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Variation in stress responses has been investigated in relation to environmental factors, species ecology, life history and fitness. Moreover, mechanistic studies have unravelled molecular mechanisms of how acute and chronic stress responses cause physiological impacts (‘damage’), and how this damage can be repaired. However, it is not yet understood how the fitness effects of damage and repair influence stress response evolution. Here we study the evolution of hormone levels as a function of stressor occurrence, damage and the efficiency of repair. We hypothesise that the evolution of stress responses depends on the fitness consequences of damage and the ability to repair that damage. To obtain some general insights, we model a simplified scenario in which an organism repeatedly encounters a stressor with a certain frequency and predictability (temporal autocorrelation). The organism can defend itself by mounting a stress response (elevated hormone level), but this causes damage that takes time to repair. We identify optimal strategies in this scenario and then investigate how those strategies respond to acute and chronic exposures to the stressor. We find that for higher repair rates, baseline and peak hormone levels are higher. This typically means that the organism experiences higher levels of damage, which it can afford because that damage is repaired more quickly, but for very high repair rates the damage does not build up. With increasing predictability of the stressor, stress responses are sustained for longer, because the animal expects the stressor to persist, and thus damage builds up. This can result in very high (and potentially fatal) levels of damage when organisms are exposed to chronic stressors to which they are not evolutionarily adapted. Overall, our results highlight that at least three factors need to be considered jointly to advance our understanding of how stress physiology has evolved: (i) temporal dynamics of stressor occurrence; (ii) relat

All organisms have a stress response system to cope with environmental threats, yet its precise form varies hugely within and across individuals, populations, and species. While the physiological mechanisms are increasingly understood, how stress responses have evolved remains elusive. Here, we show that important insights can be gained from models that incorporate physiological mechanisms within an evolutionary optimality analysis (the 'evo-mecho' approach). Our approach reveals environmental predictability and physiological constraints as key factors shaping stress response evolution, generating testable predictions about variation across species and contexts. We call for an integrated research programme combining theory, experimental evolution, and comparative analysis to advance scientific understanding of how this core physiological system has evolved.

All organisms have a stress response system to cope with environmental threats, yet its precise form varies hugely within and across individuals, populations, and species. While the physiological mechanisms are increasingly understood, how stress responses have evolved remains elusive. Here, we show that important insights can be gained from models that incorporate physiological mechanisms within an evolutionary optimality analysis (the ‘evo-mecho’ approach). Our approach reveals environmental predictability and physiological constraints as key factors shaping stress response evolution, generating testable predictions about variation across species and contexts. We call for an integrated research programme combining theory, experimental evolution, and comparative analysis to advance scientific understanding of how this core physiological system has evolved.

All organisms have a stress response system to cope with environmental threats, yet its precise form varies hugely within and across individuals, populations, and species. While the physiological mechanisms are increasingly understood, how stress responses have evolved remains elusive. Here, we show that important insights can be gained from models that incorporate physiological mechanisms within an evolutionary optimality analysis (the ‘evo-mecho’ approach). Our approach reveals environmental predictability and physiological constraints as key factors shaping stress response evolution, generating testable predictions about variation across species and contexts. We call for an integrated research programme combining theory, experimental evolution, and comparative analysis to advance scientific understanding of how this core physiological system has evolved.

In many vertebrate species, early social experience generates long-term effects on later life social behaviour. These effects are accompanied by persistent modifications in the expression of genes implicated in the stress axis. It is unknown, however, whether stress axis programming can affect the development of social competence, and if so, by which mechanism(s). Here, we used pharmacological manipulations to persistently reprogramme the hypothalamic-pituitary-interrenal axis of juvenile cooperatively breeding cichlids, Neolamprologus pulcher. During the first two months of life, juveniles were repeatedly treated with cortisol, mifepristone or control treatments. Three months after the last manipulation, we tested for treatment effects on (i) social competence, (ii) the expression of genes coding for corticotropin-releasing factor ( crf), glucocorticoid receptor ( gr1) and mineralocorticoid receptor ( mr) in the telencephalon and hypothalamus and (iii) cortisol levels. Social competence in a social challenge was reduced in cortisol-treated juveniles, which is in accordance with previous work applying early-life manipulations using different social experiences. During early life, both cortisol and mifepristone treatments induced a persistent downregulation of crf and upregulation of mr in the telencephalon. We suggest that these persistent changes in stress gene expression may represent an effective physiological mechanism for coping with stress. This article is part of the theme issue 'Developing differences: early-life effects and evolutionary medicine'.

In many vertebrate species, early social experience generates long-term effects on later life social behaviour. These effects are accompanied by persistent modifications in the expression of genes implicated in the stress axis. It is unknown, however, whether stress axis programming can affect the development of social competence, and if so, by which mechanism(s). Here, we used pharmacological manipulations to persistently reprogramme the hypothalamic-pituitary-interrenal axis of juvenile cooperatively breeding cichlids, Neolamprologus pulcher. During the first two months of life, juveniles were repeatedly treated with cortisol, mifepristone or control treatments. Three months after the last manipulation, we tested for treatment effects on (i) social competence, (ii) the expression of genes coding for corticotropin-releasing factor ( crf), glucocorticoid receptor ( gr1) and mineralocorticoid receptor ( mr) in the telencephalon and hypothalamus and (iii) cortisol levels. Social competence in a social challenge was reduced in cortisol-treated juveniles, which is in accordance with previous work applying early-life manipulations using different social experiences. During early life, both cortisol and mifepristone treatments induced a persistent downregulation of crf and upregulation of mr in the telencephalon. We suggest that these persistent changes in stress gene expression may represent an effective physiological mechanism for coping with stress. This article is part of the theme issue 'Developing differences: early-life effects and evolutionary medicine'.

Adult social behaviour can be persistently modified by early-life social experience. In rodents, such effects are induced by tactile maternal stimulation resulting in neuroendocrine modifications of the hypothalamic-pituitary-adrenal axis involved in stress responsiveness. Whether similar long-term alterations can occur in the hypothalamic-pituitary-interrenal (HPI) axis of poikilothermic vertebrates is unknown. We compared the expression of four genes of the HPI axis in adults of the cooperatively breeding cichlid Neolamprologus pulcher, which had been exposed to two early-life social treatments 1.5 years prior to brain sampling. Fish reared with parents and siblings had less brain expression of corticotropin-releasing factor and of the functional homologue of the mammalian glucocorticoid receptor (GR1) than individuals reared with same-age siblings only. Expression of the mineralocorticoid receptors (MR) did not differ between treatments, but the MR/GR1 expression ratio was markedly higher in fish reared with parents and siblings. Thus, we show here that early social experience can alter the programming of the stress axis in poikilothermic vertebrates, suggesting that this mechanism is deeply conserved within vertebrates. Moreover, we show for the first time that reprogramming of the stress axis of a vertebrate can be induced without tactile stimulation by parents.

Adult social behaviour can be persistently modified by early-life social experience. In rodents, such effects are induced by tactile maternal stimulation resulting in neuroendocrine modifications of the hypothalamic-pituitary-adrenal axis involved in stress responsiveness. Whether similar long-term alterations can occur in the hypothalamic-pituitary-interrenal (HPI) axis of poikilothermic vertebrates is unknown. We compared the expression of four genes of the HPI axis in adults of the cooperatively breeding cichlid Neolamprologus pulcher, which had been exposed to two early-life social treatments 1.5 years prior to brain sampling. Fish reared with parents and siblings had less brain expression of corticotropin-releasing factor and of the functional homologue of the mammalian glucocorticoid receptor (GR1) than individuals reared with same-age siblings only. Expression of the mineralocorticoid receptors (MR) did not differ between treatments, but the MR/GR1 expression ratio was markedly higher in fish reared with parents and siblings. Thus, we show here that early social experience can alter the programming of the stress axis in poikilothermic vertebrates, suggesting that this mechanism is deeply conserved within vertebrates. Moreover, we show for the first time that reprogramming of the stress axis of a vertebrate can be induced without tactile stimulation by parents.

The pay-off of deserting and leaving a mate to care for the offspring alone is generally assumed to depend mainly on the availability of alternative mating partners and on the potential spawning rate of males and females. Eretmodus cyanostictus is a monogamous mouthbrooding cichlid in which the clutch is successively incubated first by the female and then by the male. It has been suggested that parents are constrained to monogamy due to low remating probabilities for both sexes. We tested this hypothesis by varying the sex ratio experimentally. Mate desertion by either sex was not significantly higher when additional potential mates were present (males: 8.3%, females: 0%) than when there were no other same-sex conspecifics present (males: 0%, females: 0%). Males lost their mate to a male intruder during their incubation in 26.7% of cases. Pair members were more active and showed more aggression when same-sex conspecifics were present. Behavioural differences between treatments were strongest during the incubation period of a given sex. If no desertion takes place, sexual conflict may be expressed also on a second level, the amount of parental care each parent provides. Indeed, males took the offspring later when additional females were present, although male incubation time did not differ between treatments. A hitherto undescribed display behaviour of females was clear evidence of a conflict about the timing of shift of young. In conclusion, offering alternative mating opportunities did not strongly favour mate desertion in E. cyanostictus. It rather revealed a conflict between mates about when to shift the young

Through environmentally induced maternal effects females may fine-tune their offspring's phenotype to the conditions offspring will encounter after birth. If juvenile and adult ecologies differ, the conditions mothers experienced as juveniles may better predict their offspring's environment than the adult females' conditions. Maternal effects induced by the environment experienced by females during their early ontogeny should evolve when three ecological conditions are met: (1) Adult ecology does not predict the postnatal environmental conditions of offspring; (2) Environmental conditions for juveniles are correlated across successive generations; and (3) Juveniles occasionally settle in conditions that differ from the juvenile habitat of their mothers. By combining size-structured population counts, ecological surveys and a genetic analysis of population structure we provide evidence that all three conditions hold for Simochromis pleurospilus, a cichlid fish in which mothers adjust offspring quality to their own juvenile ecology. In particular we show (1) that the spatial niches and the habitat quality differ between juveniles and adults, and we provide genetic evidence (2) that usually fish of successive generations grow up in similar habitats, and (3) that occasional dispersal in populations with a different habitat quality is likely to occur. As adults of many species cannot predict their offspring's environment from ambient cues, life-stage specific maternal effects are likely to be common in animals. It will therefore be necessary to incorporate parental ontogeny in the study of parental effects when juveniles and adults inhabit different environments

Theoretical models predict that parents should adjust the amount of care both to their own and their partner's body condition. In most biparental species, parental duties are switched repeatedly allowing for repeated mutual adjustment of the amount of care. In the mouthbrooding cichlid Eretmodus cyanostictus, terms are switched only once with females taking the first share. The timing of the shift of the clutch between mates strongly determines both partners' brooding period and thereby their parental investment. Females signal their readiness to transfer the young several days before the male finally takes them, suggesting sexual conflict over the timing of the shift. In a lab experiment, we reduced the body condition of either the female or the male of a pair to test whether energy reserves affect the timing of the shift and whether female signalling behaviour depends on energetic state. Males with a lowered condition took the young later and incubated for a shorter period, which prolonged the incubation time of their female partners. When female condition was lowered, female and male incubation durations remained unchanged, although females signalled their readiness to shift more intensely. Our results suggest that males adjust their parental investment to own energy reserves but are unresponsive to their mate's condition. Females appear to carry the entire costs for the male's adjustment of care. We propose that intrinsic asymmetries in the scope for mutual adjustment of parental investment and the costs of negotiation crucially influence solutions of the conflict between sexes over care

Individuelle Verhaltensunterschiede (sog. „Animal personality“) werden oft als konsistent über die Zeit und situationsunabhängig angenommen. Vielfach werden solche Persönlichkeitsunterschiede zwischen Tieren einer Art durch individuelle Unterschiede im Energiehaushalt sowie Lebenszyklusvariablen (sog. ‚state variables‘, dt. Zustandsgrößen) erklärt. Dies ist in der „pace-of-life“ Hypothese zusammengefasst. In neueren Arbeiten wurde jedoch die Konsistenz von Persönlichkeitsmerkmalen über den Lebensverlauf und deren strikte Abhängigkeit von Zustandsgrößen in Frage gestellt. Die vorliegende Dissertation soll neue Einblicke in die Mechanismen, die zur Entstehung von Persönlichkeitsunterschieden im Verlauf des Lebens von Fischen, deren Situationsabhängigkeit sowie ihre Verbindung zu individuellen Unterschieden in Zustandsgrößen liefern. In einer Abfolge von fünf unabhängigen Studien untersuchte ich die genannten Annahmen und fand, dass (1) Persönlichkeitsunterschiede sich im Laufe des Lebens von Tieren vergrößern; (2) Persönlichkeitsabschätzungen bei jungen Tieren oft stärker vom experimentellen Aufbau beeinflusst werden als bei Erwachsenen; (3) der Energiehaushalt und Lebenszyklusvariablen Persönlichkeitsunterschiede sowohl unter Laborbedingungen als auch im Freiland nicht hinreichend erklären können; (4) Beziehungen zwischen Persönlichkeitsunterschieden und Unterschieden im Energiehaushalt und in Lebenszyklusvariablen fanden sich nur bei Fischen einer Population mit langsamer Lebenszyklusstrategie nicht jedoch in einer Population mit schnellem Lebenszyklus. Die vorliegende Arbeit suggeriert daher, dass sich erst im Verlauf des Lebens eines Tieres Persönlichkeitsunterschiede unvermeidbar entwickeln. Dies stellt Persönlichkeitsmessungen bei juvenilen Tieren grundsätzlich in Frage. Weiterhin scheinen Persönlichkeitsunterschiede und Zustandsgrößen unter bestimmten Umweltbedingungen und evolutiven Szenarien voneinander unabhängig zu sein., Among-individual differences in behavior (i.e., animal personality) are assumed to be consistent over time and contexts. In theory, they are often explained by individual variations in energy costs of self-maintenance as well as life history among animals (i.e., state variables), commonly expressed as the pace-of-life syndrome hypothesis. Yet, recent theories have disputed the consistency of personality types over lifetime and their rigid state-dependency. This thesis aims to offer novel insights on the mechanisms behind the emergence and development of personality over lifetime of fishes, its context dependency, and its link to individual variation in state variables. In a sequence of five independent yet interconnected studies, I tested the assumptions above and observed that (1) personality differences increased during lifetime as a function of consistent declines in the behavioral plasticity with increasing age of animals; (2) personality estimates in young animals were weaker and thus more vulnerable to experimental biases compared to adults; (3) personality variation did not depend upon individual differences in energy costs of self-maintenance and life-history traits among individuals, under both laboratory and natural settings; and (4) the relationship between behavioral, metabolic, and life-history traits was manifested only in fish populations with slow rather than fast life-history strategies. This thesis suggests that personality variation in animals might be the inevitable outcome of development, raising questions about the reliability of personality estimates in juvenile individuals. Furthermore, individual variation in personality and “states” may act independently (i.e., phenotypes are uncorrelated) under environmental conditions and evolutionary contexts that mask or select against their trade-offs.

Individuelle Verhaltensunterschiede (sog. „Animal personality“) werden oft als konsistent über die Zeit und situationsunabhängig angenommen. Vielfach werden solche Persönlichkeitsunterschiede zwischen Tieren einer Art durch individuelle Unterschiede im Energiehaushalt sowie Lebenszyklusvariablen (sog. ‚state variables‘, dt. Zustandsgrößen) erklärt. Dies ist in der „pace-of-life“ Hypothese zusammengefasst. In neueren Arbeiten wurde jedoch die Konsistenz von Persönlichkeitsmerkmalen über den Lebensverlauf und deren strikte Abhängigkeit von Zustandsgrößen in Frage gestellt. Die vorliegende Dissertation soll neue Einblicke in die Mechanismen, die zur Entstehung von Persönlichkeitsunterschieden im Verlauf des Lebens von Fischen, deren Situationsabhängigkeit sowie ihre Verbindung zu individuellen Unterschieden in Zustandsgrößen liefern. In einer Abfolge von fünf unabhängigen Studien untersuchte ich die genannten Annahmen und fand, dass (1) Persönlichkeitsunterschiede sich im Laufe des Lebens von Tieren vergrößern; (2) Persönlichkeitsabschätzungen bei jungen Tieren oft stärker vom experimentellen Aufbau beeinflusst werden als bei Erwachsenen; (3) der Energiehaushalt und Lebenszyklusvariablen Persönlichkeitsunterschiede sowohl unter Laborbedingungen als auch im Freiland nicht hinreichend erklären können; (4) Beziehungen zwischen Persönlichkeitsunterschieden und Unterschieden im Energiehaushalt und in Lebenszyklusvariablen fanden sich nur bei Fischen einer Population mit langsamer Lebenszyklusstrategie nicht jedoch in einer Population mit schnellem Lebenszyklus. Die vorliegende Arbeit suggeriert daher, dass sich erst im Verlauf des Lebens eines Tieres Persönlichkeitsunterschiede unvermeidbar entwickeln. Dies stellt Persönlichkeitsmessungen bei juvenilen Tieren grundsätzlich in Frage. Weiterhin scheinen Persönlichkeitsunterschiede und Zustandsgrößen unter bestimmten Umweltbedingungen und evolutiven Szenarien voneinander unabhängig zu sein., Among-individual differences in behavior (i.e., animal personality) are assumed to be consistent over time and contexts. In theory, they are often explained by individual variations in energy costs of self-maintenance as well as life history among animals (i.e., state variables), commonly expressed as the pace-of-life syndrome hypothesis. Yet, recent theories have disputed the consistency of personality types over lifetime and their rigid state-dependency. This thesis aims to offer novel insights on the mechanisms behind the emergence and development of personality over lifetime of fishes, its context dependency, and its link to individual variation in state variables. In a sequence of five independent yet interconnected studies, I tested the assumptions above and observed that (1) personality differences increased during lifetime as a function of consistent declines in the behavioral plasticity with increasing age of animals; (2) personality estimates in young animals were weaker and thus more vulnerable to experimental biases compared to adults; (3) personality variation did not depend upon individual differences in energy costs of self-maintenance and life-history traits among individuals, under both laboratory and natural settings; and (4) the relationship between behavioral, metabolic, and life-history traits was manifested only in fish populations with slow rather than fast life-history strategies. This thesis suggests that personality variation in animals might be the inevitable outcome of development, raising questions about the reliability of personality estimates in juvenile individuals. Furthermore, individual variation in personality and “states” may act independently (i.e., phenotypes are uncorrelated) under environmental conditions and evolutionary contexts that mask or select against their trade-offs.

Individuelle Verhaltensunterschiede (sog. „Animal personality“) werden oft als konsistent über die Zeit und situationsunabhängig angenommen. Vielfach werden solche Persönlichkeitsunterschiede zwischen Tieren einer Art durch individuelle Unterschiede im Energiehaushalt sowie Lebenszyklusvariablen (sog. ‚state variables‘, dt. Zustandsgrößen) erklärt. Dies ist in der „pace-of-life“ Hypothese zusammengefasst. In neueren Arbeiten wurde jedoch die Konsistenz von Persönlichkeitsmerkmalen über den Lebensverlauf und deren strikte Abhängigkeit von Zustandsgrößen in Frage gestellt. Die vorliegende Dissertation soll neue Einblicke in die Mechanismen, die zur Entstehung von Persönlichkeitsunterschieden im Verlauf des Lebens von Fischen, deren Situationsabhängigkeit sowie ihre Verbindung zu individuellen Unterschieden in Zustandsgrößen liefern. In einer Abfolge von fünf unabhängigen Studien untersuchte ich die genannten Annahmen und fand, dass (1) Persönlichkeitsunterschiede sich im Laufe des Lebens von Tieren vergrößern; (2) Persönlichkeitsabschätzungen bei jungen Tieren oft stärker vom experimentellen Aufbau beeinflusst werden als bei Erwachsenen; (3) der Energiehaushalt und Lebenszyklusvariablen Persönlichkeitsunterschiede sowohl unter Laborbedingungen als auch im Freiland nicht hinreichend erklären können; (4) Beziehungen zwischen Persönlichkeitsunterschieden und Unterschieden im Energiehaushalt und in Lebenszyklusvariablen fanden sich nur bei Fischen einer Population mit langsamer Lebenszyklusstrategie nicht jedoch in einer Population mit schnellem Lebenszyklus. Die vorliegende Arbeit suggeriert daher, dass sich erst im Verlauf des Lebens eines Tieres Persönlichkeitsunterschiede unvermeidbar entwickeln. Dies stellt Persönlichkeitsmessungen bei juvenilen Tieren grundsätzlich in Frage. Weiterhin scheinen Persönlichkeitsunterschiede und Zustandsgrößen unter bestimmten Umweltbedingungen und evolutiven Szenarien voneinander unabhängig zu sein., Among-individual differences in behavior (i.e., animal personality) are assumed to be consistent over time and contexts. In theory, they are often explained by individual variations in energy costs of self-maintenance as well as life history among animals (i.e., state variables), commonly expressed as the pace-of-life syndrome hypothesis. Yet, recent theories have disputed the consistency of personality types over lifetime and their rigid state-dependency. This thesis aims to offer novel insights on the mechanisms behind the emergence and development of personality over lifetime of fishes, its context dependency, and its link to individual variation in state variables. In a sequence of five independent yet interconnected studies, I tested the assumptions above and observed that (1) personality differences increased during lifetime as a function of consistent declines in the behavioral plasticity with increasing age of animals; (2) personality estimates in young animals were weaker and thus more vulnerable to experimental biases compared to adults; (3) personality variation did not depend upon individual differences in energy costs of self-maintenance and life-history traits among individuals, under both laboratory and natural settings; and (4) the relationship between behavioral, metabolic, and life-history traits was manifested only in fish populations with slow rather than fast life-history strategies. This thesis suggests that personality variation in animals might be the inevitable outcome of development, raising questions about the reliability of personality estimates in juvenile individuals. Furthermore, individual variation in personality and “states” may act independently (i.e., phenotypes are uncorrelated) under environmental conditions and evolutionary contexts that mask or select against their trade-offs.

Individuelle Verhaltensunterschiede (sog. „Animal personality“) werden oft als konsistent über die Zeit und situationsunabhängig angenommen. Vielfach werden solche Persönlichkeitsunterschiede zwischen Tieren einer Art durch individuelle Unterschiede im Energiehaushalt sowie Lebenszyklusvariablen (sog. ‚state variables‘, dt. Zustandsgrößen) erklärt. Dies ist in der „pace-of-life“ Hypothese zusammengefasst. In neueren Arbeiten wurde jedoch die Konsistenz von Persönlichkeitsmerkmalen über den Lebensverlauf und deren strikte Abhängigkeit von Zustandsgrößen in Frage gestellt. Die vorliegende Dissertation soll neue Einblicke in die Mechanismen, die zur Entstehung von Persönlichkeitsunterschieden im Verlauf des Lebens von Fischen, deren Situationsabhängigkeit sowie ihre Verbindung zu individuellen Unterschieden in Zustandsgrößen liefern. In einer Abfolge von fünf unabhängigen Studien untersuchte ich die genannten Annahmen und fand, dass (1) Persönlichkeitsunterschiede sich im Laufe des Lebens von Tieren vergrößern; (2) Persönlichkeitsabschätzungen bei jungen Tieren oft stärker vom experimentellen Aufbau beeinflusst werden als bei Erwachsenen; (3) der Energiehaushalt und Lebenszyklusvariablen Persönlichkeitsunterschiede sowohl unter Laborbedingungen als auch im Freiland nicht hinreichend erklären können; (4) Beziehungen zwischen Persönlichkeitsunterschieden und Unterschieden im Energiehaushalt und in Lebenszyklusvariablen fanden sich nur bei Fischen einer Population mit langsamer Lebenszyklusstrategie nicht jedoch in einer Population mit schnellem Lebenszyklus. Die vorliegende Arbeit suggeriert daher, dass sich erst im Verlauf des Lebens eines Tieres Persönlichkeitsunterschiede unvermeidbar entwickeln. Dies stellt Persönlichkeitsmessungen bei juvenilen Tieren grundsätzlich in Frage. Weiterhin scheinen Persönlichkeitsunterschiede und Zustandsgrößen unter bestimmten Umweltbedingungen und evolutiven Szenarien voneinander unabhängig zu sein., Among-individual differences in behavior (i.e., animal personality) are assumed to be consistent over time and contexts. In theory, they are often explained by individual variations in energy costs of self-maintenance as well as life history among animals (i.e., state variables), commonly expressed as the pace-of-life syndrome hypothesis. Yet, recent theories have disputed the consistency of personality types over lifetime and their rigid state-dependency. This thesis aims to offer novel insights on the mechanisms behind the emergence and development of personality over lifetime of fishes, its context dependency, and its link to individual variation in state variables. In a sequence of five independent yet interconnected studies, I tested the assumptions above and observed that (1) personality differences increased during lifetime as a function of consistent declines in the behavioral plasticity with increasing age of animals; (2) personality estimates in young animals were weaker and thus more vulnerable to experimental biases compared to adults; (3) personality variation did not depend upon individual differences in energy costs of self-maintenance and life-history traits among individuals, under both laboratory and natural settings; and (4) the relationship between behavioral, metabolic, and life-history traits was manifested only in fish populations with slow rather than fast life-history strategies. This thesis suggests that personality variation in animals might be the inevitable outcome of development, raising questions about the reliability of personality estimates in juvenile individuals. Furthermore, individual variation in personality and “states” may act independently (i.e., phenotypes are uncorrelated) under environmental conditions and evolutionary contexts that mask or select against their trade-offs.

In many animal taxa, size-selective predation favors fast growth early in life. However, same-aged juveniles can diverge in size due to differences in genotype, environmental conditions, and parental effects and thus may vary in competitive ability. Under food scarcity, competitively inferior juveniles may suffer suppressed growth, whereas under benign conditions, small juveniles may exhibit growth compensation and perform as well as large ones. However, studies testing this while controlling for parental effects are lacking. Here, we hand-raised cichlids, Simochromis pleurospilus, from a wide range of egg sizes and manipulated their size by differential feeding. Afterward, high- and low-ration siblings were kept in groups assigned to either a high- or low-competition environment. We investigated how the degree of competition affected aggressiveness and growth of juveniles with different feeding histories. As predicted, when competition was high, high-ration offspring grew fastest. Interestingly, when competition was weak, low-ration juveniles grew at a similar rate as high-ration ones and many were able to catch up in size. High-ration fish were more aggressive than low-ration ones, and this effect was strongest under high competition. Additionally, in the high-competition environment, received aggression was negatively related to growth, and inflicted aggression correlated positively with the growth of the aggressor. These relationships were absent under low competition. Our findings suggest that the abilities to compensate for early growth depression depend on the prevalent level of competition. Aggression is likely used to monopolize food by juvenile S. pleurospilus; however, when competition is strong, aggression cannot compensate for a size disadvantage

Mothers should adjust the size of propagules to the selective forces to which these offspring will be exposed. Usually, a larger propagule size is favored when young are exposed to high mortality risk or conspecific competition. Here we test 2 predictions on how egg size should vary with these selective agents. When offspring are cared for by parents and/or alloparents, protection may reduce the predation risk to young, which may allow mothers to invest less per single offspring. In the cooperatively breeding cichlid Neolamprologus pulcher, brood care helpers protect group offspring and reduce the latters' mortality rate. Therefore, females are expected to reduce their investment per egg when more helpers are present. In a first experiment, we tested this prediction by manipulating the helper number. In N. pulcher, helpers compete for dispersal opportunities with similar-sized individuals of neighboring groups. If the expected future competition pressure on young is high, females should increase their investment per offspring to give them a head start. In a second experiment, we tested whether females produce larger eggs when perceived neighbor density is high. Females indeed reduced egg size with increasing helper number. However, we did not detect an effect of local density on egg size, although females took longer to produce the next clutch when local density was high. We argue that females can use the energy saved by adjusting egg size to reduced predation risk to enhance future reproductive output. Adaptive adjustment of offspring size to helper number may be an important, as yet unrecognized, strategy of cooperative breeders

Social animals can greatly benefit from well-developed social skills. Because the frequency and diversity of social interactions often increase with the size of social groups, the benefits of advanced social skills can be expected to increase with group size. Variation in social skills often arises during ontogeny, depending on early social experience. Whether variation of social-group sizes affects development of social skills and related changes in brain structures remains unexplored. We investigated whether, in a cooperatively breeding cichlid, early group size (1) shapes social behavior and social skills and (2) induces lasting plastic changes in gross brain structures and (3) whether the development of social skills is confined to a sensitive ontogenetic period. Rearing-group size and the time juveniles spent in these groups interactively influenced the development of social skills and the relative sizes of four main brain regions. We did not detect a sensitive developmental period for the shaping of social behavior within the 2-month experience phase. Instead, our results suggest continuous plastic behavioral changes over time. We discuss how developmental effects on social behavior and brain architecture may adaptively tune phenotypes to their current or future environments.

Through environmentally induced maternal effects females may fine-tune their offspring's phenotype to the conditions offspring will encounter after birth. If juvenile and adult ecologies differ, the conditions mothers experienced as juveniles may better predict their offspring's environment than the adult females' conditions. Maternal effects induced by the environment experienced by females during their early ontogeny should evolve when three ecological conditions are met: (1) Adult ecology does not predict the postnatal environmental conditions of offspring; (2) Environmental conditions for juveniles are correlated across successive generations; and (3) Juveniles occasionally settle in conditions that differ from the juvenile habitat of their mothers. By combining size-structured population counts, ecological surveys and a genetic analysis of population structure we provide evidence that all three conditions hold for Simochromis pleurospilus, a cichlid fish in which mothers adjust offspring quality to their own juvenile ecology. In particular we show (1) that the spatial niches and the habitat quality differ between juveniles and adults, and we provide genetic evidence (2) that usually fish of successive generations grow up in similar habitats, and (3) that occasional dispersal in populations with a different habitat quality is likely to occur. As adults of many species cannot predict their offspring's environment from ambient cues, life-stage specific maternal effects are likely to be common in animals. It will therefore be necessary to incorporate parental ontogeny in the study of parental effects when juveniles and adults inhabit different environments.

The presence of fat stores in fish is widely used as a correlate of fish health and fitness. Techniques to measure fat content with some accuracy are available for medium-sized and large fish, but apart from morphometric indices, a noninvasive method to determine fat content in small fish has hitherto been lacking. In this study, we introduce a novel method to measure the fat content in live fish that can be applied also to small fish of less than 0.5 g of body mass. This approach relies on a precise measurement of the swim bladder volume, from which fat content can subsequently be deduced. As fat is positively buoyant, fish with larger fat stores require a smaller swim bladder to attain neutral buoyancy. To determine swim bladder volume, we developed a measuring device, which makes use of the differential compressibility of air and water. A fish is placed in a pressure-tight chamber to which a standardized amount of water is added. The resulting change in pressure Delta p is inversely proportional to the volume of the swim bladder. Using juveniles and adults of Simochromis pleurospilus (Nelissen, '78; Pisces: Tropheini) a small cichlid fish, we show that Delta p is tightly related to structural size, mass, and body condition. Most importantly, this approach allows to predict the visceral fat content of small fish more precisely than the six most commonly used morphometric body indices.

Maternal effects on offspring size can have a strong effect on fitness, as larger offspring often survive better under harsh environmental conditions. Selection should hence favour mothers that find an optimal solution to the offspring size versus number

Through environmentally induced maternal effects females may fine-tune their offspring's phenotype to the conditions offspring will encounter after birth. If juvenile and adult ecologies differ, the conditions mothers experienced as juveniles may better predict their offspring's environment than the adult females' conditions. Maternal effects induced by the environment experienced by females during their early ontogeny should evolve when three ecological conditions are met: (1) Adult ecology does not predict the postnatal environmental conditions of offspring; (2) Environmental conditions for juveniles are correlated across successive generations; and (3) Juveniles occasionally settle in conditions that differ from the juvenile habitat of their mothers. By combining size-structured population counts, ecological surveys and a genetic analysis of population structure we provide evidence that all three conditions hold for Simochromis pleurospilus, a cichlid fish in which mothers adjust offspring quality to their own juvenile ecology. In particular we show (1) that the spatial niches and the habitat quality differ between juveniles and adults, and we provide genetic evidence (2) that usually fish of successive generations grow up in similar habitats, and (3) that occasional dispersal in populations with a different habitat quality is likely to occur. As adults of many species cannot predict their offspring's environment from ambient cues, life-stage specific maternal effects are likely to be common in animals. It will therefore be necessary to incorporate parental ontogeny in the study of parental effects when juveniles and adults inhabit different environments

Theoretical models predict that parents should adjust the amount of care both to their own and their partner's body condition. In most biparental species, parental duties are switched repeatedly allowing for repeated mutual adjustment of the amount of care. In the mouthbrooding cichlid Eretmodus cyanostictus, terms are switched only once with females taking the first share. The timing of the shift of the clutch between mates strongly determines both partners' brooding period and thereby their parental investment. Females signal their readiness to transfer the young several days before the male finally takes them, suggesting sexual conflict over the timing of the shift. In a lab experiment, we reduced the body condition of either the female or the male of a pair to test whether energy reserves affect the timing of the shift and whether female signalling behaviour depends on energetic state. Males with a lowered condition took the young later and incubated for a shorter period, which prolonged the incubation time of their female partners. When female condition was lowered, female and male incubation durations remained unchanged, although females signalled their readiness to shift more intensely. Our results suggest that males adjust their parental investment to own energy reserves but are unresponsive to their mate's condition. Females appear to carry the entire costs for the male's adjustment of care. We propose that intrinsic asymmetries in the scope for mutual adjustment of parental investment and the costs of negotiation crucially influence solutions of the conflict between sexes over care

In many animal taxa, size-selective predation favors fast growth early in life. However, same-aged juveniles can diverge in size due to differences in genotype, environmental conditions, and parental effects and thus may vary in competitive ability. Under food scarcity, competitively inferior juveniles may suffer suppressed growth, whereas under benign conditions, small juveniles may exhibit growth compensation and perform as well as large ones. However, studies testing this while controlling for parental effects are lacking. Here, we hand-raised cichlids, Simochromis pleurospilus, from a wide range of egg sizes and manipulated their size by differential feeding. Afterward, high- and low-ration siblings were kept in groups assigned to either a high- or low-competition environment. We investigated how the degree of competition affected aggressiveness and growth of juveniles with different feeding histories. As predicted, when competition was high, high-ration offspring grew fastest. Interestingly, when competition was weak, low-ration juveniles grew at a similar rate as high-ration ones and many were able to catch up in size. High-ration fish were more aggressive than low-ration ones, and this effect was strongest under high competition. Additionally, in the high-competition environment, received aggression was negatively related to growth, and inflicted aggression correlated positively with the growth of the aggressor. These relationships were absent under low competition. Our findings suggest that the abilities to compensate for early growth depression depend on the prevalent level of competition. Aggression is likely used to monopolize food by juvenile S. pleurospilus; however, when competition is strong, aggression cannot compensate for a size disadvantage

Mothers should adjust the size of propagules to the selective forces to which these offspring will be exposed. Usually, a larger propagule size is favored when young are exposed to high mortality risk or conspecific competition. Here we test 2 predictions on how egg size should vary with these selective agents. When offspring are cared for by parents and/or alloparents, protection may reduce the predation risk to young, which may allow mothers to invest less per single offspring. In the cooperatively breeding cichlid Neolamprologus pulcher, brood care helpers protect group offspring and reduce the latters' mortality rate. Therefore, females are expected to reduce their investment per egg when more helpers are present. In a first experiment, we tested this prediction by manipulating the helper number. In N. pulcher, helpers compete for dispersal opportunities with similar-sized individuals of neighboring groups. If the expected future competition pressure on young is high, females should increase their investment per offspring to give them a head start. In a second experiment, we tested whether females produce larger eggs when perceived neighbor density is high. Females indeed reduced egg size with increasing helper number. However, we did not detect an effect of local density on egg size, although females took longer to produce the next clutch when local density was high. We argue that females can use the energy saved by adjusting egg size to reduced predation risk to enhance future reproductive output. Adaptive adjustment of offspring size to helper number may be an important, as yet unrecognized, strategy of cooperative breeders

The pay-off of deserting and leaving a mate to care for the offspring alone is generally assumed to depend mainly on the availability of alternative mating partners and on the potential spawning rate of males and females. Eretmodus cyanostictus is a monogamous mouthbrooding cichlid in which the clutch is successively incubated first by the female and then by the male. It has been suggested that parents are constrained to monogamy due to low remating probabilities for both sexes. We tested this hypothesis by varying the sex ratio experimentally. Mate desertion by either sex was not significantly higher when additional potential mates were present (males: 8.3%, females: 0%) than when there were no other same-sex conspecifics present (males: 0%, females: 0%). Males lost their mate to a male intruder during their incubation in 26.7% of cases. Pair members were more active and showed more aggression when same-sex conspecifics were present. Behavioural differences between treatments were strongest during the incubation period of a given sex. If no desertion takes place, sexual conflict may be expressed also on a second level, the amount of parental care each parent provides. Indeed, males took the offspring later when additional females were present, although male incubation time did not differ between treatments. A hitherto undescribed display behaviour of females was clear evidence of a conflict about the timing of shift of young. In conclusion, offering alternative mating opportunities did not strongly favour mate desertion in E. cyanostictus. It rather revealed a conflict between mates about when to shift the young