Maze design: size and number of choices impact fish performance in cognitive assays.

Although studies on fish cognition are increasing, consideration of how methodological details influence the ability to detect and measure performance is lagging. Here, in two separate experiments the authors compared latency to leave the start position, latency to make a decision, levels of participation and success rates (whether fish entered the rewarded chamber as first choice) across different physical designs. Experiments compared fish performance across (a) two sizes of T-mazes, large and standard, and a plus-maze, and (b) open choice arenas with either two or four doors. Fish in T-mazes with longer arms took longer to leave the start chamber and were less likely to participate in a trial than fish in T-mazes with shorter arms. The number of options, or complexity, in a maze significantly impacted success but did not necessarily impact behavioural measures, and did not impact the number of fish that reached a chamber. Fish in the plus-maze had similar latencies to leave the start box and time to reach any chamber as fish in the same-sized T-maze but exhibited lower overall success. Similarly, in an open choice arena, increasing the number of options - doors to potential reward chambers - resulted in lower probability of success. There was an influence of reward position in the choice arena, with rewarded chambers closest to the sides of the arena resulting in lower latencies to enter and higher probability of decision success. Together the results allow the authors to offer practical suggestions towards optimal maze design for studies of fish cognition.

Shade as enrichment: testing preferences for shelter in two model fish species.

We compared preferences shown by zebrafish Danio rerio and three-spined stickleback Gasterosteus aculeatus for shelter provided by above-tank shade and artificial plants. Zebrafish showed no preference for either shelter, whereas sticklebacks showed a preference for both shelter types over open areas and for shade over plants. Our results suggest shade may be used as enrichment for captive fish and re-emphasise the importance of species-specific welfare considerations.

The influence of stress hormones and aggression on cooperative behaviour in subordinate meerkats.

In cooperative breeders, aggression from dominant breeders directed at subordinates may raise subordinate stress hormone (glucocorticoid) concentrations. This may benefit dominants by suppressing subordinate reproduction but it is uncertain whether aggression from dominants can elevate subordinate cooperative behaviour, or how resulting changes in subordinate glucocorticoid concentrations affect their cooperative behaviour. We show here that the effects of manipulating glucocorticoid concentrations in wild meerkats (Suricata suricatta) on cooperative behaviour varied between cooperative activities as well as between the sexes. Subordinates of both sexes treated with a glucocorticoid receptor antagonist (mifepristone) exhibited significantly more pup protection behaviour (babysitting) compared to those treated with glucocorticoids (cortisol) or controls. Females treated with mifepristone had a higher probability of exhibiting pup food provisioning (pup-feeding) compared to those treated with cortisol. In males, there were no treatment effects on the probability of pup-feeding, but those treated with cortisol gave a higher proportion of the food they found to pups than those treated with mifepristone. Using 19 years of behavioural data, we also show that dominant females did not increase the frequency with which they directed aggression at subordinates at times when the need for assistance was highest. Our results suggest that it is unlikely that dominant females manipulate the cooperative behaviour of subordinates through the effects of aggression on their glucocorticoid levels and that the function of aggression directed at subordinates is probably to reduce the probability they will breed.

Intergenerational effects of ocean temperature variation: Early life benefits are short-lived in threespine stickleback.

Current climate change models predict an increase in temperature variability and extreme events such as heatwaves, and organisms need to cope with consequent changes to environmental variation. Non-genetic inheritance mechanisms can enable parental generations to prime their offspring's abilities to acclimate to environmental change-but they may also be deleterious. When parents are exposed to predictable environments, intergenerational plasticity can lead to better offspring trait performance in matching environments. Alternatively, parents exposed to variable or unpredictable environments may use plastic bet-hedging strategies to adjust the phenotypic variance among offspring. Here, we used a model species, the threespine stickleback (Gasterosteus aculeatus), to test whether putatively adaptive intergenerational effects can occur in response to shifts in environmental variation as well as to shifts in environmental mean, and whether parents employ plastic bet-hedging strategies in response to increasing environmental variation. We used a full-factorial, split-clutch experiment with parents and offspring exposed to three temperature regimes: constant, natural variation, and increased variation. We show that within-generation exposure to increased temperature variation reduces growth of offspring, but having parents that were exposed to natural temperature variation during gametogenesis may offset some early-life negative growth effects. However, these mitigating intergenerational effects do not appear to persist later in life. We found no indication that stickleback mothers plastically altered offspring phenotypic variance (egg size or clutch size) in response to temperature variation. However, lower inter-individual variance of juvenile fish morphology in offspring of increased variation parents may imply the presence of conservative bet-hedging strategies in natural populations. Overall, in our experiment, parental exposure to temperature variation had limited effects on offspring fitness-related traits. Natural levels of environmental variation promoted a potentially adaptive intergenerational response in early life development, but under more challenging conditions associated with increased environmental variation, the effect was lost.

Understanding 'Non-genetic' Inheritance: Insights from Molecular-Evolutionary Crosstalk.

Understanding the evolutionary and ecological roles of 'non-genetic' inheritance (NGI) is daunting due to the complexity and diversity of epigenetic mechanisms. We draw on insights from molecular and evolutionary biology perspectives to identify three general features of 'non-genetic' inheritance systems: (i) they are functionally interdependent with, rather than separate from, DNA sequence; (ii) precise mechanisms vary phylogenetically and operationally; and (iii) epigenetic elements are probabilistic, interactive regulatory factors and not deterministic 'epialleles' with defined genomic locations and effects. We discuss each of these features and offer recommendations for future empirical and theoretical research that implements a unifying inherited gene regulation (IGR) approach to studies of 'non-genetic' inheritance.

Rank-Related Contrasts in Longevity Arise from Extra-Group Excursions Not Delayed Senescence in a Cooperative Mammal.

In many cooperatively breeding animal societies, breeders outlive non-breeding subordinates, despite investing heavily in reproduction [1-3]. In eusocial insects, the extended lifespans of breeders arise from specialized slowed aging profiles [1], prompting suggestions that reproduction and dominance similarly defer aging in cooperatively breeding vertebrates, too [4-6]. Although lacking the permanent castes of eusocial insects, breeders of vertebrate societies could delay aging via phenotypic plasticity (similar rank-related changes occur in growth, neuroendocrinology, and behavior [7-10]), and such plastic deferment of aging may reveal novel targets for preventing aging-related diseases [11]. Here, we investigate whether breeding dominants exhibit extended longevity and delayed age-related physiological declines in wild cooperatively breeding meerkats. We show that dominants outlive subordinates but exhibit faster telomere attrition (a marker of cellular senescence and hallmark of aging [12]) and that in dominants (but not subordinates), rapid telomere attrition is associated with mortality. Our findings further suggest that, rather than resulting from specialized aging profiles, differences in longevity between dominants and subordinates are driven by subordinate dispersal forays, which become exponentially more frequent with age and increase subordinate mortality. These results highlight the need to critically examine the causes of rank-related longevity contrasts in other cooperatively breeding vertebrates, including social mole-rats, where they are currently attributed to specialized aging profiles in dominants [4].