Jordan, Alex
0000-0001-6131-9734 Berufsbeschreibung
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Modelling animal contests based on spatio-temporal dynamics
We present a general theoretical model for the spatio-temporal dynamics of animal contests. Inspired by interactions between physical particles, the model is formulated in terms of effective interaction potentials, which map typical elements of contest behaviour into empirically verifiable rules of contestant motion. This allows us to simulate the observable dynamics of contests in various realistic scenarios, notably in dyadic contests over a localized resource. Assessment strategies previously formulated in game-theoretic models, as well as the effects of fighting costs, can be described as variations in our model’s parameters. Furthermore, the trends of contest duration associated with these assessment strategies can be derived and understood within the model. Detailed description of the contestants’ motion enables the exploration of spatio-temporal properties of asymmetric contests, such as the emergence of chase dynamics. Overall, our framework aims to bridge the growing gap between empirical capabilities and theory in this widespread aspect of animal behaviour.
Parentage analysis across age cohorts reveals sex differences in reproductive skew in a group-living cichlid fish, Neolamprologus multifasciatus
Group-living animals are often faced with complex reproductive decisions, namely how to partition within-group reproduction, how to obtain extra-group reproduction, and how these two means of reproduction should be balanced. The solutions to these questions can be difficult to predict because ecological conditions can affect the scopes for within-group and extra-group reproduction in complex ways. For example, individuals that are restricted from moving freely around their habitats may have limited extra-group reproductive opportunities, but at the same time, groups may live in close proximities to one another, which could potentially have the opposite effect. The group-living cichlid fish, Neolamprologus multifasciatus, experiences such ecological conditions, and we conducted an intensive genetic parentage analysis to investigate how reproduction is distributed within and among groups for both males and females. We found that cohabiting males live in ‘high-skew’ societies, where dominant males monopolize the majority of within-group reproduction, while females live in ‘low-skew’ societies, where multiple females can produce offspring concurrently. Despite extremely short distances separating groups, we inferred only very low levels of extra-group reproduction suggesting that subordinate males have very limited reproductive opportunities. A strength of our parentage analysis lies in its inclusion of individuals that spanned a wide age range, from young fry to adults. We outline the logistical circumstances when very young offspring may not always be accessible to parentage researchers, and present strategies to overcome the challenges of inferring mating patterns from a wide age range of offspring.
Further evidence for the capacity of mirror self-recognition in cleaner fish and the significance of ecologically relevant marks
An animal that tries to remove a mark from its body that is only visible when looking into a mirror displays the capacity for mirror self-recognition (MSR), which has been interpreted as evidence for self-awareness. Conservative interpretations of existing data conclude that convincing evidence for MSR is currently restricted to great apes. Here, we address proposed shortcomings of a previous study on MSR in the cleaner wrasse Labroides dimidiatus, by varying preexposure to mirrors and by marking individuals with different colors. We found that (1) 14/14 new individuals scraped their throat when a brown mark had been provisioned, but only in the presence of a mirror; (2) blue and green color marks did not elicit scraping; (3) intentionally injecting the mark deeper beneath the skin reliably elicited spontaneous scraping in the absence of a mirror; (4) mirror-naive individuals injected with a brown mark scraped their throat with lower probability and/or lower frequency compared to mirror-experienced individuals; (5) in contrast to the mirror images, seeing another fish with the same marking did not induce throat scraping; and (6) moving the mirror to another location did not elicit renewed aggression in mirror-experienced individuals. Taken together, these results increase our confidence that cleaner fish indeed pass the mark test, although only if it is presented in ecologically relevant contexts. Therefore, we reiterate the conclusion of the previous study that either self-awareness in animals or the validity of the mirror test needs to be revised.
Social and spatial conflict drive resident aggression toward outsiders in a group-living fish
Group-living animals often experience within-group competition for resources like shelter and space, as well as for social status. Because of this conflict, residents may aggressively resist joining attempts by new members. Here, we asked whether different forms of competition mediate this response, specifically competition over 1) shelter, 2) spatial position within groups, and 3) social or sexual roles. We performed experiments on wild groups of Neolamprologus multifasciatus cichlids in Lake Tanganyika, either increasing or decreasing the number of shelters (empty snail shells) within their territories. We predicted that increases in resource abundance would reduce conflict and lower the aggression of residents toward presented conspecifics, while decreases in resources would increase aggression. We explored the effects of social conflict and spatial arrangement by introducing same or opposite sex conspecifics, at greater or lesser distances from resident subterritories. We found that changing the abundance of shells had no detectable effect on the responses of residents to presented conspecifics. Rather, aggression was strongly sex-dependent, with male residents almost exclusively aggressing presented males, and female residents almost exclusively aggressing presented females. For females, this aggression was influenced by the spatial distances between the presented conspecific and the resident female subterritory, with aggression scaling with proximity. In contrast, presentation distance did not influence resident males, which were aggressive to all presented males regardless of location. Overall, our results show that group residents respond to presented conspecifics differently depending on the type of competitive threat these potential joiners pose.
Aggression and spatial positioning of kin and non-kin fish in social groups
Group-living animals are faced with the challenge of sharing space and local resources amongst group members who may be either relatives or non-relatives. Individuals may reduce the inclusive fitness costs they incur from competing with relatives by either reducing their levels of aggression toward kin, or by maintaining physical separation between kin. In this field study, we used the group-living cichlid Neolamprologus multifasciatus to examine whether within-group aggression is reduced among group members that are kin, and whether kin occupy different regions of their group’s territory to reduce kin competition over space and local resources. We determined the kinship relationships among cohabiting adults via microsatellite genotyping and then combined these with spatial and behavioral analyses of groups in the wild. We found that aggressive contests between group members declined in frequency with spatial separation between their shelters. Female kin did not engage in aggressive contests with one another, whereas non-kin females did, despite the fact these females lived at similar distances from one another on their groups’ territories. Contests within male–male and male–female dyads did not clearly correlate with kinship. Non-kin male-male and male–female dyads lived at more variable distances from one another on their territories than their corresponding kin dyads. Together, our study indicates that contests among group members can be mediated by relatedness in a sex-dependent manner. We also suggest that spatial relationships can play an important role in determining the extent to which group members compete with one another.
Patterns of sex-biased dispersal are consistent with social and ecological constraints in a group-living cichlid fish
Background
Sex-biased dispersal is a common and widespread phenomenon that can fundamentally shape the genetic structure of the social environments in which animals live. For animals that live in and move between social groups, sex-biased dispersal can result in an asymmetry in the degree of relatedness among cohabiting males and females, which can have strong implications for their social evolution. In this study, we measured the relatedness structure within and across groups of a wild population of Neolamprologus multifasciatus, a highly-social, shell-dwelling cichlid fish endemic to Lake Tanganyika, East Africa. In total, we genotyped 812 fish from 128 social groups at 20 microsatellite loci. Neolamprologus multifasciatus live at high densities, and also experience strong ecological constraints on free movement throughout their habitat. At the same time, they exhibit sex differences in the degree of reproductive competition within their groups and this makes them an excellent model system for studying the factors associated with sex-biased dispersal.
Results
Social groups of N. multifasciatus consist of multiple males and females living together. We found that cohabiting females were unrelated to one another (Lynch-Ritland estimates of relatedness = 0.045 ± 0.15, average ± SD), while males shared much higher, albeit variable, levels of relatedness to other males in their groups (0.23 ± 0.27). We uncovered a pronounced decline in relatedness between males living in separate groups as the spatial separation between them increased, a pattern that was not evident in females. Female dispersal was also markedly constrained by the distribution and availability of nearby territories to which they could emigrate.
Conclusions
Our results indicate female-biased dispersal in N. multifasciatus. Our study also highlights how the spatial distribution of suitable dispersal destinations can influence the movement decisions of animals. We also emphasize how sex-biased dispersal can influence the relatedness structure of the social environment in which individuals interact and compete with one another.
Neural activity patterns differ between learning contexts in a social fish
Learning and decision-making are greatly influenced by context. When navigating a complex social world, individuals must quickly ascertain where to gain important resources and which group members are useful sources of such information. Such dynamic behavioural processes require neural mechanisms that are flexible across contexts. Here we examine how the social context influences the learning response during a cue discrimination task and the neural activity patterns that underlie acquisition of this novel information. Using the cichlid fish, Astatotilapia burtoni, we show that learning of the task is faster in social groups than in a non-social context. We quantify the neural activity patterns by examining the expression of Fos, an immediate-early gene, across brain regions known to play a role in social behaviour and learning (such as the putative teleost homologues of the mammalian hippocampus, basolateral amygdala and medial amygdala/BNST complex). We find that neural activity patterns differ between social and non-social contexts. Taken together, our results suggest that while the same brain regions may be involved in the learning of a cue association, the activity in each region reflects an individual's social context.
Regularly occurring bouts of retinal movements suggest an REM sleep-like state in jumping spiders
Sleep and sleep-like states are present across the animal kingdom, with recent studies convincingly demonstrating sleep-like states in arthropods, nematodes, and even cnidarians. However, the existence of different sleep phases across taxa is as yet unclear. In particular, the study of rapid eye movement (REM) sleep is still largely centered on terrestrial vertebrates, particularly mammals and birds. The most salient indicator of REM sleep is the movement of eyes during this phase. Movable eyes, however, have evolved only in a limited number of lineages-an adaptation notably absent in insects and most terrestrial arthropods-restricting cross-species comparisons. Jumping spiders, however, possess movable retinal tubes to redirect gaze, and in newly emerged spiderlings, these movements can be directly observed through their temporarily translucent exoskeleton. Here, we report evidence for an REM sleep-like state in a terrestrial invertebrate: periodic bouts of retinal movements coupled with limb twitching and stereotyped leg curling behaviors during nocturnal resting in a jumping spider. Observed retinal movement bouts were consistent, including regular durations and intervals, with both increasing over the course of the night. That these characteristic REM sleep-like behaviors exist in a highly visual, long-diverged lineage further challenges our understanding of this sleep state. Comparisons across such long-diverged lineages likely hold important questions and answers about the visual brain as well as the origin, evolution, and function of REM sleep.
Fast estimation of plant growth dynamics using deep neural networks
Background
In recent years, there has been an increase of interest in plant behaviour as represented by growth-driven responses. These are generally classified into nastic (internally driven) and tropic (environmentally driven) movements. Nastic movements include circumnutations, a circular movement of plant organs commonly associated with search and exploration, while tropisms refer to the directed growth of plant organs toward or away from environmental stimuli, such as light and gravity. Tracking these movements is therefore fundamental for the study of plant behaviour. Convolutional neural networks, as used for human and animal pose estimation, offer an interesting avenue for plant tracking. Here we adopted the Social LEAP Estimates Animal Poses (SLEAP) framework for plant tracking. We evaluated it on time-lapse videos of cases spanning a variety of parameters, such as: (i) organ types and imaging angles (e.g., top-view crown leaves vs. side-view shoots and roots), (ii) lighting conditions (full spectrum vs. IR), (iii) plant morphologies and scales (100 μm-scale Arabidopsis seedlings vs. cm-scale sunflowers and beans), and (iv) movement types (circumnutations, tropisms and twining).
Results
Overall, we found SLEAP to be accurate in tracking side views of shoots and roots, requiring only a low number of user-labelled frames for training. Top views of plant crowns made up of multiple leaves were found to be more challenging, due to the changing 2D morphology of leaves, and the occlusions of overlapping leaves. This required a larger number of labelled frames, and the choice of labelling “skeleton” had great impact on prediction accuracy, i.e., a more complex skeleton with fewer individuals (tracking individual plants) provided better results than a simpler skeleton with more individuals (tracking individual leaves).
Conclusions
In all, these results suggest SLEAP is a robust and versatile tool for high-throughput automated tracking of plants, presenting a new avenue for research focusing on plant dynamics.
Spatiotemporal dynamics of animal contests arise from effective forces between contestants
Competition among animals for resources, notably food, territories, and mates, is ubiquitous at all scales of life. This competition is often resolved through contests among individuals, which are commonly understood according to their outcomes and in particular, how these outcomes depend on decision-making by the contestants. Because they are restricted to end-point predictions, these approaches cannot predict real-time or real-space dynamics of animal contest behavior. This limitation can be overcome by studying systems that feature typical contest behavior while being simple enough to track and model. Here, we propose to use such systems to construct a theoretical framework that describes real-time movements and behaviors of animal contestants. We study the spatiotemporal dynamics of contests in an orb-weaving spider, in which all the common elements of animal contests play out. The confined arena of the web, on which interactions are dominated by vibratory cues in a two-dimensional space, simplifies the analysis of interagent interactions. We ask whether these seemingly complex decision-makers can be modeled as interacting active particles responding only to effective forces of attraction and repulsion due to their interactions. By analyzing the emergent dynamics of “contestant particles,” we provide mechanistic explanations for real-time dynamical aspects of animal contests, thereby explaining competitive advantages of larger competitors and demonstrating that complex decision-making need not be invoked in animal contests to achieve adaptive outcomes. Our results demonstrate that physics-based classification and modeling, in terms of effective rules of interaction, provide a powerful framework for understanding animal contest behaviors.