Social motivation and conflict resolution tactics as potential building blocks of sociality in cichlid fishes

Even closely related and ecologically similar cichlid species of Lake Tanganyika exhibit an impressive diversity of social systems, and therefore these fishes offer an excellent opportunity to examine the evolution of social behaviour. Sophisticated social relationships are thought to have evolved via a building block design where more fundamental social behaviours and cognitive processes have been combined, incrementally modified, and elaborated over time. Here, we studied two of these putative social building blocks in two closely related species of cichlids: Neolamprologus pulcher, a group-living species, and Telmatochromis temporalis, a non-grouping species. Otherwise well matched in ecology, this pair of species provide an excellent comparison point to understand how behavioural processes may have been modified in relation to the evolution of sociality. Using social assays in both the laboratory and in the field, we explored each species’ motivation to interact with conspecifics, and each species’ conflict resolution tactics. We found that individuals of the group living species, N. pulcher, displayed higher social motivation and were more likely to produce submission displays than were individuals of the non-grouping species, T. temporalis. We argue that the motivation to interact with conspecifics is a necessary prerequisite for the emergence of group living, and that the use of submission reduces the costs of conflict and facilitates the maintenance of close social proximity. These results suggest that social motivation and conflict resolution tactics are associated with social complexity, and that these behavioural traits may be functionally significant in the evolution and maintenance of sociality

Developmental plasticity of the stress response in female but not in male guppies

To survive, animals must respond appropriately to stress. Stress responses are costly, so early-life experiences with potential stressors could adaptively tailor adult stress responses to local conditions. However, how multiple stressors influence the development of the stress response remains unclear, as is the role of sex. Trinidadian guppies (Poecilia reticulata) are small fish with extensive life-history differences between the sexes and population variation in predation pressure and social density. We investigated how sex and early-life experience influence hormonal stress responses by manipulating conspecific density and perceived predation risk during development. In adults, we sampled cortisol twice to measure initial release and change over time in response to a recurring stressor. The sexes differed considerably in their physiological stress response. Males released more cortisol for their body mass than females and did not reduce cortisol release over time. By contrast, all females, except those reared at high density together with predation cues, reduced cortisol release over time. Cortisol responses of males were thus less dynamic in response to current circumstances and early-life experiences than females, consistent with life-history differences between the sexes. Our study underscores the importance of early-life experiences, interacting ecological factors and sex differences in the organization of the stress response

Head up displays are a submission signal in the group-living daffodil cichlid.

Dominance hierarchies can reduce conflict within social groups and agonistic signals can help to establish and maintain these hierarchies. Behaviours produced by subordinates in response to aggression are often assumed to function as signals of submission, however, these behaviours may serve other purposes, for example, defence or escape. For a behaviour to act as a submission signal, the receiver must respond by reducing their likelihood of further aggression towards the signaller. In the current study, we examine the receiver response to a putative signal of submission, the head up display, within established social groups of the cooperatively breeding fish, the daffodil cichlid (Neolamprologus pulcher). We found that when subordinate signallers produce the head up display in response to aggression from the breeder male, he exhibited a longer latency to behave aggressively towards that individual again. We also report that head up displays are rarely produced without being elicited by aggression, and the number of head up displays correlates with the amount of aggression received. Our results demonstrate that the head up display is used as a signal of submission in the daffodil cichlid and provide insight into intragroup communication in an emerging model system for the study of social behaviour

Submissive behaviour is mediated by sex, social status, relative body size and shelter availability in a social fish

Acting submissively may inhibit aggression and facilitate the termination of contests without further escalation. The need to minimize conflict is vital in highly social species where within 23 group interactions are frequent, and aggression can dampen group productivity. Within social groups, individual group members may modulate their use of submissive signals depending on their phenotype, the value of the contested resource, their relationship to the receiver of the signal and the characteristics of the local environment. We predicted that submissive behaviour would be more common when signallers had limited ability to flee from conflict, when signallers were of a low rank within the group, when signallers and receivers differed substantially in body size (and thus in fighting ability), and when signallers and receivers were of opposite sex and therefore not directly in competition over reproductive opportunities. We tested these predictions using social network analyses on detailed behavioural observations from 27 social groups of the cooperatively breeding cichlid fish Neolamprologus pulcher. Congruent with our prediction, submissive behaviour was more common when there were fewer shelters available, suggesting that constraints on fleeing behaviour may increase the use of submission. Also fitting with predictions, submissive behaviour was more common with increasing body size asymmetry between the competitors, among lower ranked fish and in interactions between opposite-sex dyads, which supports the idea that signalling submission is adaptive in contests over low-value resources. Our findings suggest that subordinate N.pulcher are primarily concerned with being tolerated within the social group and may use submissive behaviour to avoid escalated conflict. They offer a window into the factors that influence signals of submission in a highly social vertebrate

Horses show individual level lateralisation when inspecting an unfamiliar and unexpected stimulus.

Animals must attend to a diverse array of stimuli in their environments. The emotional valence and salience of a stimulus can affect how this information is processed in the brain. Many species preferentially attend to negatively valent stimuli using the sensory organs on the left side of their body and hence the right hemisphere of their brain. Here, we investigated the lateralisation of visual attention to the rapid appearance of a stimulus (an inflated balloon) designed to induce an avoidance reaction and a negatively valent emotional state in 77 Italian saddle horses. Horses' eyes are laterally positioned on the head, and each eye projects primarily to the contralateral hemisphere, allowing eye use to be a proxy for preferential processing in one hemisphere of the brain. We predicted that horses would inspect the novel and unexpected stimulus with their left eye and hence right hemisphere. We found that horses primarily inspected the balloon with one eye, and most horses had a preferred eye to do so, however, we did not find a population level tendency for this to be the left or the right eye. The strength of this preference tended to decrease over time, with the horses using their non-preferred eye to inspect the balloon increasingly as the trial progressed. Our results confirm a lateralised eye use tendency when viewing negatively emotionally valent stimuli in horses, in agreement with previous findings. However, there was not any alignment of lateralisation at the group level in our sample, suggesting that the expression of lateralisation in horses depends on the sample population and testing context

Subordinate submissive responses are predicted by dominant behaviour in a cooperatively breeding fish

Abstract In complex social groups, animals rely on communication to facilitate priority access to resources and minimise the costs of conflict. Animals typically have more aggression signals than submission signals. However, some social species do show multiple submission signals, and the context in which these different signals are used is often not well understood. In the current study, we assessed agonistic interactions within groups of the cooperatively breeding daffodil cichlid fish (Neolamprologus pulcher) to investigate the relationship between the aggressive behaviours of the dominant breeding pair, and the submissive responses of the highest ranked subordinate within the group. Daffodil cichlids may respond to aggression by fleeing or by the production of either a tail quiver display or a head up display. Among the two submission signals, the tail quiver display was used more frequently in response to a threat display, while head up displays were produced approximately equally in response to both threat displays and overt aggression. An exaggerated version of the head up display was given more often in response to overt aggression, suggesting a graded submissive response both within and between the two submission signals. Within fish, the frequency of head up displays, but not tail quiver displays, correlated positively with the frequency of threat displays received. The current study helps us to better understand the use of submission signals in a highly social vertebrate and sheds light on submission as an understudied aspect of communication

Heart and brain: Change in cardiac entropy is related to lateralised visual inspection in horses

Cerebral lateralisation is the tendency for an individual to preferentially use one side of their brain and is apparent in the biased use of paired sensory organs. Horses vary in eye use when viewing a novel stimulus which may be due to different physiological reactions. To understand the interplay between physiology and lateralisation, we presented a novel object (an inflated balloon) to 20 horses while electrocardiogram traces were collected. We measured the amount of time each horse looked at the balloon with each eye. We calculated ‘sample entropy’ as a measure of non-linear heart rate variability both prior to and during the stimulus presentation. A smaller drop in sample entropy values between the habituation phase and the sample presentation indicates the maintenance of a more complex signal associated with a relaxed physiological state. Horses that spent longer viewing the balloon with their left eye had a greater reduction in sample entropy, while time spend looking with the right eye was unrelated to the change in sample entropy. Therefore, the horses that exhibited a greater reduction in sample entropy tended to use their right hemisphere more, which may take precedence in emotional reactions. These results may help to explain the variation in lateralisation observed among horses

Isotocin neuronal phenotypes differ among social systems in cichlid fishes

Social living has evolved numerous times across a diverse array of animal taxa. An open question is how the transition to a social lifestyle has shaped, and been shaped by, the underlying neurohormonal machinery of social behaviour. The nonapeptide neurohormones, implicated in the regulation of social behaviours, are prime candidates for the neuroendocrine substrates of social evolution. Here, we examined the brains of eight cichlid fish species with divergent social systems, comparing the number and size of preoptic neurons that express the nonapeptides isotocin and vasotocin. While controlling for the influence of phylogeny and body size, we found that the highly social cooperatively breeding species (n = 4) had fewer parvocellular isotocin neurons than the less social independently breeding species (n = 4), suggesting that the evolutionary transition to group living and cooperative breeding was associated with a reduction in the number of these neurons. In a complementary analysis, we found that the size and number of isotocin neurons significantly differentiated the cooperatively breeding from the independently breeding species. Our results suggest that isotocin is related to sociality in cichlids and may provide a mechanistic substrate for the evolution of sociality

Consistency and flexibility in solving spatial tasks: different horses show different cognitive styles

Individual animals vary in their behaviour and reactions to novel situations. These differences may extend to differences in cognition among individuals. We tested twenty-six horses for their ability to detour around symmetric and asymmetric obstacles. All of the animals were able to get around the barrier to reach a food target, but varied in their approach. Some horses moved slowly but were more accurate in choosing the shortest way. Other horses acted quickly, consistently detoured in the same direction, and did not reliably choose the shortest way. The remaining horses shifted from a faster, directionally consistent response with the symmetric barrier, to a slower but more accurate response with the asymmetric barrier. The asymmetric barrier induced a reduction in heart rate variability, suggesting that this is a more demanding task. The different approaches used to solve the asymmetric task may reflect distinct cognitive styles in horses, which vary among individuals, and could be linked to different personality traits. Understanding equine behaviour and cognition can inform horse welfare and management

No evidence that footedness in pheasants influences cognitive performance in tasks assessing colour discrimination and spatial ability

The differential specialization of each side of the brain facilitates the parallel processing of information and has been documented in a wide range of animals. Animals that are more lateralized as indicated by consistent preferential limb use are commonly reported to exhibit superior cognitive ability as well as other behavioural advantages.We assayed the lateralization of 135 young pheasants (Phasianus colchicus), indicated by their footedness in a spontaneous stepping task, and related this measure to individual performance in either 3 assays of visual or spatial learning and memory. We found no evidence that pronounced footedness enhances cognitive ability in any of the tasks. We also found no evidence that an intermediate footedness relates to better cognitive performance. This lack of relationship is surprising because previous work revealed that pheasants have a slight population bias towards right footedness, and when released into the wild, individuals with higher degrees of footedness were more likely to die. One explanation for why extreme lateralization is constrained was that it led to poorer cognitive performance, or that optimal cognitive performance was associated with some intermediate level of lateralization. This stabilizing selection could explain the pattern of moderate lateralization that is seen in most non-human species that have been studied. However, we found no evidence in this study to support this explanation