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Information on fish preference for environmental conditions can be a useful tool to offer them what they want, an important requirement for welfare purposes. Giving that such an approach deals with psychological states of the fish, we investigated whether psychological stress affects fish preference. In this study, we found that the aversive condition of crossing a white open field did not change individual zebrafish's preference for color background or for plant enrichments, despite a great individual variability of preference response. We conclude that individual preferences for environmental conditions are maintained even following a psychological stressor, and highlight that variability in preference options among individuals raises questions about how to best improve welfare of grouped zebrafish.

Animal welfare assessment techniques try to take into consideration the specific needs and wants of the animal in question. Providing enrichment (the addition of physical objects or conspecifics in the housing environment) is often a way to give captive animals the opportunity to choose who or what they interact with and how they spend their time. A fundamental component of the aquatic environment that is often overlooked in captivity, however, is the ability for the animal to choose to engage in physical exercise. For many animals, including fish, exercise is an important aspect of their life history, and is known to have many health benefits, including positive changes in the brain and behavior. Here we present a method for assessing habitat preferences in captive animals. The protocol could easily be adapted to look at a variety of environmental factors (e.g., gravel versus sand as a substrate, plastic plants versus live plants, low flow versus high flow of water) in different aquatic species, or for use with terrestrial species. Statistical assessment of preference is carried out using Jacob's preference index, which ranks the habitats from -1 (avoidance) to +1 (most preferred). With this information, it can be determined what the animal wants from a welfare perspective, including their preferred location.

Populations that have recently diverged offer a powerful model for studying evolution. Ecological differences are expected to generate divergent selection on multiple traits, including neurobiological ones. Animals must detect, process, and act on information from their surroundings and the form of this information can be highly dependent on the environment. We might expect different environments to generate divergent selection not only on the sensory organs, but also on the brain regions responsible for processing sensory information. Here, we test this hypothesis using recently evolved reproductively isolated species pairs of threespine stickleback fish Gasterosteus aculeatus that have well-described differences in many morphological and behavioral traits correlating with ecological differences. We use a state-of-the-art method, magnetic resonance imaging, to get accurate volumetric data for 2 sensory processing regions, the olfactory bulbs and optic tecta. We found a tight correlation between ecology and the size of these brain regions relative to total brain size in 2 lakes with intact species pairs. Limnetic fish, which rely heavily on vision, had relatively larger optic tecta and smaller olfactory bulbs compared with benthic fish, which utilize olfaction to a greater extent. Benthic fish also had larger total brain volumes relative to their body size compared with limnetic fish. These differences were erased in a collapsed species pair in Enos Lake where anthropogenic disturbance has led to intense hybridization. Together these data indicate that evolution of sensory processing regions can occur rapidly and independently.

While differences in individual personality are common in animal populations, understanding the ecological significance of variation has not yet been resolved. Evidence suggests that personality may influence learning and memory; a finding that could improve our understanding of the evolutionary processes that produce and maintain intraspecific behavioural heterogeneity. Here, we tested whether boldness, the most studied personality trait in fish, could predict learning ability in brook trout. After quantifying boldness, fish were trained to find a hidden food patch in a maze environment. Stable landmark cues were provided to indicate the location of food and, at the conclusion of training, cues were rearranged to test for learning. There was a negative relationship between boldness and learning as shy fish were increasingly more successful at navigating the maze and locating food during training trials compared to bold fish. In the altered testing environment, only shy fish continued using cues to search for food. Overall, the learning rate of bold fish was found to be lower than that of shy fish for several metrics suggesting that personality could have widespread effects on behaviour. Because learning can increase plasticity to environmental change, these results have significant implications for fish conservation., (Copyright © 2016 Elsevier B.V. All rights reserved.)

To test whether swimming skills can be improved by exposure to structurally complex environments, juvenile rainbow trout Oncorhynchus mykiss were reared in either physically enriched or plain tanks for 2 months and were then screened to test their ability to swim along a channel while avoiding obstructions. The results show that even a relatively short exposure period to enrichment improves O. mykiss swimming agility., (© 2016 The Fisheries Society of the British Isles.)

The ways in which challenging environments during development shape the brain and behaviour are increasingly being addressed. To date, studies typically consider only single variables, but the real world is more complex. Many factors simultaneously affect the brain and behaviour, and whether these work independently or interact remains untested. To address this, zebrafish (Danio rerio) were reared in a two-by-two design in housing that varied in structural complexity and/or exposure to a stressor. Fish experiencing both complexity (enrichment objects changed over time) and mild stress (daily net chasing) exhibited enhanced learning and were less anxious when tested as juveniles (between 77 and 90 days). Adults tested (aged 1 year) were also less anxious even though fish were kept in standard housing after three months of age (i.e. no chasing or enrichment). Volumetric measures of the brain using magnetic resonance imaging (MRI) showed that complexity alone generated fish with a larger brain, but this increase in size was not seen in fish that experienced both complexity and chasing, or chasing alone. The results highlight the importance of looking at multiple variables simultaneously, and reveal differential effects of complexity and stressful experiences during development of the brain and behaviour., (© 2016 The Authors.)

HIGHLIGHTS This study tested the effects of adolescent-stress on adult learning and memory.Adolescent-stressed rats had enhanced reversal learning compared to unstressed rats.Adolescent-stress exposure made working memory more vulnerable to disturbance.Adolescent-stress did not affect adult associative learning or reference memory. Exposure to acute stress can cause a myriad of cognitive impairments, but whether negative experiences continue to hinder individual as they age is not as well understood. We determined how chronic unpredictable stress during adolescence affects multiple learning and memory processes in adulthood. Using male Sprague Dawley rats, we measured learning (both associative and reversal) and memory (both reference and working) starting 110 days after completion of an adolescent-stress treatment. We found that adolescent-stress affected adult cognitive abilities in a context-dependent way. Compared to rats reared without stress, adolescent-stressed rats exhibited enhanced reversal learning, an indicator of behavioral flexibility, but showed no change in associative learning and reference memory abilities. Working memory, which in humans is thought to underpin reasoning, mathematical skills, and reading comprehension, may be enhanced by exposure to adolescent-stress. However, when adolescent-stressed animals were tested after a novel disturbance, they exhibited a 5-fold decrease in working memory performance while unstressed rats continued to exhibit a linear learning curve. These results emphasize the capacity for stress during adolescence to transform the cognitive abilities of adult animals, even after stress exposure has ceased and animals have resided in safe environments for the majority of their lifespans.

Spatial abilities allow animals to retain and cognitively manipulate information about their spatial environment and are dependent upon neural structures that mature during adolescence. Exposure to stress in adolescence is thought to disrupt neural maturation, possibly compromising cognitive processes later in life. We examined whether exposure to chronic unpredictable stress in adolescence affects spatial ability in late adulthood. We evaluated spatial learning, reference and working memory, as well as long-term retention of visuospatial cues using a radial arm water maze. We found that stress in adolescence decreased the rate of improvement in spatial learning in adulthood. However, we found no overall performance impairments in adult reference memory, working memory, or retention caused by adolescent-stress. Together, these findings suggest that adolescent-stress may alter the strategy used to solve spatial challenges, resulting in performance that is more consistent but is not refined by incorporating available spatial information. Interestingly, we also found that adolescent-stressed rats showed a shorter latency to begin the water maze task when re-exposed to the maze after an overnight delay compared with control rats. This suggests that adolescent exposure to reoccurring stressors may prepare animals for subsequent reoccurring challenges. Overall, our results show that stress in adolescence does not affect all cognitive processes, but may affect cognition in a context-dependent manner.

Recently we have shown that adult rats exposed to chronic stress during adolescence increase foraging performance in high-threat conditions by 43% compared to rats reared without stress. Our findings suggest that stress during adolescence can prepare rats to better function under future threat, which supports hypotheses describing an adaptive role for the long-term consequences of early stress (e.g. the thrifty phenotype and maternal mismatch hypotheses). These hypotheses often predict that early stress will impair performance in low-threat conditions later in life. However, we did not find any difference in performance under low-threat conditions between adolescent-stressed and unstressed adult animals. To understand why stress during adolescence may affect performance in high-threat but not in low-threat conditions, we discuss our findings in the framework of the Yerkes-Dodson law, a key precept of psychology that has been used for over a century to describe how stress affects performance.

The capacity for specialization and radiation make fish an excellent group in which to investigate the depth and variety of animal cognition. Even though early observations of fish using tools predates the discovery of tool use in chimpanzees, fish cognition has historically been somewhat overlooked. However, a recent surge of interest is now providing a wealth of material on which to draw examples, and this has required a selective approach to choosing the research described below. Our goal is to illustrate the necessity for basing cognitive investigations on the ecological and evolutionary context of the species at hand. We also seek to illustrate the importance of ecology and the environment in honing a range of sensory systems that allow fish to glean information and support informed decision-making. The various environments and challenges with which fish interact require equally varied cognitive skills, and the solutions that fish have developed are truly impressive. Similarly, we illustrate how common ecological problems will frequently produce common cognitive solutions. Below, we focus on four topics: spatial learning and memory, avoiding predators and catching prey, communication, and innovation. These are used to illustrate how both simple and sophisticated cognitive processes underpin much of the adaptive behavioral flexibility exhibited throughout fish phylogeny. Never before has the field had such a wide array of interdisciplinary techniques available to access both cognitive and mechanistic processes underpinning fish behavior. This capacity comes at a critical time to predict and manage fish populations in an era of unprecedented global change., (© 2015 John Wiley & Sons, Ltd.)

Exposure to stress during adolescence can cause long-term changes in behavior and cognition. Anxiety diagnoses rise during adolescence and are increased by adverse experiences. Currently, it is unknown how long stress during adolescence alters anxiety in adulthood. We found that rats exposed to chronic unpredictable stress during adolescence expressed altered behavior 6.5 months later; showing increased anxiety in a feeding test in a novel environment. Although behavioral changes indicative of anxiety were detected in late adulthood, the basal levels of fecal corticoid metabolites in prior-stressed rats did not differ from unstressed, control rats., (Copyright © 2014 Elsevier B.V. All rights reserved.)

An assessment of consciousness in nonverbal animals requires a framework for research that extends testing methods beyond subjective report. This chapter proposes a working definition of consciousness in terms of temporal representation that provides the critical link between internal phenomenology and external behavior and neural structure. Our claim is that consciousness represents the present moment as distinct from the past and the future in order to flexibly respond to stimuli. We discuss behavioral and neural evidence that indicates the capacity for both flexible response and temporal representation, and we illustrate these capacities in fish, a taxonomic group that challenges human intuitions about consciousness.

Fish exhibit diverse cognitive capacities: they cooperate, punish, develop cultural traditions, learn to map their environment and communicate their intentions to one another. Skills such as these have helped fish radiate to colonize the many and diverse aquatic niches available. Prey fish are no exception to this, and several recent studies have shown them to be a rich resource for understanding the evolutionary ecology of animal cognition. Many fish have to cope with the threat of predation, but some environments contain more predators than others. These environments deliver the opportunity to investigate how predation pressure shapes fish cognition and behaviour. Here we compared fish from two high and two low predation habitats in their ability to learn a sequential choice spatial task. We also investigated their ability to solve the maze after it was rearranged. Fish from high predation sites made more errors as they learned to navigate the maze than fish from low predation sites. The fish also varied in the cues that they learned to help them solve the task. These did not vary by levels of predation pressure, rather, they differed between rivers, with fish from one river learning to use landmark cues, and those from the other river learning the sequence of left and right turns. As the different populations varied in how well they learned to navigate through a reconfigured maze, it seems likely that predation pressure is not the only factor influencing spatial behavior in these fish., (Copyright © 2014 Elsevier B.V. All rights reserved.)

Living in challenging environments can influence the behavior of animals in a number of ways. For instance, populations of prey fish that experience frequent, nonlethal interactions with predators have a high proportion of individuals that express greater reaction to risk and increased activity and exploration-collectively known as temperament traits. Temperament traits are often correlated, such that individuals that are risk-prone also tend to be active and explore more. Spatial learning, which requires the integration of many sensory cues, has also been shown to vary in fish exposed to different levels of predation threat. Fish from areas of low predation risk learn to solve spatial tasks faster than fish from high predation areas. However, it is not yet known whether simpler forms of learning, such as learning associations between two events, are similarly influenced. Simple forms of associative learning are likely to be affected by temperament because a willingness to approach and explore novel situations could provide animals with a learning advantage. However, it is possible that routine-forming and inflexible traits associated with risk-prone and increased exploratory behavior may act in the opposite way and make risk-prone individuals poorer at learning associations. To investigate this, we measured temperament in Panamanian bishop fish (Brachyrhaphis episcopi) sampled from a site known to contain many predators. The B. episcopi were then tested with an associative learning task. Within this population, fish that explored more were faster at learning a cue that predicted access to food, indicating a link between temperament and basic learning abilities.

This study describes how three individual fish, Atlantic cod (Gadus morhua L.), developed a novel behaviour and learnt to use a dorsally attached external tag to activate a self-feeder. This behaviour was repeated up to several hundred times, and over time these fish fine-tuned the behaviour and made a series of goal-directed coordinated movements needed to attach the feeder's pull string to the tag and stretch the string until the feeder was activated. These observations demonstrate a capacity in cod to develop a novel behaviour utilizing an attached tag as a tool to achieve a goal. This may be seen as one of the very few observed examples of innovation and tool use in fish.

Farming fish for human consumption continues to expand as an industry and, with this increasing interaction with captive fish populations, there is now a growing interest in determining how to create good welfare for the fish we farm. This article summarises recent advances in our understanding of pain and stress responses in fish and how these relate to farmed fish welfare. Over the last decade several studies have examined whether or not fish feel pain, how aversive the experience is, and how such experiences may be mitigated through the use of analgesics. The basic neural mechanisms that enable the detection of tissue damage, i.e. nociceptive mechanisms, appear to be broadly conserved from fish through to birds and mammals, however, there is debate about the extent of the negative feelings associated with pain and whether these are truly experienced by fish. The stress response that helps fish to cope with various challenges also appears to be largely conserved across vertebrates, and the physiological changes that occur in response to acute and chronic stress in fish are similar to those described for mammals. Therefore, fish appear to have the innate ability to experience negative states such as pain and stress in a way analogous to that experienced by other vertebrates. There are multiple situations in which farmed fish may experience pain and stress and there is now a growing recognition that, to deliver appropriate welfare, on-farm practices and procedures will have to change. It is also the case that the welfare requirements of the different fish species that we farm vary, with some species coping better in captive rearing environments than others. The topic of fish welfare is relatively new and more research on stress responses, allostasis, pain thresholds and analgesics is required to help promote good fish welfare.

Aluminum (Al) toxicity occurs frequently in natural aquatic ecosystems as a result of acid deposition and natural weathering processes. Detrimental effects of Al toxicity on aquatic organisms are well known and can have consequences for survival. Fish exposed to Al in low pH waters will experience physiological and neuroendocrine changes that disrupt homeostasis and alter behavior. To investigate the effects of Al exposure on both the brain and behavior, Atlantic salmon (Salmo salar) kept in water treated with Al (pH 5.7, 0.37±0.04 μmol 1(-1) Al) for 2 weeks were compared with fish kept in under control conditions (pH 6.7, <0.04 μmol 1(-1) Al). Fish exposed to Al and acidic conditions had increased Al accumulation in the gills and decreased gill Na(+), K(+)-ATPase activity, which impaired osmoregulatory capacity and caused physiological stress, indicated by elevated plasma cortisol and glucose levels. Here we show for the first time that exposure to Al in acidic conditions also impaired learning performance in a maze task. Al toxicity also reduced the expression of NeuroD1 transcript levels in the forebrain of exposed fish. As in mammals, these data show that exposure to chronic stress, such as acidified Al, can reduce neural plasticity during behavioral challenges in salmon, and may impair the ability to cope with new environments.

Different kinds of experience during early life can play a significant role in the development of an animal's behavioural phenotype. In natural contexts, this influences behaviours from anti-predator responses to navigation abilities. By contrast, for animals reared in captive environments, the homogeneous nature of their experience tends to reduce behavioural flexibility. Studies with cage-reared rodents indicate that captivity often compromises neural development and neural plasticity. Such neural and behavioural deficits can be problematic if captive-bred animals are being reared with the intention of releasing them as part of a conservation strategy. Over the last decade, there has been growing interest in the use of environmental enrichment to promote behavioural flexibility in animals that are bred for release. Here, we describe the positive effects of environmental enrichment on neural plasticity and cognition in juvenile Atlantic salmon (Salmo salar). Exposing fish to enriched conditions upregulated the forebrain expression of NeuroD1 mRNA and improved learning ability assessed in a spatial task. The addition of enrichment to the captive environment thus promotes neural and behavioural changes that are likely to promote behavioural flexibility and improve post-release survival.

Animals that experience adverse events in early life often have life-long changes to their physiology and behavior. Long-term effects of stress during early life have been studied extensively, but less attention has been given to the consequences of negative experiences solely during the adolescent phase. Adolescence is a particularly sensitive period of life when regulation of the glucocorticoid "stress" hormone response matures and specific regions in the brain undergo considerable change. Aversive experiences during this time might, therefore, be expected to generate long-term consequences for the adult phenotype. Here we investigated the long-term effects of exposure to chronic unpredictable stress during adolescence on adult decision-making, coping response, cognitive bias, and exploratory behavior in rats. Rats exposed to chronic unpredictable stress (e.g., isolation, crowding, cage tilt) were compared to control animals that were maintained in standard, predictable conditions throughout development. Unpredictable stress during adolescence resulted in a suite of long-term behavioral and cognitive changes including a negative cognitive bias [F (1, 12) = 5.000, P < 0.05], altered coping response [T (1, 14) = 2.216, P = 0.04], and accelerated decision-making [T (1, 14) = 3.245, P = 0.01]. Exposure to chronic stress during adolescence also caused a short-term increase in boldness behaviors; in a novel object test 15 days after the last stressor, animals exposed to chronic unpredictable stress had decreased latencies to leave a familiar shelter and approach a novel object [T (1, 14) = 2.240, P = 0.04; T (1, 14) = 2.419, P = 0.03, respectively]. The results showed that stress during adolescence has long-term impacts on behavior and cognition that affect the interpretation of ambiguous stimuli, behavioral response to adverse events, and how animals make decisions.

Most fishes experiencing challenging environments are able to adjust and adapt their physiology and behaviour to help them cope more effectively. Much of this flexibility is supported and influenced by cognition and neural plasticity. The understanding of fish cognition and the role played by different regions of the brain has improved significantly in recent years. Techniques such as lesioning, tract tracing and quantifying changes in gene expression help in mapping specialized brain areas. It is now recognized that the fish brain remains plastic throughout a fish's life and that it continues to be sensitive to environmental challenges. The early development of fish brains is shaped by experiences with the environment and this can promote positive and negative effects on both neural plasticity and cognitive ability. This review focuses on what is known about the interactions between the environment, the telencephalon and cognition. Examples are used from a diverse array of fish species, but there could be a lot to be gained by focusing research on neural plasticity and cognition in fishes for which there is already a wealth of knowledge relating to their physiology, behaviour and natural history, e.g. the Salmonidae., (© 2012 The Authors. Journal of Fish Biology © 2012 The Fisheries Society of the British Isles.)

In this study we evaluated Pavlovian conditioned food anticipatory behaviour as a potential indicator for stress in groups of Atlantic salmon, and compared this with the physiological stress responses of cortisol excretion into water and hyper-consumption of oxygen. We hypothesised that environmental stress would result in reduced feeding motivation. To assess this, we measured the strength of anticipatory behaviour during a period of flashing light that signalled arrival of food. Further, we expected that fish given a reduced food ration would be less sensitive to environmental stress than fish fed full ration. The fish responded to an acute temperature fluctuation with hyper-consumption of oxygen that decreased in line with the temperature, and elevated cortisol excretion up to 1h after the stressor. These physiological responses did not differ significantly between the food ration groups. The anticipatory behaviour was significantly reduced after the stressor and returned to control levels after 1 to 2 h in the reduced ration group, but not until after 3 to 4 h in the full ration group. Our results show that acute stress can be measured in terms of changes to feeding motivation, and that it is a more sensitive indicator of stress that influences the fish over a longer time period than measures of change in cortisol excretion., (Copyright © 2011 Elsevier Inc. All rights reserved.)

Mature female Atlantic salmon Salmo salar were given intraperitoneal cortisol implants 1 week prior to stripping to examine the influence of simulated maternal stress on offspring boldness and social dominance. Behavioural tests originally designed to investigate stress responsiveness and coping styles in salmonids (i.e. feeding in isolation, dominance tests and acute confinement) were carried out on the offspring 1·5 years after hatching. In the feeding test, there were no differences between the two treatment groups in total feeding score or number of pellets eaten, but offspring from the cortisol-implanted females made more unsuccessful feeding attempts than offspring from control females. In dominance tests, there was no difference between controls and cortisol-treated fish regarding propensity to become socially dominant. A higher proportion of individuals with bite marks, however, was observed in the cortisol group when compared to controls. Cortisol-treated offspring that gained dominant rank in the dominance tests performed more aggressive acts after stable dominance-subordinate relationships were established compared to control winners. During acute confinement stress, offspring from cortisol-implanted females showed a reduction in the proportion of time they were moving compared to the controls. These results indicate that the maternal endocrine state at spawning affects several aspects of progeny behaviour potentially related to subsequent success and survival in farmed S. salar., (© 2011 The Authors. Journal of Fish Biology © 2011 The Fisheries Society of the British Isles.)

Two temperament traits, tendency to explore and activity level, were measured in a tropical poeciliid fish, the Panamanian bishop Brachyrhaphis episcopi. Open-field arena tests were used to quantify how predation pressure shapes activity levels and exploratory behaviours. Fish behaviour differed between high and low-predation populations. Fish that experienced higher levels of predation were both more explorative and more active. There were also some individual differences within populations; fish varied in their levels of exploration and activity in a novel open arena, but these differences were not related to sex or size. Together with previous studies on this species, these results indicate that there is a behavioural syndrome associated with predation pressure. Fish from high-predation populations are bolder, more explorative and more active than those from low-predation populations., (© 2011 The Authors. Journal of Fish Biology © 2011 The Fisheries Society of the British Isles.)

Aggression is often positively correlated with other behavioural traits such as boldness and activity levels. Comparisons across populations can help to determine factors that promote the evolution of such traits. We quantified these behaviours by testing the responses of wild-caught poeciliid fish, Brachyrhaphis episcopi, to mirror image stimuli. This species occurs in populations that experience either high or low levels of predation pressure. Previous studies have shown that B. episcopi from low predation environments are less bold than those that occur with many predators. We therefore predicted that fish from high predation populations would be more aggressive and more active than fish from low predation populations. However, we found the opposite - low predation fish approached a mirror and a novel object more frequently than high predation fish suggesting that 'boldness' and aggression were higher in low predation populations, and that population-level boldness measures may vary depending on context. When tested individually, low predation fish inspected their mirror image more frequently. Females, but not males, from low predation sites were also more aggressive towards their mirror image. Variation in female aggression may be driven by a trade-off between food availability and predation risk. This suggests that the relationship between aggression and boldness has been shaped by adaptation to environmental conditions, and not genetic constraints., (Copyright © 2010 Elsevier B.V. All rights reserved.)

Since the collapse of the pelagic fisheries off southwest Africa in the late 1960s, jellyfish biomass has increased and the structure of the Benguelan fish community has shifted, making the bearded goby (Sufflogobius bibarbatus) the new predominant prey species. Despite increased predation pressure and a harsh environment, the gobies are thriving. Here we show that physiological adaptations and antipredator and foraging behaviors underpin the success of these fish. In particular, body-tissue isotope signatures reveal that gobies consume jellyfish and sulphidic diatomaceous mud, transferring "dead-end" resources back into the food chain.

To determine whether enhancing the survival of new recruits is a sensible target for the restorative management of depleted coral-reef fish populations, settlement-stage ambon damsel fish Pomacentrus amboinensis were captured, tagged and then either released immediately onto small artificial reefs or held in aquaria for 1 week prior to release. Holding conditions were varied to determine whether they affected survival of fish: half the fish were held in bare tanks (non-enriched) and the other half in tanks containing coral and sand (enriched). Holding fish for this short period had a significantly positive effect on survivorship relative to the settlement-stage treatment group that were released immediately. The enrichment of holding conditions made no appreciable difference on the survival of fish once released onto the reef. It did, however, have a positive effect on the survival of fish while in captivity, thus supporting the case for the provision of simple environmental enrichment in fish husbandry. Collecting and holding settlement-stage fish for at least a week before release appear to increase the short-term survival of released fish; whether it is an effective method for longer-term enhancement of locally depleted coral-reef fish populations will require further study.

1. There is growing interest in the causes and consequences of animal temperaments. Temperament behaviours often have heritable components, but ecological variables can also affect them. Numerous variables are likely to differ between habitats, and these may interact to influence temperament behaviours. 2. Temperament behaviours may be correlated within populations (behavioural syndromes), although the underlying causes of such correlations are currently unclear. 3. We analysed three different temperament behaviours and learning ability in three-spined sticklebacks, Gasterosteus aculeatus, to determine how different ecological variables influence them both within and between populations. We selected populations from four ponds and four rivers that varied naturally in their exposure to predators. 4. High-predation river populations were significantly less bold than a high-predation pond and low-predation river populations, and low-predation pond populations were significantly less bold than a high-predation pond population. Within populations, temperament behaviours were correlated in one high-predation river population only. 5. These results suggest that multiple ecological factors can interact to affect temperament behaviours between populations, and also correlations in those behaviours within populations.

In many estuarine sites, morphological and genetic differences between anadromous and freshwater threespine sticklebacks are maintained despite breeding in sympatry. Here, we investigate the maintenance of this morphological divergence in a natural hybrid zone in the River Tyne, Scotland. We provide a morphological description of the hybrid zone, and using a Bayesian MCMC approach, identified distinct anadromous and freshwater genetic clusters. Anadromous and freshwater sticklebacks breed in spatial and temporal sympatry in the lower reaches of the River Tyne. The frequency of hybrids within these sites (33%) indicates prezygotic isolation is not complete, and suggests that assortative mating is not strong. However, significant heterozygote deficit and cytonuclear disequilibrium in juveniles collected from sympatric sites confirms that barriers to gene flow exist between the morphs in the wild. In addition, we found no evidence of a directional bias in hybridisation, although hybrids with anadromous mothers were more common because anadromous females outnumbered freshwater females within the hybrid zone. We discuss the potential contribution of temporal, spatial, and sexual prezygotic barriers to the observed reproductive isolation as well as postzygotic selection against hybrid zygotes or fry.

We subjected fish from regions of high and low levels of predation pressure in four independent streams to a mild stressor and recorded their opercular beat rates. Fish from low-predation areas showed higher maximum, minimum and mean opercular beat frequencies than fish from high-predation regions. The change in opercular beat frequency (scope) was also significantly greater in fish from low- than in fish from high-predation regions. Under normal activity levels, however, low predation fish showed a reduced opercular beat frequency, which may be indicative of reduced activity levels or metabolic rate. Opercular beat frequency was negatively correlated with standard length as one would expect based on higher metabolic rates in smaller fish. We suggest that these contrasting stress responses are most likely the result of differential exposure to predators in fish from high- and low-predation areas. We argue that reduced stress responses in high-predation areas evolved to prevent excessive energy expenditure by modulating the fright response.

The release of hatchery-reared fishes for restoring threatened and endangered populations is one of the most controversial issues in applied ecology. A central issue has been to determine whether releases cause extinction of local wild populations. This may arise either through domesticated or non-local fishes hybridizing with wild fishes, or through inappropriate behavioural interactions; for example, many hatchery fishes show exaggerated aggressive and competitive behaviour and out-compete wild counterparts. The impact of the impoverished hatchery environment in shaping behaviour is only now receiving attention. Attempts to counteract hatchery-related behavioural deficiencies have utilized intensive training programmes shortly before the fishes are released. However, we show here that simple exposure to variable spatial and foraging cues in the standard hatchery environment generates fishes with enhanced behavioural traits that are probably associated with improved survival in the wild. It appears that fishes need to experience a varying and changeable environment to learn and develop flexible behaviour. Using variable hatchery rearing environments to generate suitable phenotypes in combination with a knowledge of appropriate local genotypes, rehabilitation of wild fishes is likely to succeed, where to date it has largely failed.

Female three-spined sticklebacks have been found to use visual cues when responding sexually towards courting males, often preferring more intensely red-coloured males, and males with blue rather than silver irises. However, traditionally the literature has failed to test preference across the full spectral range to which females might be sensitive, limiting analysis to the human-visible wavelengths of the electromagnetic spectrum. We studied the effects that the addition of ultraviolet wavelengths has on the mate-choice preferences of female sticklebacks using a two-choice paradigm. We found that females preferred males that were viewed across the full spectrum to males whose display lacked an ultraviolet component. Using suitable controls we were able to establish that female preference was sexually motivated and was not caused by a general preference for the manipulated light conditions. Our results indicate that female preference may be due to an enhancement in visual contrast when males are viewed in full spectrum conditions.

Differential use of each hemisphere of the brain for specific tasks is a widespread phenomenon that appears to have arisen in the early history of tetrapod lineage. Despite a high degree of conformity in the development of lateralization among the tetrapods, some variation exists. The mechanisms underlying this variation remain largely unresolved. We exposed fish from regions of high and low predation pressure to a series of visual experiences, including viewing an empty compartment, a novel object and a live predator. Fish from each region differed in their preferential use of each eye to view the scenes. For example, fish from high predation regions viewed a live predator by using their right eye, whereas fish from low predation sites showed no eye preference. These results suggest that the degree of lateralization varies between populations of the same species that have been exposed to different ecological/evolutionary pressures.

This study aimed to assess fear responses to a novel object while experiencing a noxious event to determine whether nociception or fear will dominate attention in a fish in novel object testing paradigm. This experimentally tractable animal model was used to investigate (1) the degree of neophobia to a novel object while experiencing noxious stimulation, (2) the response of the fish after removing the fear-causing event by using a familiar object, and (3) the effects of removing the nociceptive response by morphine administration and examining the response to a novel object. Control animals displayed a classic fear response to the novel objects and spent most of their time moving away from this stimulus, as well as showing an increase in respiration rate when the novel object was presented. In contrast, noxiously stimulated animals spent most of their time in close proximity to the novel object and showed no additional increase in respiration rate to novel object presentation. There was evidence of a slight hypoalgesia in noxiously stimulated animals. The responses to familiar objects demonstrated that by familiarizing the animal with the object, fear was removed from the experiment. Both control and noxiously treated animals responded in similar ways to a novel object by spending the majority of their time in close proximity. Treatment with morphine reduced effects of noxious stimulation and appears to be an effective analgesic. After morphine administration, the acid-injected animals showed a neophobic response to a novel object and this was similar to the response of the control fish, with a similar amount of time spent moving away from the object and an increase in ventilation in response to the novel object. Morphine affected the fear response because both groups approached the novel object more quickly than the non-morphine controls. These results suggest that nociception captures the animal's attention with only a relatively small amount of attention directed at responding to the fear of the novel object.

Nociception is the detection of a noxious tissue-damaging stimulus and is sometimes accompanied by a reflex response such as withdrawal. Pain perception, as distinct from nociception, has been demonstrated in birds and mammals but has not been systematically studied in lower vertebrates. We assessed whether a fish possessed cutaneous nociceptors capable of detecting noxious stimuli and whether its behaviour was sufficiently adversely affected by the administration of a noxious stimulus. Electrophysiological recordings from trigeminal nerves identified polymodal nociceptors on the head of the trout with physiological properties similar to those described in higher vertebrates. These receptors responded to mechanical pressure, temperatures in the noxious range (more than 40 degrees C) and 1% acetic acid, a noxious substance. In higher vertebrates nociceptive nerves are either A-delta or C fibres with C fibres being the predominating fibre type. However, in the rainbow trout A-delta fibres were most common, and this offers insights into the evolution of nociceptive systems. Administration of noxious substances to the lips of the trout affected both the physiology and the behaviour of the animal and resulted in a significant increase in opercular beat rate and the time taken to resume feeding, as well as anomalous behaviours. This study provides significant evidence of nociception in teleost fishes and furthermore demonstrates that behaviour and physiology are affected over a prolonged period of time, suggesting discomfort.

It is widely believed that male mammals have better spatial ability than females. A large number of evolutionary hypotheses have been proposed to explain these differences, but few species have been tested. The authors critically review the proposed evolutionary explanations for sex differences in spatial cognition and conclude that most of the hypotheses are either logically flawed or, as yet, have no substantial support. Few of the data exclusively support or exclude any current hypotheses. The hypothesis with the strongest support proposes that range size was the selection pressure that acted to increase spatial ability. The authors suggest ways in which these hypotheses could be tested by presenting explicit predictions and suggesting suitable test species or conditions.

Among the theories that have been proposed to explain the evolution of altruism are direct reciprocity and indirect reciprocity. The idea of the latter is that helping someone or refusing to do so has an impact on one's reputation within a group. This reputation is constantly assessed and reassessed by others and is taken into account by them in future social interactions. Generosity in indirect reciprocity can evolve if and only if it eventually leads to a net benefit in the long term. Here, we show that this key assumption is met. We let 114 students play for money in an indirect and a subsequent direct reciprocity game. We found that although being generous, i.e., giving something of value to others, had the obvious short-term costs, it paid in the long run because it builds up a reputation that is rewarded by third parties (who thereby themselves increase their reputation). A reputation of being generous also provided an advantage in the subsequent direct reciprocity game, probably because it builds up trust that can lead to more stable cooperation.

'Good genes' models of sexual selection suggest that elaborate male sexual ornaments have evolved as reliable signals of male quality because only males of high genetic viability are able to develop and maintain them. Females benefit from choosing such individuals if quality is heritable. A key prediction is that the offspring of males with elaborate mating displays will perform better than those of less elaborate males, but it has proved difficult to demonstrate such an effect independently of the effects of differences in parental investment. We tested for 'good genes' linked to male ornamentation in the three-spined stickleback Gasterosteus aculeatus using in vitro fertilization to generate maternal half-siblings, which were raised without parental care. Maternal half-siblings sired by brightly coloured males grew less quickly than half-siblings sired by dull males but were more resistant to a controlled disease challenge. Among the offspring that became infected, those with brighter fathers had higher white blood cell counts. This suggests that highly ornamented males confer disease resistance on their offspring. The association with reduced growth suggests a mechanism for the maintenance of heritable variation in both disease resistance and male sexual coloration.

In a number of mammalian species, males appear to have superior spatial abilities to females. The favoured explanations for this cognitive difference are hormonal, with higher testosterone levels in males than females leading to better spatial performance, and evolutionary, where sexual selection has favoured males with increased spatial abilities for either better navigational skills in hunting or to enable an increased territory size. However, an alternative explanation for this sex difference focuses on the role of varying levels of oestrogen in females in spatial cognition (the 'fertility and parental care' hypothesis). One possibility is that varying oestrogen levels result in variation in spatial learning and memory so that, when tested across the oestrous cycle, females perform as well as males on days of low oestrogen but more poorly on days of high oestrogen. If day in the oestrous cycle is not taken into account then, across an experiment, any sex differences found would always produce male superiority. We used a spatial working memory task in a Morris water maze to test the spatial learning and memory abilities of male and female rats. The rats were tested across a number of consecutive days during which the females went through four oestrous cycles. We found no overall sex differences in latencies to reach a submerged platform in a Morris water maze but, on the day of oestrus (low oestrogen), females took an extra swim to learn the platform's location (a 100% increase over the other days in the cycle). Female swim speed also varied across the oestrous cycle but females were no less active on the day of oestrus. These results oppose the predictions of the fertility and parental care hypothesis.

Recent work has shown that mice with subclinical parasitic infections suffer impaired spatial learning and memory, as assayed in an open-field water maze. Although the mechanism underlying this effect is not clear, the phenomenon has been reported following infection with both a protozoan parasite (Eimeria vermiformis) and a gastrointestinal nematode (Heligmosomoides polygyrus). In a variety of experiments, we examined the effects of a different gastrointestinal nematode, Strongyloides ratti, on the ability of rats and mice to learn a spatial or a discrimination task. Animals were tested at various stages post-infection, with different levels of infection, using different lines of S. ratti and with varying experimental protocols. All animals learned the tasks, but we found no evidence of an effect of S. ratti infection on learning or memory. Even rats infected with approximately 5000 S. ratti larvae, a dose which has an impact on rat body size, showed no deficit in learning ability. Various reasons for the conflict between our results and those previously reported for E. vermiformis and H. polygyrus are discussed. Our results show that impaired learning and memory following parasitic infection is not a ubiquitous or at least easily replicated phenomenon.

Operant studies on pigeons using slide-projected images suggest that photographs of geographical locations might be used as a research tool to study the importance of visual landmarks in homing. Before using this method, however, it is necessary to show that pigeons do see photographic slides as representing real world locations. After reviewing the evidence for picture-to-object correspondence for geographical locations in pigeons, we report the results of an experiment designed to test whether outdoor experience at a location affected homing pigeons' ability to categorise slides of that versus another location displayed in an operant set-up. Four birds visited one location immediately before each experimental session; four birds visited an irrelevant location. No effect of outdoor experience was found on acquisition, or transfer to novel stimuli. The possible reasons for limitations on picture-to-object correspondence are discussed.

Predictors of the attitudinal measure of orientation toward help-seeking for emotional problems have been shown to include demographic, network, and personality variables. This research determined whether these same variables predict the behavioral outcome measure of help-seeking, both in general and from professional services in particular. Help-seeking in response to emotional problems was studied in a sample of Australian adolescents. General help-seeking was predicted by more symptoms of psychological distress, being female, availability of social support, knowing someone who had sought professional help, and the personality characteristics of high private self-consciousness and willingness to disclose mental health. When only those with evident emotional distress were considered, only gender and willingness to disclose remained significant predictors. These same variables did not account for those who sought professional help rather than relying upon their informal network. Level of psychological distress was the only significant predictor of professional consultation. Psychological symptoms and gender were shown to be more relevant predictors of the behavioral measure of help-seeking than network or personality characteristics.

This paper focuses on the one third of retirees who report difficulty in making the transition from work to retirement. On the basis of Levy's [16] research, it is hypothesized that there are different styles of poor adjustment that have different consequences for the retirees experiencing them. Data from 487 male and female retirees identify four poor adjustment styles--poor health, negativism, change adaptation and retirement reluctance. While the latter two responses to retirement tend to be problems only in the short term, the former two may have far reaching consequences. Health related retirement and negativism about retiring were linked with low activity and involvement, poor physical and mental health, inadequate income and low life satisfaction in the years following retirement.

This paper outlines and evaluates a newly developed measure of neuroticism, the Scale of Emotional Arousability (SEA). The 15-item scale is economical to use, avoids the consistent pattern of negatively worded items that plagues the neuroticism (N) Scale of the Eysenck Personality Inventory (EPI) and comprises of items that focus specifically on emotional arousability, thereby bringing the measurement of neuroticism in line with Eysenck's (1967) original conceptualization. The SEA shows every indication of being psychometrically sound and correlates with related constructs in much the same way as the N Scale of the EPI. The SEA is recommended as an alternative to the EPI for research purposes, particularly where there is a need to measure independently such related constructs as neurosis, self-esteem and private self-consciousness.