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  • 1.
    Abbey-Lee, Robin N.
    et al.
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Kreshchenko, Anastasia
    Mechanical and Aeronautical Engineering Division L5, Department of Mechanical, Aerospace & Civil Engineering, Dalton Nuclear Institute, FSE Research Institutes,The University of Manchester, UK.
    Fernandez Sala, Xavier
    Linköping University, Department of Physics, Chemistry and Biology. Linköping University, Faculty of Science & Engineering.
    Petkova, Irina
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering. School of Biological Sciences, Centre for Ecology,Evolution and Behaviour, Royal Holloway University of London, Egham UK.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Effects of monoamine manipulations on the personality and gene expression of three-spined sticklebacks2019In: Journal of Experimental Biology, ISSN 0022-0949, E-ISSN 1477-9145, Vol. 222, no 20, article id jeb211888Article in journal (Refereed)
    Abstract [en]

    Among-individual behavioral differences (i.e. animal personality) are commonly observed across taxa, although the underlying, causal mechanisms of such differences are poorly understood. Animal personality has been correlated with physiological functions as well as fitness-related traits. Variation in many aspects of monoamine systems, such as metabolite levels and gene polymorphisms, has been linked to behavioral variation. Therefore, here we experimentally investigated the potential role of monoamines in explaining individual variation in personality, using two common pharmaceuticals that respectively alter the levels of serotonin and dopamine in the brain: fluoxetine and ropinirole. We exposed three-spined sticklebacks, a species that shows animal personality, to either chemical alone or to a combination of the two chemicals, for 18 days. During the experiment, fish were assayed at four time points for the following personality traits: exploration, boldness, aggression and sociability. To quantify brain gene expression on short- and longer-term scales, fish were sampled at two time points. Our results show that monoamine manipulations influence fish behavior. Specifically, fish exposed to either fluoxetine or ropinirole were significantly bolder, and fish exposed to the two chemicals together tended to be bolder than control fish. Our monoamine manipulations did not alter the gene expression of monoamine or stress-associated neurotransmitter genes, but control, untreated fish showed covariation between gene expression and behavior. Specifically, exploration and boldness were predicted by genes in the dopaminergic, serotonergic and stress pathways, and sociability was predicted by genes in the dopaminergic and stress pathways. These results add further support to the links between monoaminergic systems and personality, and show that exposure to monoamines can causally alter animal personality.

  • 2.
    Abbey-Lee, Robin N.
    et al.
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Kreshchenko, Anastasia
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Fernandez Sala, Xavier
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Petkova, Irina
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Effects of monoamine manipulations on the personality and gene expression of three-spined sticklebacks2019Data set
    Abstract [en]

    Among-individual behavioral differences (i.e. animal personality) are commonly observed across taxa, although the underlying, causal mechanisms of such differences are poorly understood. Animal personality has been implicated in correlations with physiological functions as well as affecting fitness-related traits. Variation in many aspects of monoamine systems, such as metabolite levels and gene polymorphisms, has been linked to behavioral variation. Therefore, here we investigated the potential role of monoamines in explaining individual variation in personality, using two common pharmaceuticals that respectively alter the levels of serotonin and dopamine in the brain: fluoxetine and ropinirole. We exposed three- spined sticklebacks, a species that shows animal personality, to either chemical alone or to a combination of the two chemicals, for 18 days. During the experiment, fish were assayed at four time points for the following personality traits: exploration, boldness, aggression and sociability. To quantify brain gene expression on short- and longer-term scales, fish were sampled at two time points. Our results show that monoamine manipulations influence fish behavior. Specifically, fish exposed to either fluoxetine or ropinirole were significantly bolder, and fish exposed to the two chemicals together tended to be bolder than control fish. Our monoamine manipulations did not alter the gene expression of monoamine or stress-associated neurotransmitter genes, but control, untreated fish showed covariation between gene expression and behavior. Specifically, exploration and boldness were predicted by genes in the dopaminergic, serotonergic and stress pathways, and sociability was predicted by genes in the dopaminergic and stress pathways. These results add further support to the links between monoaminergic systems and personality, and show that exposure to monoamines can causally alter animal personality.

  • 3.
    Abbey-Lee, Robin N.
    et al.
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Uhrig, Emily J.
    Department of Zoology, Stockholm University, 10691 Stockholm, Sweden.
    Zidar, Josefina
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Favati, Anna
    Department of Zoology, Stockholm University, 10691 Stockholm, Sweden.
    Almberg, Johan
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Dahlblom, Josefin
    Department of Neuroscience, Uppsala Biomedical Centre BMC, Uppsala University, Uppsala, Sweden.
    Winberg, Svante
    Department of Neuroscience, Uppsala Biomedical Centre BMC, Uppsala University, Uppsala, Sweden.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    The influence of rearing on behavior, brain monoamines and gene expression in three-spined sticklebacks2018Data set
    Abstract [en]
    1. The causes of individual variation in behavior are often not well understood, and potential underlying mechanisms include both intrinsic and extrinsic factors, such as early environmental, physiological, and genetic differences.
    2. In an exploratory laboratory study, we raised three-spined sticklebacks (Gasterosteus aculeatus) under 4 different environmental conditions (simulated predator environment, complex environment, variable social environment, and control). We investigated how these manipulations related to behavior, brain physiology and gene expression later in life, with focus on brain dopamine and serotonin levels, turnover rates, and gene expression.
    3. The different rearing environments influenced behavior and gene expression, but did not alter monoamine levels or metabolites. Specifically, compared to control fish, fish exposed to a simulated predator environment tended to be less aggressive, more exploratory, and more neophobic; and fish raised in both complex and variable social environments tended to be less neophobic. Exposure to a simulated predator environment tended to lower expression of dopamine receptor DRD4A, a complex environment increased expression of dopamine receptor DRD1B, while a variable social environment tended to increase serotonin receptor 5-HTR2B and increased serotonin transporter SLC6A4A expression. Despite both behavior and gene expression varying with early environment, there was no evidence that gene expression mediated the relationship between early environment and behavior.
    4. Our results confirm that environmental conditions early in life can affect phenotypic variation. However, the mechanistic pathway of the monoaminergic systems translating early environmental variation into observed behavioral responses was not detected.
  • 4.
    Abbey-Lee, Robin N.
    et al.
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Uhrig, Emily
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Zidar, Josefina
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Favati, A.
    Department of Zoology, Stockholm University, Stockholm, Sweden.
    Almberg, J.
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Dahlbom, J.
    Department of Neuroscience, Uppsala Biomedical Centre BMC, Uppsala University, Uppsala, Sweden.
    Winberg, S.
    Department of Neuroscience, Uppsala Biomedical Centre BMC, Uppsala University, Uppsala, Sweden.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    The Influence of Rearing on Behavior, Brain Monoamines, and Gene Expression in Three-Spined Sticklebacks2018In: Brain, behavior, and evolution, ISSN 0006-8977, E-ISSN 1421-9743, Vol. 91, no 4, p. 201-213Article in journal (Refereed)
    Abstract [en]

    The causes of individual variation in behavior are often not well understood, and potential underlying mechanisms include both intrinsic and extrinsic factors, such as early environmental, physiological, and genetic differences. In an exploratory laboratory study, we raised three-spined sticklebacks <i>(Gasterosteus aculeatus)</i> under 4 different environmental conditions (simulated predator environment, complex environment, variable social environment, and control). We investigated how these manipulations related to behavior, brain physiology, and gene expression later in life, with focus on brain dopamine and serotonin levels, turnover rates, and gene expression. The different rearing environments influenced behavior and gene expression, but did not alter monoamine levels or metabolites. Specifically, compared to control fish, fish exposed to a simulated predator environment tended to be less aggressive, more exploratory, and more neophobic; and fish raised in both complex and variable social environments tended to be less neophobic. Exposure to a simulated predator environment tended to lower expression of dopamine receptor DRD4A, a complex environment increased expression of dopamine receptor DRD1B, while a variable social environment tended to increase serotonin receptor 5-HTR2B and serotonin transporter SLC6A4A expression. Despite both behavior and gene expression varying with early environment, there was no evidence that gene expression mediated the relationship between early environment and behavior. Our results confirm that environmental conditions early in life can affect phenotypic variation. However, the mechanistic pathway of the monoaminergic systems translating early environmental variation into observed behavioral responses was not detected.

  • 5.
    Abbey-Lee, Robin
    et al.
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Uhrig, Emily
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering. Southern Oregon Univ, OR 97520 USA.
    Garnham, Laura
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Lundgren, Kristoffer
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Child, Sarah
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering. Univ Manchester, England.
    Lovlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Experimental manipulation of monoamine levels alters personality in crickets2018In: Scientific Reports, ISSN 2045-2322, E-ISSN 2045-2322, Vol. 8, article id 16211Article in journal (Refereed)
    Abstract [en]

    Animal personality has been described in a range of species with ecological and evolutionary consequences. Factors shaping and maintaining variation in personality are not fully understood, but monoaminergic systems are consistently linked to personality variation. We experimentally explored how personality was influenced by alterations in two key monoamine systems: dopamine and serotonin. This was done using ropinirole and fluoxetine, two common human pharmaceuticals. Using the Mediterranean field cricket (Gryllus bimaculatus), we focused on the personality traits activity, exploration, and aggression, with confirmed repeatability in our study. Dopamine manipulations explained little variation in the personality traits investigated, while serotonin manipulation reduced both activity and aggression. Due to limited previous research, we created a dose-response curve for ropinirole, ranging from concentrations measured in surface waters to human therapeutic doses. No ropinirole dose level strongly influenced cricket personality, suggesting our results did not come from a dose mismatch. Our results indicate that the serotonergic system explains more variation in personality than manipulations of the dopaminergic system. Additionally, they suggest that monoamine systems differ across taxa, and confirm the importance of the mode of action of pharmaceuticals in determining their effects on behaviour.

  • 6.
    Cauchoix, M.
    et al.
    CNRS, France; Inst Adv Study Toulouse, France.
    Chow, P. K. Y.
    Univ Exeter, England; Hokkaido Univ, Japan.
    van Horik, J. O.
    Univ Exeter, England.
    Atance, C. M.
    Univ Ottawa, Canada.
    Barbeau, E. J.
    UPS, France.
    Barragan-Jason, G.
    Inst Adv Study Toulouse, France.
    Bize, P.
    Univ Aberdeen, Scotland.
    Boussard, A.
    Stockholm Univ, Sweden.
    Buechel, S. D.
    Stockholm Univ, Sweden.
    Cabirol, A.
    Univ Paul Sabatier, France.
    Cauchard, L.
    Univ Montreal, Canada.
    Claidiere, N.
    Aix Marseille Univ, France.
    Dalesman, S.
    Aberystwyth Univ, Wales.
    Devaud, J. M.
    Univ Paul Sabatier, France.
    Didic, M.
    AP HM Timone, France; Inst Neurosci Syst, France.
    Doligez, B.
    Univ Lyon 1, France.
    Fagot, J.
    Aix Marseille Univ, France.
    Fichtel, C.
    German Primate Ctr, Germany; Univ Gottingen, Germany; Leibniz Sci Campus Primate Cognit, Germany.
    Henke-von der Malsburg, J.
    German Primate Ctr, Germany; Univ Gottingen, Germany; Leibniz Sci Campus Primate Cognit, Germany.
    Hermer, E.
    Univ Gottingen, Germany.
    Huber, L.
    Leibniz Sci Campus Primate Cognit, Germany.
    Huebner, F.
    German Primate Ctr, Germany; Univ Gottingen, Germany; Leibniz Sci Campus Primate Cognit, Germany.
    Kappeler, P. M.
    German Primate Ctr, Germany; Univ Gottingen, Germany; Leibniz Sci Campus Primate Cognit, Germany.
    Klein, S.
    Univ Paul Sabatier, France.
    Langbein, J.
    Leibniz Inst Farm Anim Biol, Germany.
    Langley, E. J. G.
    Univ Exeter, England.
    Lea, S. E. G.
    Univ Exeter, England.
    Lihoreau, M.
    Univ Paul Sabatier, France.
    Lovlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Matzel, L. D.
    Rutgers State Univ, NJ USA.
    Nakagawa, S.
    Univ New South Wales, Australia; Univ New South Wales, Australia.
    Nawroth, C.
    Leibniz Inst Farm Anim Biol, Germany.
    Oesterwind, S.
    Univ Rostock, Germany.
    Sauce, B.
    Rutgers State Univ, NJ USA.
    Smith, E. A.
    Univ Lincoln, England.
    Sorato, Enrico
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Tebbich, S.
    Univ Vienna, Austria.
    Wallis, L. J.
    Univ Vienna, Austria; Eotvos Lorand Univ, Hungary.
    Whiteside, M. A.
    Univ Exeter, England.
    Wilkinson, A.
    Univ Lincoln, England.
    Chaine, A. S.
    CNRS, France; Inst Adv Study Toulouse, France.
    Morand-Ferron, J.
    Univ Ottawa, Canada.
    The repeatability of cognitive performance: a meta-analysis2018In: Philosophical Transactions of the Royal Society of London. Biological Sciences, ISSN 0962-8436, E-ISSN 1471-2970, Vol. 373, no 1756, article id 20170281Article in journal (Refereed)
    Abstract [en]

    Behavioural and cognitive processes play important roles in mediating an individuals interactions with its environment. Yet, while there is a vast literature on repeatable individual differences in behaviour, relatively little is known about the repeatability of cognitive performance. To further our understanding of the evolution of cognition, we gathered 44 studies on individual performance of 25 species across six animal classes and used meta-analysis to assess whether cognitive performance is repeatable. We compared repeatability (R) in performance (1) on the same task presented at different times (temporal repeatability), and (2) on different tasks that measured the same putative cognitive ability (contextual repeatability). We also addressed whether R estimates were influenced by seven extrinsic factors (moderators): type of cognitive performance measurement, type of cognitive task, delay between tests, origin of the subjects, experimental context, taxonomic class and publication status. We found support for both temporal and contextual repeatability of cognitive performance, with mean R estimates ranging between 0.15 and 0.28. Repeatability estimates were mostly influenced by the type of cognitive performance measures and publication status. Our findings highlight the widespread occurrence of consistent inter-individual variation in cognition across a range of taxa which, like behaviour, may be associated with fitness outcomes. This article is part of the theme issue Causes and consequences of individual differences in cognitive abilities.

  • 7.
    Dean, Rebecca
    et al.
    University of Oxford, UK.
    Cornwallis, Charlie K
    University of Oxford, UK.
    Løvlie, Hanne
    University of Oxford, UK; Stockholm University, Sweden.
    Worley, Kirsty
    University of Oxford, UK; University of East Anglia, UK.
    Richardson, David S.
    University of East Anglia, UK.
    Pizzari, Tommaso
    University of Oxford, UK.
    Male reproductive senescence causes potential for sexual conflict over mating2010In: Current Biology, ISSN 0960-9822, E-ISSN 1879-0445, Vol. 20, no 13, p. 1192-1196Article in journal (Refereed)
    Abstract [en]

    The realization that senescence, age-dependent declines in survival and reproductive performance, pervades natural populations has brought its evolutionary significance into sharper focus. However, reproductive senescence remains poorly understood because it is difficult to separate male and female mechanisms underpinning reproductive success. We experimentally investigated male reproductive senescence in feral fowl, Gallus gallus domesticus, where socially dominant males monopolize access to females and the ejaculates of multiple males compete for fertilization. We detected the signal of senescence on multiple determinants of male reproductive success. The effect of age on status was dependent upon the intensity of intrasexual competition: old males were less likely to dominate male-biased groups where competition is intense but were as likely as young males to dominate female-biased groups. Mating and fertilization success declined sharply with male age largely as a result of population-level patterns. These age-dependent declines translated into sexually antagonistic payoffs: old males fertilized more eggs when they were dominant, but this resulted in females suffering a drastic reduction in fertility. Thus, male senescence causes potential for sexual conflict over mating, and the intensity of this conflict is modulated socially, by the probability of old males dominating reproductive opportunities.

  • 8.
    Favati, Anna
    et al.
    Department of Zoology, Stockholm University, Stockholm, Sweden.
    Leimar, Olof
    Department of Zoology, Stockholm University, Stockholm, Sweden.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, The Institute of Technology.
    Personality Predicts Social Dominance in Male Domestic Fowl2014In: PLoS ONE, ISSN 1932-6203, E-ISSN 1932-6203, Vol. 9, no 7, p. e103535-Article in journal (Refereed)
    Abstract [en]

    Individuals in social species commonly form dominance relationships, where dominant individuals enjoy greater access to resources compared to subordinates. A range of factors such as sex, age, body size and prior experiences has to varying degrees been observed to affect the social status an individual obtains. Recent work on animal personality (i.e. consistent variation in behavioural responses of individuals) demonstrates that personality can co-vary with social status, suggesting that also behavioural variation can play an important role in establishment of status. We investigated whether personality could predict the outcome of duels between pairs of morphologically matched male domestic fowl (Gallus gallus domesticus), a species where individuals readily form social hierarchies. We found that males that more quickly explored a novel arena, or remained vigilant for a longer period following the playback of a warning call were more likely to obtain a dominant position. These traits were uncorrelated to each other and were also uncorrelated to aggression during the initial part of the dominance-determining duel. Our results indicate that several behavioural traits independently play a role in the establishment of social status, which in turn can have implications for the reproductive success of different personality types.

  • 9.
    Favati, Anna
    et al.
    Stockholm University, Sweden.
    Leimar, Olof
    Stockholm University, Sweden.
    Radesäter, Tommy
    Stockholm University, Sweden.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, The Institute of Technology. Stockholm University, Sweden.
    Social status and personality: stability in social state can promote consistency of behavioural responses2014In: Proceedings of the Royal Society of London. Biological Sciences, ISSN 0962-8452, E-ISSN 1471-2954, Vol. 281, no 1774, p. 20132531-Article in journal (Refereed)
    Abstract [en]

    Stability of ‘state’ has been suggested as an underlying factor explainingbehavioural stability and animal personality (i.e. variation among, andconsistency within individuals in behavioural responses), but the possibilitythat stable social relationships represent such states remains unexplored.Here, we investigated the influence of social status on the expression andconsistency of behaviours by experimentally changing social status betweenrepeated personality assays. We used male domestic fowl (Gallus gallusdomesticus), a social species that forms relatively stable dominance hierarchies,and showed that behavioural responses were strongly affected bysocial status, but also by individual characteristics. The level of vigilance,activity and exploration changed with social status, whereas boldnessappeared as a stable individual property, independent of status. Furthermore,variation in vocalization predicted future social status, indicatingthat individual behaviours can both be a predictor and a consequence ofsocial status, depending on the aspect in focus. Our results illustrate thatsocial states contribute to both variation and stability in behaviouralresponses, and should therefore be taken into account when investigatingand interpreting variation in personality.

  • 10.
    Favati, Anna
    et al.
    Department of Zoology, Stockholm University, Stockholm 106 91, Sweden.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Leimar, Olof
    Department of Zoology, Stockholm University, Stockholm 106 91, Sweden.
    Individual aggression, but not winner–losereffects, predicts social rank in male domestic fowl2017In: Behavioral Ecology, ISSN 1045-2249, E-ISSN 1465-7279, Vol. 28, no 3, p. 874-882Article in journal (Refereed)
    Abstract [en]

    Many factors can affect the probability for an individual to obtain a high social rank, including size, weaponry, and behavioral attributes

    such as aggression. Recent experiences of winning or losing can also affect the chances of winning future contests, commonly

    referred to as “winner–loser effects”. Individuals often differ in behavior in a consistent way, including in aggression, thereby showing

    differences in personality. However, the relative importance of recent experience and aspects of personality in determining rank,

    as well as the extent to which winning or losing affects aggression, has rarely been studied. Here, we investigate these questions

    using male domestic fowl. We matched males for body size, comb size, and aggression in pair-wise duels to: 1) study the effect of

    contest outcome on aggression and 2) compare the effect of individual aggression and contest experience on future social status in

    small groups. We found that aggression was a highly repeatable personality trait and that aggression increased after winning and

    decreased after losing. Nevertheless, such winner–loser effects were not enough to increase the odds of becoming dominant in a

    small group. Instead, aggressiveness measured prior to a contest experience best predicted future rank. Boldness and exploration did

    not predict rank and of the 2, only boldness was positively correlated with aggressiveness. We conclude that for male domestic fowl

    in contests among phenotypically matched contestants, aggressiveness is more important for obtaining high rank than winner–loser

    effects, or other aspects of personality.

  • 11.
    Favati, Anna
    et al.
    Stockholm Univ, Sweden.
    Uden, Eva
    Stockholm Univ, Sweden.
    Leimar, Olof
    Stockholm Univ, Sweden.
    Lovlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Personality remains: no effect of 3-week social status experience on personality in male fowl2018In: Behavioral Ecology, ISSN 1045-2249, E-ISSN 1465-7279, Vol. 29, no 2, p. 312-320Article in journal (Refereed)
    Abstract [en]

    Behavioral responses of male fowl did not depend on social rank after 3 weeks in stable groups, but were consistent over time for an individual. Theory suggests that stable social states, for example, stable social hierarchies, may lead to consistent variation in behavior, that is, variation in personality. Our results suggest that variation in personality is not a consequence of variation in social status and that personality is more important than current social position in determining individual behavior in stable groups.Individuals often differ in behavior in a consistent way, that is, they show variation in personality. Understanding the processes explaining the emergence and maintenance of this variation is a current major topic in the field of animal behavioral research. Recent theoretical models predict that differences in various "states" can generate individual variation in behavior. Previous studies have mainly focused on endogenous states like metabolic rate or energy reserves, but theory also suggests that states based on social interactions could play important roles in shaping personality. We have earlier demonstrated short-term status-dependent variation in behavior in the domestic fowl (Gallus gallus domesticus), but whether such behavioral variation remains also after a longer period of time, is unknown. Therefore, we examine the influence of social status on variation in behavior, using experimental manipulation of social status in pairs of male domestic fowl. We scored males in 3 personality assays (novel arena test, novel object test, and aggression test) before and after 3 weeks in pairs as either dominant or subordinate. We observed individual consistency of behavior despite alteration of social status. We further found no support for social status acting as a state that generates variation in personality over the used time interval: social status had no significant effect on the change in behavioral responses between repeated personality tests. Our results suggest that personality is more important than current social situation for describing individual behavior in stable groups.

  • 12.
    Favati, Anna
    et al.
    Department of Zoology, Stockholm University, Stockholm, Sweden.
    Zidar, Josefina
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Thorpe, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Jensen, Per
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Lovlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    The ontogeny of personality traits in the redjunglefowl, Gallus gallus2016In: Behavioral Ecology, ISSN 1045-2249, E-ISSN 1465-7279, Vol. 27, no 2, p. 484-493Article in journal (Refereed)
    Abstract [en]

    Consistent behavioral differences among individuals, that is, personality, are described in numerous species. Nevertheless, thedevelopment of behavioral consistency over ontogeny remains unclear, including whether the personality of individuals is consistentthroughout life, and if adult personality can be predicted already at young age. We investigated the ontogeny of personality in thered junglefowl (Gallus gallus) by scoring personality of hatchlings at 5 time points through adulthood, including before and after themajor developmental stages of becoming independent and sexual mature. We use the conceptual framework laid out by Stamps andGroothuis (2010a) to holistically investigate the observed changes in behavioral response over ontogeny. We demonstrate that meanvalues of behavioral responses changed across ontogeny and stabilized after independence. Rank-order consistencies of behavioralresponses were overall low across independence and sexual maturation. Only in 1 case could low rank-order consistencies potentiallybe explained by different phenotypes displaying different amounts of change in behavior; more explorative individuals decreased inexploration after independence, while less explorative individuals remained so. Correlations among behavior varied across ontogenyand weakened after sexual maturation. Our results demonstrate that both absolute values and consistency of behavioral traits maychange across ontogeny and that individual developmental trajectories and adult personality only to some extent can be predictedearly in life. These results have implications for future studies on personality, highlighting that the life stage at which individuals arescored affects the observed consistency of behavioral responses.

  • 13.
    Garnham, Laura
    et al.
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Sophisticated fowl: The complex behaviour andcognitive skills of chickens and red junglefowl2018In: Behavioral Sciences, ISSN 2076-328X, Vol. 8, no 13Article in journal (Refereed)
    Abstract [en]

    The world’s most numerous bird, the domestic chicken, and their wild ancestor, the red junglefowl, have long been used as model species for animal behaviour research. Recently, this research has advanced our understanding of the social behaviour, personality, and cognition of fowl, and demonstrated their sophisticated behaviour and cognitive skills. Here, we overview some of this research, starting with describing research investigating the well-developed senses of fowl, before presenting how socially and cognitively complex they can be. The realisation that domestic chickens, our most abundant production animal, are behaviourally and cognitively sophisticated should encourage an increase in general appraise and fascination towards them. In turn, this should inspire increased use of them as both research and hobby animals, as well as improvements in their unfortunately often poor welfare.

  • 14.
    Gillingham, Mark
    et al.
    University of Oxford, UK.
    Richardson, David
    University of East Anglia, UK.
    Løvlie, Hanne
    University of Oxford, UK.
    Moynihan, Anna
    University of Oxford, UK; University of Edinburgh, UK.
    Worley, Kirsty
    University of Oxford, UK; University of East Anglia, UK.
    Pizzari, Tom
    University of Oxford, UK.
    Cryptic preference for MHC-dissimilar females in male red junglefowl, Gallus gallus2009In: Proceedings of the Royal Society of London. Biological Sciences, ISSN 0962-8452, E-ISSN 1471-2954, Vol. 276, no 1659, p. 1083-1092Article in journal (Refereed)
    Abstract [en]

    An increasing number of studies test the idea that females increase offspring fitness by biasing fertilization in favour of genetically compatible partners; however, few have investigated or controlled for corresponding preferences in males. Here, we experimentally test whether male red junglefowl, Gallus gallus, prefer genetically compatible females, measured by similarity at the major histocompatibility complex (MHC), a key gene complex in vertebrate immune function. Theory predicts that because some degree of MHC heterozygosity favours viability, individuals should prefer partners that carry MHC alleles different from their own. While male fowl showed no preference when simultaneously presented with an MHC-similar and an MHC-dissimilar female, they showed a 'cryptic' preference, by allocating more sperm to the most MHC-dissimilar of two sequentially presented females. These results provide the first experimental evidence that males might respond to the MHC similarity of a female through differential ejaculate expenditure. By revealing that cryptic male behaviours may bias fertilization success in favour of genetically compatible partners, this study demonstrates the need to experimentally disentangle male and female effects when studying preferences for genetically compatible partners.

  • 15.
    Girndt, Antje
    et al.
    Max Planck Institute Ornithol, Germany; Imperial Coll London, England; University of Konstanz, Germany.
    Cockburn, Glenn
    Max Planck Institute Ornithol, Germany; University of Konstanz, Germany.
    Sanchez-Tojar, Alfredo
    Max Planck Institute Ornithol, Germany; Imperial Coll London, England; University of Konstanz, Germany.
    Lovlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Schroeder, Julia
    Max Planck Institute Ornithol, Germany; Imperial Coll London, England.
    Method matters: Experimental evidence for shorter avian sperm in faecal compared to abdominal massage samples2017In: PLoS ONE, ISSN 1932-6203, E-ISSN 1932-6203, Vol. 12, no 8, article id e0182853Article in journal (Refereed)
    Abstract [en]

    Birds are model organisms in sperm biology. Previous work in zebra finches, suggested that sperm sampled from males faeces and ejaculates do not differ in size. Here, we tested this assumption in a captive population of house sparrows, Passer domesticus. We compared sperm length in samples from three collection techniques: female dummy, faecal and abdominal massage samples. We found that sperm were significantly shorter in faecal than abdominal massage samples, which was explained by shorter heads and midpieces, but not flagella. This result might indicate that faecal sampled sperm could be less mature than sperm collected by abdominal massage. The female dummy method resulted in an insufficient number of experimental ejaculates because most males ignored it. In light of these results, we recommend using abdominal massage as a preferred method for avian sperm sampling. Where avian sperm cannot be collected by abdominal massage alone, we advise controlling for sperm sampling protocol statistically.

  • 16.
    Hayward, A
    et al.
    Centre for Ecology and Conservation, University of Exeter, Penryn, UK.
    Tsuboi, M
    Department of Animal Ecology, Uppsala University, Uppsala, Sweden.
    Owusu, C
    Department of Animal Ecology, Uppsala University, Uppsala, Sweden.
    Kotrschal, A
    Department of Zoology, Stockholm University, Stockholm, Sweden.
    Buechel, S D
    Department of Zoology, Stockholm University, Stockholm, Sweden.
    Zidar, Josefina
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Cornwallis, C K
    Department of Ecology, Lund University, Lund, Sweden.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Kolm, N
    Department of Zoology, Stockholm University, Stockholm, Sweden.
    Evolutionary associations between host traits and parasite load: insights from Lake Tanganyika cichlids.2017In: Journal of Evolutionary Biology, ISSN 1010-061X, E-ISSN 1420-9101, Vol. 30, no 6, p. 1056-1067Article in journal (Refereed)
    Abstract [en]

    Parasite diversity and abundance (parasite load) vary greatly among host species. However, the influence of host traits on variation in parasitism remains poorly understood. Comparative studies of parasite load have largely examined measures of parasite species richness and are predominantly based on records obtained from published data. Consequently, little is known about the relationships between host traits and other aspects of parasite load, such as parasite abundance, prevalence and aggregation. Meanwhile, understanding of parasite species richness may be clouded by limitations associated with data collation from multiple independent sources. We conducted a field study of Lake Tanganyika cichlid fishes and their helminth parasites. Using a Bayesian phylogenetic comparative framework, we tested evolutionary associations between five key host traits (body size, gut length, diet breadth, habitat complexity and number of sympatric hosts) predicted to influence parasitism, together with multiple measures of parasite load. We find that the number of host species that a particular host may encounter due to its habitat preferences emerges as a factor of general importance for parasite diversity, abundance and prevalence, but not parasite aggregation. In contrast, body size and gut size are positively related to aspects of parasite load within, but not between species. The influence of host phylogeny varies considerably among measures of parasite load, with the greatest influence exerted on parasite diversity. These results reveal that both host morphology and biotic interactions are key determinants of host-parasite associations and that consideration of multiple aspects of parasite load is required to fully understand patterns in parasitism.

  • 17.
    Hedlund, Louise
    et al.
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Lovlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Personality and production: Nervous cows produce less milk2015In: Journal of Dairy Science, ISSN 0022-0302, E-ISSN 1525-3198, Vol. 98, no 9, p. 5819-5828Article in journal (Refereed)
    Abstract [en]

    The objective of this study was to investigate relationships between animal personality (i.e., consistency in behavioral responses, also called temperament) and milk production in dairy cows. There has recently been a growing research interest in animal personality, which in production animals can have an important impact on welfare and production potential. Despite this, the relationship between personality and milk production in dairy cows remains unclear. Here we investigate links between behavioral responses during milking and in personality tests (responses to novel object and social isolation) with milk production in 2 breeds of dairy cattle, Swedish Red and White and Holstein. The milk production parameters investigated were energy-corrected milk (in kg) for the cows first lactation and energy-corrected milk for their current lactation. Overall, cows that stepped more during milking or spent more time facing the herd during social isolation produced less milk in their first lactation. Cows that vocalized more during isolation had a lower current milk production. Variation in other behavioral responses showed limited relationships with milk production. Taken together, our results support a relationship between behavioral responses and milk production, where cows showing signs of nervousness produce less milk. However, observed relationships are dependent on the milk measure used, behavior, and breed investigated, supporting that the relationship between behavior and production traits is not straightforward.

  • 18.
    Lisney, Thomas J.
    et al.
    Evolutionary Biology Centre, Uppsala University, Sweden.
    Rubene, Diana
    Evolutionary Biology Centre, Uppsala University, Sweden.
    Roza, Jani
    Evolutionary Biology Centre, Uppsala University, Sweden.
    Løvlie, Hanne
    Stockholm University, Sweden.
    Håstad, Olle
    Uppsala University, Sweden; Swedish University of Agricultural Sciences, Uppsala, Sweden.
    Ödeen, Anders
    volutionary Biology Centre, Uppsala University, Sweden.
    Behavioural assessment of flicker fusion frequency in chicken, Gallus gallus domesticus2011In: Vision Research, ISSN 0042-6989, E-ISSN 1878-5646, Vol. 51, no 12, p. 1324-1332Article in journal (Refereed)
    Abstract [en]

    To interact with its visual environment, an organism needs to perceive objects in both space and time. High temporal resolution is hence important to the fitness of diurnally active animals, not least highly active aerial species such as birds. However, temporal resolution, as assessed by flicker fusion frequency (FFF; the stimulus frequency at which a flickering light stimulus can no longer be resolved and appears continuous) or critical flicker fusion frequency (CFF; the highest flicker fusion frequency at any light intensity) has rarely been assessed in birds. In order to further our understanding of temporal resolution as a function of light intensity in birds we used behavioural experiments with domestic chickens (Gallus gallus domesticus) from an old game breed 'Gammalsvensk dvärghöna' (which is morphologically and behaviourally similar to the wildtype ancestor, the red jungle fowl, G. gallus), to generate an 'Intensity/FFF curve' (I/FFF curve) across full spectrum light intensities ranging from 0.2 to 2812 cd m⁻². The I/FFF curve is double-branched, resembling that of other chordates with a duplex retina of both rods and cones. Assuming that the branches represent rod and cone mediated responses respectively, the break point between them places the transition between scotopic and photopic vision at between 0.8 and 1.9 cd m⁻². Average FFF ranged from 19.8 Hz at the lowest light intensity to a CFF 87.0 Hz at 1375 cd m⁻². FFF dropped slightly at the highest light intensity. There was some individual variation with certain birds displaying CFFs of 90-100 Hz. The FFF values demonstrated by this non-selected breed appear to be considerably higher than other behaviourally derived FFF values for similar stimuli reported for white and brown commercial laying hens, indicating that the domestication process might have influenced temporal resolution in chicken.

  • 19.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Cryptic Female Choice2016In: Oxfords Bibliographies in Evolutionary BiologyArticle, review/survey (Refereed)
    Abstract [en]

    Sexual selection (see the Oxford Bibliographies article “Sexual Selection”) is a powerful evolutionary force, selecting fortraits that increase the reproductive success of individuals. Before copulation, sexual selection can occur throughintrasexual selection, typically observed as competition among individuals of the same sex for access to mating partnersof the other sex (see Oxford Bibliographies article on Evolutionary Biology “Male-Male Competition”), and intersexualselection, observed as (typically female) mate choice (see Oxford Bibliographies article on Evolutionary Biology “MateChoice”). When females are sexually promiscuous and mate with multiple males (which is more the rule than theexception in the animal kingdom), these two processes have the potential to continue also after copulation: intrasexualselection as sperm competition (Oxford Bibliographies article on Evolutionary Biology “Sperm Competition”), andintersexual selection as cryptic female choice. The term cryptic is applied because this form of female choice can be hardto observe (e.g., when it occurs inside the female reproductive tract) and hard to quantify with classical measures ofreproductive success (e.g., mating success). In addition, this form of female choice is hard to disentangle from otherepisodes of sexual selection (see below). The framework used to understand female choice occurring after (or sometimesduring) copulation is currently somewhat divergent, since some authors adopt a very broad definition of cryptic femalechoice, while others apply a more conservative definition (see discussion of this under Definition and History). Crypticfemale choice is a relatively young research topic (it first started properly after the publication of Eberhard’s seminal bookFemale Control: Sexual Selection by Cryptic Female Choice [Eberhard 1996, cited under General Overviews] in 1996). Itwas realized early on in the history of the field that a broad range of mechanisms across a variety of species exist throughwhich females can potentially bias the outcome of a copulation (e.g., ejaculate ejection, differential sperm storage, spermchoice—see section Mechanisms and Processes Used as Cryptic Female Choice). As a consequence, measures ofprecopulatory processes or sperm competition can be misleading in species with cryptic female choice, due to femalepostcopulatory influences on fertilization. Yet, although there is no doubt that females have great potential to bias paternityat the postcopulatory stage, cryptic female choice is the least studied of the processes through which sexual selection canoccur (e.g., compared to sperm competition, or male-male competition). This is probably because demonstration of crypticfemale choice is notoriously difficult. It can be challenging to separate pre- from postcopulatory processes, the interactionof male adaptations to sperm competition and female influences on fertilization, and variation in differential embryomortality from female-induced biases in paternity (see Potential Pitfalls in the Study of Cryptic Female Choice). Thestudies that have convincingly been able to separate these processes and demonstrate cryptic female choice are currentlyprimarily from insect, bird, and externally fertilizing species (see Mechanisms and Processes Used as Cryptic FemaleChoice). I here present when we may expect to observe cryptic female choice, how females may benefit from crypticfemale choice, some techniques that can

  • 20.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering. Linköping University.
    Cryptic female choice2016In: ENCYCLOPEDIA OF EVOLUTIONARY PSYCHOLOGICAL SCIENCEArticle in journal (Refereed)
  • 21.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Introduction to animal personality2017In: The Ethology of Domestic Animals / [ed] Per Jensen, CABI Publishing, 2017, 3, p. 104-118Chapter in book (Refereed)
  • 22.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Personliga höns2017Other (Other (popular science, discussion, etc.))
  • 23.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Personliga höns2017In: Svenska rasfjäderfäförbundets tidskrift, ISSN 1650-7258, no 4, p. 4p. 1-4Article in journal (Other (popular science, discussion, etc.))
  • 24.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, The Institute of Technology.
    Supporters of sperm: The 12th  Biology of Spermatozoa meeting, Hassop Hall, Derbyshire, UK2014In: Spermatogenesis, ISSN 2156-5554, Vol. 4, no 1, article id e27596Article in journal (Other academic)
    Abstract [en]

    The Biology of Spermatozoa (BoS) meetings have run on a biannual basis since the early 1990s. They are dedicated to the fascinating research topic of sperm and their complicated route to fertilization. The BoS meetings focus on sperm, but they also explore additional supporting factors important in fertilization, such as those present in seminal and ovarian fluid, as well as the genomic bases of sperm biology. Here, I present a report of the recent BoS meeting, and showcase some of the highlights of this year’s meeting.

  • 25.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Tio saker du troligen inte visste om höns2017Other (Other (popular science, discussion, etc.))
  • 26.
    Løvlie, Hanne
    et al.
    Stockholm University, Sweden.
    Cornwallis, Charles K.
    University of Sheffield, UK.
    Pizzari, Tommaso
    University of Oxford, UK.
    Male mounting alone reduces female promiscuity in the fowl2005In: Current Biology, ISSN 0960-9822, E-ISSN 1879-0445, Vol. 15, no 13, p. 1222-1227Article in journal (Refereed)
    Abstract [en]

    The fertilization success of an insemination is at risk when a female has the possibility to copulate with multiple males, generating sperm competition and sexual conflict over remating. Female propensity to remate is often reduced after copulation, and a staggering diversity of highly derived male traits that discourage female promiscuity have been investigated. However, it is difficult to separate the effect of such specialized traits and insemination products from the more basic effect that the act of mounting per se may have on female remating. Here, we use a novel approach that separates the influence of mounting from that of insemination on female remating in the promiscuous feral fowl. Mounting alone caused a transient but drastic reduction in female propensity to remate, and-crucially-the number of sperm that a female obtained from a new male. Therefore, like other taxa, female fowl show a reduction in promiscuity after copulation, but this is entirely due to mounting alone. This effect of mounting, independent of insemination and fertilization, indicates that even copulations that deliver little or no semen, a puzzling behavior common in many species including the fowl, may play a crucial role in sperm competition.

  • 27.
    Løvlie, Hanne
    et al.
    Edward Grey Institute, University of Oxford, UK Stockholm University, Stockholm, Sweden.
    Gillingham, Mark A. F.
    Edward Grey Institute, University of Oxford, UK.
    Worley, Kirsty
    Edward Grey Institute, University of Oxford, UK .
    Pizzari, Tommaso
    Edward Grey Institute, University of Oxford, UK.
    Richardson, David S.
    School of Biological Sciences, University of East Anglia, Norwich, UK.
    Cryptic female choice favours sperm from major histocompatibility complex-dissimilar males2013In: Proceedings of the Royal Society of London. Biological Sciences, ISSN 0962-8452, E-ISSN 1471-2954, Vol. 280, no 1769Article in journal (Refereed)
    Abstract [en]

    Cryptic female choice may enable polyandrous females to avoid inbreedingor bias offspring variability at key loci after mating. However, the role ofthese genetic benefits in cryptic female choice remains poorly understood.Female red junglefowl, Gallus gallus, bias sperm use in favour of unrelatedmales. Here, we experimentally investigate whether this bias is driven byrelatedness per se, or by similarity at the major histocompatibility complex(MHC), genes central to vertebrate acquired immunity, where polymorphismis critical to an individual’s ability to combat pathogens. Throughexperimentally controlled natural matings, we confirm that selection againstrelated males’ sperm occurs within the female reproductive tract but demonstratethat this is more accurately predicted by MHC similarity: controllingfor relatedness per se, more sperm reached the eggs when partners wereMHC-dissimilar. Importantly, this effect appeared largely owing to similarityat a single MHC locus (class I minor). Further, the effect of MHCsimilarity was lost following artificial insemination, suggesting that malephenotypic cues might be required for females to select sperm differentially.These results indicate that postmating mechanisms that reduce inbreedingmay do so as a consequence of more specific strategies of cryptic femalechoice promoting MHC diversity in offspring.

  • 28.
    Løvlie, Hanne
    et al.
    Animal Ecology, Department of Ecology and Genetics, Uppsala University, Uppsala, Sweden.
    Immonen, Elina
    Animal Ecology, Department of Ecology and Genetics, Uppsala University, Uppsala, Sweden.
    Gustavsson, Emil
    Animal Ecology, Department of Ecology and Genetics, Uppsala University, Uppsala, Sweden.
    Kazancioglu, Emil
    Animal Ecology, Department of Ecology and Genetics, Uppsala University, Uppsala, Sweden.
    Arnqvist, Göran
    Animal Ecology, Department of Ecology and Genetics, Uppsala University, Uppsala, Sweden.
    The influence of mitonuclear genetic variation on personality in seed beetles2014In: Proceedings of the Royal Society of London. Biological Sciences, ISSN 0962-8452, E-ISSN 1471-2954, Vol. 281, no 1796Article in journal (Refereed)
    Abstract [en]

    There is a growing awareness of the influence of mitochondrial genetic variation on life-history phenotypes, particularly via epistatic interactions with nuclear genes. Owing to their direct effect on traits such as metabolic and growth rates, mitonuclear interactions may also affect variation in behavioural types or personalities (i.e. behavioural variation that is consistent within individuals, but differs among individuals). However, this possibility is largely unexplored. We used mitonuclear introgression lines, where three mitochondrial genomes were introgressed into three nuclear genetic backgrounds, to disentangle genetic effects on behavioural variation in a seed beetle. We found within-individual consistency in a suite of activity-related behaviours, providing evidence for variation in personality. Composite measures of overall activity of individuals in behavioural assays were influenced by both nuclear genetic variation and by the interaction between nuclear and mitochondrial genomes. More importantly, the degree of expression of behavioural and life-history phenotypes was correlated and mitonuclear genetic variation affected expression of these concerted phenotypes. These results show that mitonuclear genetic variation affects both behavioural and life-history traits, and they provide novel insights into the maintenance of genetic variation in behaviour and personality.

  • 29.
    Løvlie, Hanne
    et al.
    Stockholm University, Sweden.
    Pizzari, Tommaso
    University of Oxford, UK.
    Sex in the morning or in the evening? Females adjust daily mating patterns to the intensity of sexual harassment2007In: American Naturalist, ISSN 0003-0147, E-ISSN 1537-5323, Vol. 170, no 1Article in journal (Refereed)
    Abstract [en]

    Selection on males to mate at a higher rate than females often results in male harassment of females and counteracting female responses. When the reproductive value of copulation changes over time, these mating strategies are expected to be time dependent. Here, we demonstrate that variation in the intensity of male harassment leads to drastic changes in female daily mating patterns. In feral populations of fowl Gallus gallus domesticus, male harassment is intense, particularly in the evening when inseminations are most likely to result in fertilization. We experimentally manipulated the intensity of male harassment through similar-sized groups of different sex ratios. Male mating propensity was always higher than females', particularly in male-biased groups and in the evening, when males were closer to and more likely to approach females. Females counteracted male harassment by escalating resistance to mating and--crucially--by shifting their daily mating pattern: in strongly female-biased groups with relaxed sexual harassment, females solicited sex in the evening, while in male-biased groups, they solicited sex in the morning, thus avoiding harassment in the evening. Together, these results indicate that intersexual conflict may occur not only over mating rates but also over when in the day to copulate.

  • 30.
    Løvlie, Hanne
    et al.
    Linköping University, The Institute of Technology. Linköping University, Department of Physics, Chemistry and Biology, Biology. Stockholm University, Sweden.
    Zidar, Josefina
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, The Institute of Technology. Stockholm University, Sweden.
    Berneheim, Christina
    School of Biological Sciences, The University of Edinburgh, Edinburgh, UK.
    A cry for help: female distress calling during copulation is context dependent2014In: Animal Behaviour, ISSN 0003-3472, E-ISSN 1095-8282, Vol. 92, p. 151-157Article in journal (Refereed)
    Abstract [en]

    Owing to selection for increased mating propensity, males often expose females to sexual harassment.Consequently, females may evolve counterstrategies to retain control of mating. Females can do thisdirectly by resisting copulations, or indirectly by manipulating other males to intervene and prevent thecopulation. Uttering copulation calls may be one indirect method for females to trigger male intervention.Copulation calls are commonly observed in mammals, primarily in primates, and also in some birds.Female fowl, Gallus gallus, sometimes utter calls during copulation, particularly in forced copulationswith low-ranking males. These loud calls, called distress calls, attract other males and can result indisruption of the copulation, and subsequent mating with the intervening male if he is high ranking.Consequently, uttering such calls can act both to abort a coerced copulation and to generate novel opportunitiesfor females to copulate with higher-ranking males. Nevertheless, uttering loud calls can carrycosts, such as attracting predators. Females are therefore predicted to utter copulation calls primarilywhen doing so offers benefits, which for female fowl requires the presence of another high-ranking male.We tested this prediction by altering the social environment of female domestic fowl, G. g. domesticus.We found that females uttered copulation calls more frequently during copulations in the presence ofdominant ‘observer’ males than in their absence. Thus, we provide evidence of context-dependent utteranceof female calls during copulations in a bird. This type of female vocalization is rarely investigatedin nonprimate vertebrates, but increased research in this field offers potential to improve understandingof female mate choice strategies and the dynamics of sexual selection.

  • 31.
    Maklakov, Alexei A
    et al.
    Department of Animal Ecology, Evolutionary Biology Centre, Uppsala University, Sweden.
    Immler, Simone
    Department of Evolutionary Biology, Evolutionary Biology Centre, Uppsala, Sweden.
    Løvlie, Hanne
    Department of Animal Ecology, Evolutionary Biology Centre, Uppsala University, Sweden.
    Flis, Ilona
    Department of Evolutionary Biology, Evolutionary Biology Centre, Uppsala, Sweden.
    Friberg, Urban
    Department of Evolutionary Biology, Evolutionary Biology Centre, Uppsala, Sweden.
    The effect of sexual harassment on lethal mutation rate in female Drosophila melanogaster2013In: Proceedings of the Royal Society of London. Biological Sciences, ISSN 0962-8452, E-ISSN 1471-2954, Vol. 280, no 1750Article in journal (Refereed)
    Abstract [en]

    The rate by which new mutations are introduced into a population may have far-reaching implications for processes at the population level. Theory assumes that all individuals within a population have the same mutation rate, but this assumption may not be true. Compared with individuals in high condition, those in poor condition may have fewer resources available to invest in DNA repair, resulting in elevated mutation rates. Alternatively, environmentally induced stress can result in increased investment in DNA repair at the expense of reproduction. Here, we directly test whether sexual harassment by males, known to reduce female condition, affects female capacity to alleviate DNA damage in Drosophila melanogaster fruitflies. Female gametes can repair double-strand DNA breaks in sperm, which allows manipulating mutation rate independently from female condition. We show that male harassment strongly not only reduces female fecundity, but also reduces the yield of dominant lethal mutations, supporting the hypothesis that stressed organisms invest relatively more in repair mechanisms. We discuss our results in the light of previous research and suggest that social effects such as density and courtship can play an important and underappreciated role in mediating condition-dependent mutation rate.

  • 32.
    Meziane, El Kahina
    et al.
    Univ Cambridge, England.
    Potts, Nicola D.
    Univ Cambridge, England.
    Viertlboeck, Birgit C.
    Ludwig Maximillian Univ, Germany.
    Lovlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Krupa, Andrew P.
    Univ Sheffield, England.
    Burke, Terry A.
    Univ Sheffield, England.
    Brown, Stewart
    Aviagen Ltd, Scotland.
    Watson, Kellie A.
    Univ Edinburgh, Scotland.
    Richardson, David S.
    Univ East Anglia, England.
    Pizzari, Tommaso
    Univ Oxford, England.
    Goebel, Thomas W.
    Ludwig Maximillian Univ, Germany.
    Kaufman, Jim
    Univ Cambridge, England.
    Bi-Functional Chicken Immunoglobulin-Like Receptors With a Single Extracellular Domain (ChIR-AB1): Potential Framework Genes Among a Relatively Stable Number of Genes Per Haplotype2019In: Frontiers in Immunology, ISSN 1664-3224, E-ISSN 1664-3224, Vol. 10, article id 2222Article in journal (Refereed)
    Abstract [en]

    The leukocyte receptor complex (LRC) in humans encodes many receptors with immunoglobulin-like (Ig-like) extracellular domains, including the killer Ig-like receptors (KIRs) expressed on natural killer (NK) cells among others, the leukocyte Ig-like receptors (LILRs) expressed on myeloid and B cells, and an Fc receptor (FcR), all of which have important roles in the immune response. These highly-related genes encode activating receptors with positively-charged residues in the transmembrane region, inhibitory receptors with immuno-tyrosine based motifs (ITIMs) in the cytoplasmic tail, and bi-functional receptors with both. The related chicken Ig-like receptors (ChIRs) are almost all found together on a microchromosome, with over 100 activating (A), inhibitory (B), and bi-functional (AB) genes, bearing either one or two extracellular Ig-like domains, interspersed over 500-1,000 kB in the genome of an individual chicken. Sequencing studies have suggested rapid divergence and little overlap between ChIR haplotypes, so we wished to begin to understand their genetics. We chose to use a hybridization technique, reference strand-mediated conformational analysis (RSCA), to examine the ChIR-AB1 family, with a moderate number of genes dispersed across the microchromosome. Using fluorescently-labeled references (FLR), we found that RSCA and sequencing of ChIR-AB1 extracellular exon gave two groups of peaks with mobility correlated with sequence relationship to the FLR. We used this system to examine widely-used and well-characterized experimental chicken lines, finding only one or a few simple ChIR haplotypes for each line, with similar numbers of peaks overall. We found much more complicated patterns from a broiler line from a commercial breeder and a flock of red junglefowl, but trios of parents and offspring from another commercial chicken line show that the complicated patterns are due to heterozygosity, indicating a relatively stable number of peaks within haplotypes of these birds. Some ChIR-AB1 peaks were found in all individuals from the commercial lines, and some of these were shared with red junglefowl and the experimental lines derived originally from egg-laying chickens. Overall, this analysis suggests that there are some simple features underlying the apparent complexity of the ChIR locus.

  • 33.
    Olofsson, Martin
    et al.
    Department of Zoology, Stockholm University, Sweden.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, The Institute of Technology. Stockholm University, Stockholm, Sweden Uppsala University, Uppsala, Sweden.
    Tibblin, Jessika
    Department of Zoology, Stockholm University, Sweden.
    Jakobsson, Sven
    Department of Zoology, Stockholm University, Sweden.
    Wiklund, Christer
    Department of Zoology, Stockholm University, Sweden.
    Eyespot display in the peacock butterfly triggers antipredator behaviors in naïve adult fowl2013In: Behavioral Ecology, ISSN 1045-2249, E-ISSN 1465-7279, Vol. 24, no 1, p. 305-310Article in journal (Refereed)
    Abstract [en]

    Large conspicuous eyespots have evolved in multiple taxa and presumably function to thwart predator attacks. Traditionally,large eyespots were thought to discourage predator attacks because they mimicked eyes of the predators’ own predators.However, this idea is controversial and the intimidating properties of eyespots have recently been suggested to simply be a consequenceof their conspicuousness. Some lepidopteran species include large eyespots in their antipredation repertoire. In thepeacock butterfly, Inachis io, eyespots are typically hidden during rest and suddenly exposed by the butterfly when disturbed.Previous experiments have shown that small wild passerines are intimidated by this display. Here, we test whether eyespots alsointimidate a considerably larger bird, domestic fowl, Gallus gallus domesticus, by staging interactions between birds and peacockbutterflies that were sham-painted or had their eyespots painted over. Our results show that birds typically fled when peacockbutterflies performed their display regardless of whether eyespots were visible or painted over. However, birds confronting butterflieswith visible eyespots delayed their return to the butterfly, were more vigilant, and more likely to utter alarm calls associatedwith detection of ground-based predators, compared with birds confronting butterflies with eyespots painted over. Becauseproduction of alarm calls and increased vigilance are antipredation behaviors in the fowl, their reaction suggests that eyespotsmay elicit fear rather than just an aversion to conspicuous patterns. Our results, therefore, suggest that predators perceive largelepidopteran eyespots as belonging to the eyes of a potential predator. Key words: chicken, predator–prey interactions, startledisplay.

  • 34.
    Petkova, Irina
    et al.
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering. Royal Holloway Univ London, England.
    Abbey-Lee, Robin
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Lovlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Parasite infection and host personality: Glugea-infected three-spined sticklebacks are more social2018In: Behavioral Ecology and Sociobiology, ISSN 0340-5443, E-ISSN 1432-0762, Vol. 72, no 11, article id UNSP 173Article in journal (Refereed)
    Abstract [en]

    The existence of animal personality is now well-documented, although the causes and consequences of this phenomenon are still largely unclear. Parasite infection can have pervasive effects on hosts, including altering host behaviour, and may thus contribute to differences in host personality. We investigated the relationship between the three-spined stickleback and its common parasite Glugea anomala, with focus on differences in host personality. Naturally infected and uninfected individuals were assayed for the five personality traits activity, exploration, boldness, sociability, and aggression. If infected fish behaved differently from uninfected, to benefit this parasite with horizontal transmission, we predicted behaviour increasing interactions with other sticklebacks to increase. Infection status explained differences in host personality. Specifically, Glugea-infected individuals were more social than uninfected fish. This confirms a link between parasite infection and host behaviour, and a relationship which may improve the horizontal transmission of Glugea. However, future studies need to establish the consequences of this for the parasite, and the causality of the parasite-host personality relationship. Significance statement Parasite infection that alters host behaviour could be a possible avenue of research into the causes of animal personality. We studied the link between infection and personality using the three-spined stickleback and its parasite Glugea anomala. We predicted that infected individuals would be more prone to interact with other sticklebacks, since this would improve transmission of this parasite. The personality of uninfected and naturally infected fish was measured and we observed that Glugea-infected sticklebacks were more social. Our results confirm a link between parasitism and variation in host personality.

  • 35.
    Pizzari, Tommaso
    et al.
    Swedish University of Agricultural Sciences, Skara, Sweden.
    Cornwallis, Charles K.
    University of Sheffield, UK.
    Løvlie, Hanne
    Stockholm University, Sweden.
    Jakobsson, Sven
    Stockholm University, Sweden.
    Birkhead, Tim R.
    University of Sheffield, UK.
    Sophisticated sperm allocation in male fowl2003In: Nature, ISSN 0028-0836, E-ISSN 1476-4687, Vol. 426, p. 70-74Article in journal (Refereed)
    Abstract [en]

    When a female is sexually promiscuous, the ejaculates of different males compete for the fertilization of her eggs; the more sperm a male inseminates into a female, the more likely he is to fertilize her eggs. Because sperm production is limited and costly, theory predicts that males will strategically allocate sperm (1) according to female promiscuity, (2) saving some for copulations with new females, and (3) to females producing more and/or better offspring. Whether males allocate sperm in all of these ways is not known, particularly in birds where the collection of natural ejaculates only recently became possible. Here we demonstrate male sperm allocation of unprecedented sophistication in the fowl Gallus gallus. Males show status-dependent sperm investment in females according to the level of female promiscuity; they progressively reduce sperm investment in a particular female but, on encountering a new female, instantaneously increase their sperm investment; and they preferentially allocate sperm to females with large sexual ornaments signalling superior maternal investment. Our results indicate that female promiscuity leads to the evolution of sophisticated male sexual behaviour.

  • 36.
    Pizzari, Tommaso
    et al.
    University of Leeds, UK.
    Løvlie, Hanne
    Stockholm University, Sweden.
    Cornwallis, Charles K.
    University of Sheffield, UK.
    Sex-specific, counter-acting responses to inbreeding in a bird2004In: Proceedings of the Royal Society of London. Biological Sciences, ISSN 0962-8452, E-ISSN 1471-2954, Vol. 271, no 1553, p. 2115-2121Article in journal (Refereed)
    Abstract [en]

    Inbreeding often depresses offspring fitness. Because females invest more than males in a reproductive event, inbreeding is expected to be more costly to mothers than fathers, creating a divergence between the reproductive interests of each sex and promoting sex-specific inbreeding strategies. Males and females may bias the probability of inbreeding by selecting copulation partners, and, in sexually promiscuous species, through male strategic sperm investment in different females and female selection of the sperm of different males. However, these processes are often difficult to study, and the way that different male and female strategies interact to determine inbreeding remains poorly understood. Here we demonstrate sex-specific, counteracting responses to inbreeding in the promiscuous red junglefowl, Gallus gallus. First, a male was just as likely to copulate with his full-sib sister as with an unrelated female. In addition, males displayed a tendency to: (i) initiate copulation faster when exposed to an unrelated female than when exposed to a sister, and (ii) inseminate more sperm into sisters than into unrelated females. Second, females retained fewer sperm following insemination by brothers, thus reducing the risk of inbreeding and counteracting male inbreeding strategies.

  • 37.
    Qvarnström, Anna
    et al.
    Uppsala University, Sweden.
    Rudh, Andreas
    Uppsala University, Sweden.
    Edström, Torkel
    Uppsala University, Sweden.
    Ödeen, Anders
    Uppsala University, Sweden.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, The Institute of Technology. Stockholm University, Sweden.
    Tullberg, Birgitta S
    Stockholm University, Sweden.
    Coarse dark patterning functionally constrains adaptive shifts from aposematism to crypsis in strawberry poison frogs2014In: Evolution, ISSN 0014-3820, E-ISSN 1558-5646, Vol. 68, no 10, p. 2793-2803Article in journal (Refereed)
    Abstract [en]

    Ecological specialization often requires tight coevolution of several traits, which may constrain future evolutionary pathways and make species more prone to extinction. Aposematism and crypsis represent two specialized adaptations to avoid predation. We tested whether the combined effects of color and pattern on prey conspicuousness functionally constrain or facilitate shifts between these two adaptations. We combined data from 17 natural populations of strawberry poison frogs, Oophaga pumilio with an experimental approach using digitalized images of frogs and chickens as predators. We show that bright coloration often co-occurs with coarse patterning among the natural populations. Dull green frogs with coarse patterning are rare in nature but in the experiment they were as easily detected as bright red frogs suggesting that this trait combination represents a transient evolutionary state toward aposematism. Hence, a gain of either bright color or coarse patterning leads to conspicuousness, but a transition back to crypsis would be functionally constrained in populations with both bright color and coarse patterning by requiring simultaneous changes in two traits. Thus, populations (or species) signaling aposematism by conspicuous color should be less likely to face an evolutionary dead end and more likely to radiate than populations with both conspicuous color and coarse patterning.

  • 38.
    Rosher, Charlotte
    et al.
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering. Faculty of Life Sciences, University of Manchester, M13 9LP Manchester, UK.
    Favati, Anna
    Department of Zoology, Stockholm University, Sweden.
    Dean, Rebecca
    School of Biological Sciences, Clayton Campus, Monash University, Victoria 3800, Australia, and Department of Genetics, Evolution and Environment, University College London.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Relatedness and age reduce aggressive maleinteractions over mating in domestic fowl2017In: Behavioral Ecology, ISSN 1045-2249, E-ISSN 1465-7279, Vol. 28, no 3, p. 760-766Article in journal (Refereed)
    Abstract [en]

    Altruistic behaviour represents a fundamental challenge in evolutionary biology. It is often best understood through kin selection,

    where favourable behaviour is directed towards relatives. Kin selection can take place when males cooperate to enhance the reproductive

    success of relatives. Here, we focus on reduced male–male competition over mating as a case of cooperation, by examining

    male tolerance of matings by related and unrelated competitors. A suitable model for exploring whether relatedness affects male–male

    interactions over mating is the domestic fowl,

    Gallus gallus domesticus. In this species, males form social hierarchies and dominant

    males commonly interrupt subdominant males’ copulation attempts. We investigated whether dominant male fowl differentially direct

    aggressive interactions towards unrelated and related subordinate males during mating attempts. Dominant male fowl were found to

    interrupt mating attempts of male relatives less often than those of unrelated males. We further tested whether male age mediates the

    magnitude of kin tolerance behaviour. However, we found no support for this as both young and old dominant males were less likely to

    interrupt related, compared to unrelated, subdominant males’ copulations during male–male interactions. Our results, consistent with

    kin selection, provide a rare experimental demonstration of relatedness relaxing male–male competition over mating.

  • 39.
    Segami Marzal, Julia Carolina
    et al.
    Uppsala university, Uppsala, Sweden.
    Rudh, Andreas
    Uppsala university, Uppsala, Sweden.
    Rogell, Björn
    Stockholm university, Stockholm, Sweden.
    Ödeen, Anders
    Uppsala university, Uppsalal, Sweden.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Roscher, Charlotte
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering. Faculty of Life Sciences, University of Manchester, UK.
    Qvarnström, Anna
    Uppsala university, Uppsala, Sweden.
    Cryptic female Strawberry poison frogs experience elevated predation risk when associating with an aposematic partner2017In: Ecology and Evolution, ISSN 2045-7758, E-ISSN 2045-7758, Vol. 7, no 2, p. 744-750Article in journal (Refereed)
    Abstract [en]

    Population divergence in sexual signals may lead to speciation through prezygotic isolation.

    Sexual signals can change solely due to variation in the level of natural selection

    acting against conspicuousness. However, directional mate choice (i.e., favoring conspicuousness)

    across different environments may lead to gene flow between populations,

    thereby delaying or even preventing the evolution of reproductive barriers and

    speciation. In this study, we test whether natural selection through predation upon

    mate-choosing

    females can favor corresponding changes in mate preferences. Our

    study system, Oophaga pumilio, is an extremely color polymorphic neotropical frog

    with two distinctive antipredator strategies: aposematism and crypsis. The conspicuous

    coloration and calling behavior of aposematic males may attract both cryptic and

    aposematic females, but predation may select against cryptic females choosing aposematic

    males. We used an experimental approach where domestic fowl were encouraged

    to find digitized images of cryptic frogs at different distances from aposematic

    partners. We found that the estimated survival time of a cryptic frog was reduced

    when associating with an aposematic partner. Hence, predation may act as a direct

    selective force on female choice, favoring evolution of color assortative mating that, in

    turn, may strengthen the divergence in coloration that natural selection has

    generated

  • 40.
    Sorato, Enrico
    et al.
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Zidar, Josefina
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Garnham, Laura
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Wilson, Alastair
    Centre for Ecology and Conservation, University of Exeter, Penryn Campus, Penryn, Cornwall, UK.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Heritabilities and co-variation among cognitive traits in red junglefowl2018In: Philosophical Transactions of the Royal Society of London. Biological Sciences, ISSN 0962-8436, E-ISSN 1471-2970, Vol. 373, no 1756, article id 20170285Article in journal (Refereed)
    Abstract [en]

    Natural selection can act on between-individual variation in cognitive abilities, yet evolutionary responses depend on the presence of underlying genetic variation. It is, therefore, crucial to determine the relative extent of genetic versus environmental control of these among-individual differences in cognitive traits to understand their causes and evolutionary potential. We investigated heritability of associative learning performance and of a cognitive judgement bias (optimism), as well as their covariation, in a captive pedigree-bred population of red junglefowl (Gallus gallusn > 300 chicks over 5 years). We analysed performance in discriminative and reversal learning (two facets of associative learning), and cognitive judgement bias, by conducting animal models to disentangle genetic from environmental contributions. We demonstrate moderate heritability for reversal learning, and weak to no heritability for optimism and discriminative learning, respectively. The two facets of associative learning were weakly negatively correlated, consistent with hypothesized trade-offs underpinning individual cognitive styles. Reversal, but not discriminative learning performance, was associated with judgement bias; less optimistic individuals reversed a previously learnt association faster. Together these results indicate that genetic and environmental contributions differ among traits. While modular models of cognitive abilities predict a lack of common genetic control for different cognitive traits, further investigation is required to fully ascertain the degree of covariation between a broader range of cognitive traits and the extent of any shared genetic control.

  • 41.
    Tan, Cedric K. W.
    et al.
    University of Oxford, England .
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, The Institute of Technology.
    Greenway, Elisabeth
    University of Oxford, England University of St Andrews, Scotland .
    Goodwin, Stephen F.
    University of Oxford, England .
    Pizzari, Tommaso
    University of Oxford, England .
    Wigby, Stuart
    University of Oxford, England .
    Correction: Sex-specific responses to sexual familiarity, and the role of olfaction in Drosophila: a new analysis confirms original results (vol 280, 20131691, 2013)2014In: Proceedings of the Royal Society of London. Biological Sciences, ISSN 0962-8452, E-ISSN 1471-2954, Vol. 281, no 1783Article in journal (Other academic)
    Abstract [en]

    n/a

  • 42.
    Tan, Cedric K.W.
    et al.
    Edward Grey Institute, Department of Zoology, University of Oxford, UK.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, The Institute of Technology. Oxford University, Oxford, UK.
    Greenway, Elisabeth
    Edward Grey Institute, Department of Zoology, University of Oxford, UK.
    Goodwin, Stephen F.
    Department of Physiology Anatomy and Genetics, University of Oxford, UK.
    Pizzari, Tommaso
    Edward Grey Institute, Department of Zoology, University of Oxford, UK.
    Wigby, Stuart
    Edward Grey Institute, Department of Zoology, University of Oxford, UK.
    Sex-specific responses to sexual familiarity, and the role of olfaction in Drosophila2013In: Proceedings of the Royal Society of London. Biological Sciences, ISSN 0962-8452, E-ISSN 1471-2954, Vol. 280, no 1771, p. 20131691-Article in journal (Refereed)
    Abstract [en]

    Studies of mating preferences have largely neglected the potential effects ofindividuals encountering their previous mates (‘directly sexually familiar’),or new mates that share similarities to previous mates, e.g. from the samefamily and/or environment (‘phenotypically sexually familiar’). Here, weshow that male and female Drosophila melanogaster respond to the direct andphenotypic sexual familiarity of potential mates in fundamentally differentways. We exposed a single focal male or female to two potential partners. Inthe first experiment, one potential partner was novel (not previously encountered)and one was directly familiar (their previous mate); in the secondexperiment, one potential partner was novel (unrelated, and from a differentenvironment from the previous mate) and one was phenotypically familiar(from the same family and rearing environment as the previous mate). Wefound that males preferentially courted novel females over directly or phenotypicallyfamiliar females. By contrast, females displayed aweak preference fordirectly and phenotypically familiar males over novel males. Sex-specificresponses to the familiarity of potential mates were significantly weaker orabsent in Orco1 mutants, which lack a co-receptor essential for olfaction, indicatinga role for olfactory cues in mate choice over novelty. Collectively, ourresults show that direct and phenotypic sexual familiarity is detected througholfactory cues and play an important role in sex-specific sexual behaviour.

  • 43.
    Tan, Cedric K.W.
    et al.
    Edward Grey Institute, Department of Zoology, University of Oxford, Oxford, U.K..
    Løvlie, Hanne
    Edward Grey Institute, Department of Zoology, University of Oxford, Oxford, U.K. Uppsala University, Uppsala, Sweden.
    Pizzari, Tommaso
    Edward Grey Institute, Department of Zoology, University of Oxford, Oxford, U.K..
    Wigby, Stuart
    Edward Grey Institute, Department of Zoology, University of Oxford, Oxford, U.K..
    No evidence for precopulatory inbreeding avoidance in Drosophila melanogaster2012In: Animal Behaviour, ISSN 0003-3472, E-ISSN 1095-8282, Vol. 83, no 6, p. 1433-1441Article in journal (Refereed)
    Abstract [en]

    Inbreeding depression can lead to the evolution of inbreeding avoidance before or after mating. However, despite widespread evidence of inbreeding depression, studies of inbreeding avoidance have generated different results across populations or species. These differences could potentially reflect the confounding effects of factors such as magnitude of inbreeding depression, sex, social familiarity, state of primary sexual receptivity and mating history. We examined the influence of these proximate factors on precopulatory inbreeding avoidance in a laboratory-adapted, outbred population of Drosophila melanogaster. We found a significant but low coefficient of inbreeding depression based on egg–adult viability measures. Controlling for sex-specific responses, familiarity, sexual receptivity and mating history, we found no evidence of precopulatory inbreeding avoidance. Mate choice of virgins was random with respect to relatedness and measurements of courtship frequency, mating latency and mating duration did not indicate any preference for unrelated partners. In fact, the only evidence for differential sexual behaviour in response to relatedness was that males first mated to unrelated females were significantly faster to remate with related females than with unrelated females. These results suggest that inbreeding avoidance may be limited in outbred populations of D. melanogaster, and fit theoretical predictions that inbreeding is not selected against in either sex when the coefficient of inbreeding depression is relatively low

  • 44.
    Tsuboi, Masahito
    et al.
    Uppsala Univ, Dept Ecol & Genet Anim Ecol, Evolutionary Biol Ctr, SE-75236 Uppsala, Sweden.
    Husby, Arild
    Univ Helsinki, Dept Biosci, FI-00014 Helsinki, Finland.
    Kotrschal, Alexander
    Stockholm Univ, Dept Zool Ethol, SE-10691 Stockholm, Sweden.
    Hayward, Alexander
    Stockholm Univ, Dept Zool Ethol, SE-10691 Stockholm, Sweden.
    Buechel, Severine D.
    ETH, Inst Integrat Biol IBZ, CH-8092 Zurich, Switzerland.
    Zidar, Josefina
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, The Institute of Technology.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, The Institute of Technology.
    Kolm, Niclas
    Stockholm Univ, Dept Zool Ethol, SE-10691 Stockholm, Sweden.
    Comparative support for the expensivetissue hypothesis: Big brains are correlatedwith smaller gut and greater parentalinvestment in Lake Tanganyika cichlids2014In: Evolution, ISSN 0014-3820, E-ISSN 1558-5646, Vol. 263, p. 33-38Article in journal (Refereed)
    Abstract [en]

    The brain is one of the most energetically expensive organs in the vertebrate body. Consequently, the energetic requirementsof encephalization are suggested to impose considerable constraints on brain size evolution. Three main hypotheses concerninghow energetic constraints might affect brain evolution predict covariation between brain investment and (1) investment intoother costly tissues, (2) overall metabolic rate, and (3) reproductive investment. To date, these hypotheses have mainly beentested in homeothermic animals and the existing data are inconclusive. However, there are good reasons to believe that energeticlimitations might play a role in large-scale patterns of brain size evolution also in ectothermic vertebrates. Here, we test thesehypotheses in a group of ectothermic vertebrates, the Lake Tanganyika cichlid fishes. After controlling for the effect of sharedancestry and confounding ecological variables, we find a negative association between brain size and gut size. Furthermore, wefind that the evolution of a larger brain is accompanied by increased reproductive investment into egg size and parental care. Ourresults indicate that the energetic costs of encephalization may be an important general factor involved in the evolution of brainsize also in ectothermic vertebrates.

  • 45.
    Tsuboi, Masahito
    et al.
    Evolutionary Biology Centre, Department of Ecology and Genetics/Animal Ecology, Uppsala University, Uppsala, Sweden.
    Kotrschal, Alexander
    Department of Zoology/Ethology, Stockholm University, Stockholm, Sweden.
    Hayward, Alexander
    Department of Zoology/Ethology, Stockholm University, Stockholm, Sweden.
    Buechel, Severine Denise
    Department of Zoology/Ethology, Stockholm University, Stockholm, Sweden.
    Zidar, Josefina
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Lovlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Kolm, Niclas
    Department of Zoology/Ethology, Stockholm University, Stockholm, Sweden.
    Evolution of brain-body allometry in Lake Tanganyika cichlids.2016In: Evolution, ISSN 0014-3820, E-ISSN 1558-5646, Vol. 70, no 7, p. 1559-1568Article in journal (Refereed)
    Abstract [en]

    Brain size is strongly associated with body size in all vertebrates. This relationship has been hypothesized to be an important constraint on adaptive brain size evolution. The essential assumption behind this idea is that static (i.e., within species) brain-body allometry has low ability to evolve. However, recent studies have reported mixed support for this view. Here, we examine brain-body static allometry in Lake Tanganyika cichlids using a phylogenetic comparative framework. We found considerable variation in the static allometric intercept, which explained the majority of variation in absolute and relative brain size. In contrast, the slope of the brain-body static allometry had relatively low variation, which explained less variation in absolute and relative brain size compared to the intercept and body size. Further examination of the tempo and mode of evolution of static allometric parameters confirmed these observations. Moreover, the estimated evolutionary parameters indicate that the limited observed variation in the static allometric slope could be a result of strong stabilizing selection. Overall, our findings suggest that the brain-body static allometric slope may represent an evolutionary constraint in Lake Tanganyika cichlids.

  • 46.
    Zidar, Josefina
    et al.
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Balogh, Alexandra
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, The Institute of Technology.
    Favati, Anna
    Stockholm University, Sweden.
    Jensen, Per
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Leimar, Olof
    Stockholm University, Sweden.
    Lovlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    A comparison of animal personality and coping styles in the red junglefowl2017In: Animal Behaviour, ISSN 0003-3472, E-ISSN 1095-8282, Vol. 130, p. 209-220Article in journal (Refereed)
    Abstract [en]

    There is an increased focus in biology on consistent behavioural variation. Several terms are used to describe this variation, including animal personality and coping style. Both terms describe between individual consistency in behavioural variation; however, they differ in the behavioural assays typically used, the expected distribution of response variables, and whether they incorporate variation in behavioural flexibility. Despite these differences, the terms are often used interchangeably. We conducted experiments using juvenile and adult red junglefowl, Gallus gallus, as subjects to explore the degree to which animal personality and coping styles overlap. We demonstrate that animal personality and coping styles can be described in this species, and that shyer individuals had more flexible responses, as expected for coping styles. Behavioural responses from both personality and coping style assays had continuous distributions, and were not clearly separated into two types. Behavioural traits were not correlated and, hence, there was no evidence of a behavioural syndrome. Further, behavioural responses obtained in personality assays did not correlate with those from coping style tests. Animal personality and coping styles are therefore not synonymous in the red junglefowl. We suggest that the terms animal personality and coping style are not equivalent and should not be used interchangeably. (C) 2017 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

  • 47.
    Zidar, Josefina
    et al.
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Balogh, Alexandra
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering. Stockholm Univ, Sweden.
    Favati, Anna
    Stockholm Univ, Sweden.
    Jensen, Per
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Leimar, Olof
    Stockholm Univ, Sweden.
    Sorato, Enrico
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Lovlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    The relationship between learning speed and personality is age- and task-dependent in red junglefowl2018In: Behavioral Ecology and Sociobiology, ISSN 0340-5443, E-ISSN 1432-0762, Vol. 72, no 10, article id UNSP 168Article in journal (Refereed)
    Abstract [en]

    Cognition is fundamental to animals lives and an important source of phenotypic variation. Nevertheless, research on individual variation in animal cognition is still limited. Further, although individual cognitive abilities have been suggested to be linked to personality (i.e., consistent behavioral differences among individuals), few studies have linked performance across multiple cognitive tasks to personality traits. Thus, the interplays between cognition and personality are still unclear. We therefore investigated the relationships between an important aspect of cognition, learning, and personality, by exposing young and adult red junglefowl (Gallus gallus) to multiple learning tasks (discriminative, reversal, and spatial learning) and personality assays (novel arena, novel object, and tonic immobility). Learning speed was not correlated across learning tasks, and learning speed in discrimination and spatial learning tasks did not co-vary with personality. However, learning speed in reversal tasks was associated with individual variation in exploration, and in an age-dependent manner. More explorative chicks learned the reversal task faster than less explorative ones, while the opposite association was found for adult females (learning speed could not be assayed in adult males). In the same reversal tasks, we also observed a sex difference in learning speed of chicks, with females learning faster than males. Our results suggest that the relationship between cognition and personality is complex, as shown by its task- and age-dependence, and encourage further investigation of the causality and dynamics of this relationship.Significance statementIn the ancestor of todays chickens, the red junglefowl, we explored how personality and cognition relate by exposing both chicks and adults to several learning tasks and personality assays. Our birds differed in personality and learning speed, while fast learners in one task did not necessarily learn fast in another (i.e., there were no overall smarter birds). Exploration correlated with learning speed in the more complex task of reversal learning: faster exploring chicks, but slower exploring adult females, learned faster, compared to less explorative birds. Other aspects of cognition and personality did not correlate. Our results suggest that cognition and personality are related, and that the relationship can differ depending on task and age of the animal.

  • 48.
    Zidar, Josefina
    et al.
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Campderrich, Irene
    Department of Animal Environment and Health, Swedish University of Agricultural Sciences, Uppsala, Sweden; Neiker-Tecnalia, Department of Animal Production, Vitoria-Gasteiz, Spain.
    Jansson, Emilie
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Wichman, Anette
    Department of Animal Environment and Health, Swedish University of Agricultural Sciences, Uppsala, Sweden.
    Winberg, Svante
    Department of Neuroscience, Uppsala Biomedical Centre BMC, Uppsala University, Uppsala, Sweden.
    Keeling, Linda
    Department of Animal Environment and Health, Swedish University of Agricultural Sciences, Uppsala, Sweden.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, Faculty of Science & Engineering.
    Environmental complexity buffers against stress-induced negative judgement bias in female chickens2018In: Scientific Reports, ISSN 2045-2322, E-ISSN 2045-2322, Vol. 8, no 5404Article in journal (Refereed)
    Abstract [en]

    Cognitive processes are often biased by emotions. In humans, affective disorders are accompanied by pessimistic judgement, while optimistic judgement is linked to emotional stability. Similar to humans, animals tend to interpret ambiguous stimuli negatively after experiencing stressful events, although the long-lasting impact on judgement bias has rarely been investigated. We measure judgement bias in female chicks (Gallus gallus domesticus) after exposure to cold stress, and before and after exposure to additional unpredictable stressors. Additionally, we explore if brain monoamines can explain differences in judgement bias. Chicks exposed to cold stress did not differ in judgement bias compared to controls, but showed sensitivity to additional stressors by having higher motivation for social reinstatement. Environmental complexity reduced stress-induced negative judgement bias, by maintaining an optimistic bias in individuals housed in complex conditions even after stress exposure. Moreover, judgement bias was related to dopamine turnover rate in mesencephalon, with higher activity in individuals that had a more optimistic response. These results demonstrate that environmental complexity can buffer against negative effects of additive stress and that dopamine relates to judgement bias in chicks. These results reveal that both internal and external factors can mediate emotionally biased judgement in animals, thus showing similarities to findings in humans.

  • 49.
    Zidar, Josefina
    et al.
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, The Institute of Technology.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology, Biology. Linköping University, The Institute of Technology.
    Fåglars Luktsinne2012In: Fauna och flora : populär tidskrift för biologi, ISSN 0014-8903, Vol. 107, no 4, p. 26-29Article in journal (Other (popular science, discussion, etc.))
  • 50.
    Zidar, Josefina
    et al.
    Stockholm University, Sweden.
    Løvlie, Hanne
    Linköping University, Department of Physics, Chemistry and Biology. Linköping University, The Institute of Technology. Stockholm University, Stockholm, Sweden.
    Scent of the enemy: behavioural responses to predator faecal odour in the fowl2012In: Animal Behaviour, ISSN 0003-3472, E-ISSN 1095-8282, Vol. 84, no 3, p. 547-554Article in journal (Refereed)
    Abstract [en]

    Chemical communication is used by diverse organisms in a variety of contexts and can have strong fitness consequences for the individuals involved. However, despite the extensive use of birds as models for many research areas in biology, avian olfaction has been poorly investigated. Studies on bird species that lack well-developed olfactory organs and those investigating responses to predator odours are particularly scarce. We investigated behavioural responses of the domestic fowl, Gallus gallus domesticus, a ground-living species with intermediate olfactory bulb size, to several predator and nonpredator faecal odours. We found that the birds spent less time foraging and were more vigilant when exposed to predator faecal odour compared with nonpredator faecal odour. Individuals showed a similar response when exposed to increased amounts of faeces. Taken together, our results demonstrate that domestic fowl can distinguish between herbivore and predator faecal odour, and respond to predator olfactory cues alone, without prior experience. Our results have implications for the understanding of predator-prey interactions and responses to olfactory cues in general, and for chemical communication in avian species more specifically.

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