in the presence of predators. This non-intuitive observation may arise as a result of bold individuals being more likely to engage in anti-predator behaviors. Contrary to our prediction, bold indi- viduals were less likely to perform a fast-start response to a predator threat. This correlation was consistent among populations and species but was only significant in hybrids. The observed correlation between boldness and anti-predator behavior could im- pact hybrid fitness and the evolvability of hybrid lineages. More generally, our findings suggest that hybridization could influence the integration of behavioral phenotypes, as has been amply documented for morphology. Animal personality and behavioral syndromes could therefore play an important role in the evolutionary fate of natural hybrids [Current Zoology 61 （4）： 596-603, 2015].

Boldness is related to the development of the cephalic crest in the male of the river blenny Salariafluviatilis （Asso, 1801）

Boldness is defined as the tendency of an individual to take risks when exposed to novel objects or situations. The main aim of this work was to dilucidate if boldness was related to the development of the secondary sexual traits （SSTs） in the male of the fiver blenny Salariafluviatilis, a freshwater fish which takes care of eggs. As a second objective SSTs effect on pa rental status adoption was also be explored. Wild young fish were caught in the Segre River （Ebro basin） in November 2010. Fish were kept in five aquaria in groups of eight males and eight females with artificial nests. Boldness （boldness score and hesitancy） of males （initially with little SSTs development） was tested as the time required to exit a refuge when exposed to a new environ ment. Fish were mantained in the same aquaria for several months, and males＇ SSTs development and reproductive behaviour were monitored. Boldness score was negatively associated to cephalic crest development. Hesitancy was negatively related to male length and was also influenced by the interaction between male length and cephalic crest. Parental status acquisition was positively related to the male＇s length, marginally related to anal gland development, and nonrelated to cephalic crest. These re sults suggest an intrasexual role of the crest as an informative trait of boldness （honest signal） which might not necessarily be chosen by the females. It is also discussed that correlational selection could be responsible for linking boldness and growth rate in this species. Sexual selection, in addition to having directed the evolution of secondary sexual traits, might have favored certain personality traits （e.g. boldness） associated with the achievement of a larger body length .