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bioRxiv2017

Two sides of the same coin: monetary incentives concurrently improve and bias confidence judgments.

Decisions are accompanied by a feeling of confidence, i.e., a belief about the decision being correct. Confidence accuracy is critical, notably in high-stakes situations such as medical or financial decisionmaking. Here, we investigated how incentive motivation influences confidence accuracy by combining a perceptual task with a confidence incentivization mechanism. Importantly, by varying the magnitude and valence (gains or losses) of monetary incentives, we orthogonalized their motivational and affective components. Corroborating theories of rational decision-making and motivation, our results first reveal that the motivational value of incentives improves aspects of confidence accuracy. However, in line with a value-confidence interaction hypothesis we further show that the affective value of incentives concurrently biases confidence reports, thus degrading confidence accuracy. Finally, we demonstrate that the motivational and affective effects of incentives differentially impact how confidence builds on perceptual evidence. Altogether, these findings may provide new hints about confidence miscalibration in healthy or pathological contexts.

animal behavior and cognition↗

bioRxiv2017

Captive-bred Orangutans voluntarily choose to reward themselves with cabbage containing greater amounts of anthocyanin in a two-alternative decision task.

Anthocyanins are an important part of the human diet and the most commonly consumed plant secondary metabolites. They are potent antioxidants, and in several recent studies the ingestion of anthocyanins has been linked to positive health benefits for humans. Here, we show that when given a choice between two alternative samples of cabbage to ingest, captive born orangutans (n = 6) voluntarily chose the sample that contained greater amounts of anthocyanin. This occurred when they had to decide between samples of red cabbage (Brassica oleracea var. capitata f. rubra) from the same plant (p<0.05), and samples from green cabbage (Brassica oleracea var. capitata) (p<0.01). This indicates that anthocyanin holds a reward value for these hominids. There was no difference in L*a*b* colour between ingested and discarded samples in red cabbage, but when the choice was between two green samples, the animals chose samples that were more green and yellow. There was also no difference in the amount of lightness (L*) between chosen and discarded samples of either plant. It is therefore unclear if the animals use leaf colour in decision-making. In addition to other macro nutrients provided by plants, anthocyanin is also chosen by these endangered apes.

animal behavior and cognition↗

bioRxiv2017

Group size constraints may mask underlying similarities in social structure: a comparison of female elephant societies

We report on female Asian elephant social structure in Nagarahole and Bandipur National Parks (Kabini population), southern India, and examine the role of group size in affecting the outcome of social structure analysis in female elephants, which show high fission-fusion dynamics. Based on five years of data, we found the Kabini association network structured into highly modular communities that we call clans. We then modified the dataset (to obtain the Kabini 500-m dataset) to match sampling methods previously used in a study each of Asian (Uda Walawe) and African savannah (Samburu) elephants, so that network and association statistics could be compared across populations. Measures of association and network structure previously used were more similar amongst the Asian elephant populations compared to Samburu. The Samburu population formed a hierarchically-nested multilevel society whereas the Asian populations did not. However, we found hierarchical clustering levels in all three populations using Louvain community detection. Moreover, the average community sizes obtained through the Louvain method were not significantly different across populations, indicating basic similarities in social structure. Since fission-fusion dynamics allow for community members to form groups of different sizes, we examined the effect of average group size on association and network statistics. Higher average association index and degree, and lower average path length in Samburu compared to the Kabini 500-m dataset were explained by the larger average group size in Samburu. Thus, underlying similarities in the social networks of species showing fission-fusion dynamics may be obscured by differences in average group size.\n\nSignificance StatementVarious measures of associations and social network analyses have been used to compare social structures of different populations. We studied the social structure of female Asian elephants in a southern Indian population and compared it with those of a Sri Lankan Asian elephant population and an African savannah elephant population. We showed that, while there were social differences between the Asian and African savannah elephant populations using previous methods, there were basic similarities across all three populations using a method of network community detection. This discrepancy across analyses partly stemmed from differences in average group size between populations. Average group size in fission-fusion societies variously affected different association and network statistics, which has implications for inferences about social structure.

animal behavior and cognition↗

bioRxiv2017

Caenorhabditis elegans as an emerging model for studying the basic biology of anorectic effects of nicotine

Nicotine decreases food intake, and smokers often report that they smoke to control their weight. To see whether similar phenomena could be observed in the model organism Caenorhabditis elegans, we challenged drug-na{iota}ve nematodes with a chronic low (0.01 mM) and high (1 mM) nicotine concentration for 55 h (from hatching to adulthood). After that, we recorded changes in their behavior in a nicotine gradient, where they could choose a desired nicotine concentration. By using a combination of behavioral and morphometric methods, we found that both nicotine and food modulate worm behavior. In the presence of food the nematodes adapted to the low nicotine concentration, when placed in the gradient, chose a similar nicotine concentration like C. elegans adapted to the high nicotine concentration. However, in the absence of food, the nematodes adapted to the low nicotine concentration, when placed in the gradient of this alkaloid, chose a similar nicotine concentration like naive worms. The nematodes growing up in the presence of high concentrations of nicotine had a statistically smaller body size, compared to the control condition, and the presence of food did not cause any enhanced slowing movement. These results provide a platform for more detailed molecular and cellular studies of nicotine addiction and food intake in this model organism.

animal behavior and cognition↗

bioRxiv2017

Histone 4 acetylation regulates behavioral individuality in zebrafish

Animals can show very different behaviors even in isogenic populations, but the underlying mechanisms to generate this variability remain elusive. We found that laboratory and isogenic zebrafish (Danio rerio) larvae showed consistent individual behaviors when swimming freely in identical wells or in reaction to stimuli. We also found that this behavioral inter-individual variability was reduced when we impaired the histone deacetylation pathway. Individuals with high levels of histone H4 acetylation, and specifically H4K12, behaved similar to the average of the population, but those with low levels deviated from it. More precisely, we found a set of genomic regions whose histone H4 acetylation is reduced with the distance between the individual and the average population behavior. We found evidence that this modulation depends on a complex of Yin-yang 1 (YY1) and histone deacetylase 1 (HDAC1) that binds to and deacetylates these regions. These changes were not only maintained at the transcriptional level but also amplified, as most target regions were located near genes encoding transcription factors. We suggest that stochasticity in the histone deacetylation pathway participates the generation of genetic-independent behavioral inter-individual variability.

animal behavior and cognition↗

bioRxiv2017

Associative Learning from Replayed Experience

We develop an extension of the Rescorla-Wagner model of associative learning. In addition to learning from the current trial, the new model supposes that animals store and replay previous trials, learning from the replayed trials using the same learning rule. This simple idea provides a unified explanation for diverse phenomena that have proved challenging to earlier associative models, including spontaneous recovery, latent inhibition, retrospective revaluation, and trial spacing effects. For example, spontaneous recovery is explained by supposing that the animal replays its previous trials during the interval between extinction and test. These include earlier acquisition trials as well as recent extinction trials, and thus there is a gradual re-acquisition of the conditioned response. We present simulation results for the simplest version of this replay idea, where the trial memory is assumed empty at the beginning of an experiment, all experienced trials are stored and none removed, and sampling from the memory is performed at random. Even this minimal replay model is able to explain the challenging phenomena, illustrating the explanatory power of an associative model enhanced by learning from remembered as well as real experiences.

animal behavior and cognition↗

bioRxiv2017

Visual memories are stored along a compressed timeline

In continuous recognition the recency effect manifests as a decrease in accuracy and a sublinear increase in response time (RT) with the lag of a repeated stimulus. The recency effect could result from the gradual weakening of mnemonic traces. Alternatively, the recency effect could result from a search through a compressed timeline of recent experience. These two hypotheses make very different predictions about the shape of response time distributions. Using highly-memorable pictures to mitigate changes in accuracy enabled a detailed examination of the effect of recency on retrieval dynamics. The recency at which pictures were repeated ranged over two orders of magnitude across three experiments. Analysis of the RT distributions showed that the time at which memories became accessible changed with the recency of the probe, as predicted by a serial search model suggesting that visual memories can be accessed by sequentially scanning along a compressed representation of the past.

animal behavior and cognition↗

bioRxiv2017

Reduced foraging investment as an adaptation to patchy food sources: a phasic army ant simulation

Colonies of several ant species within the subfamily Dorylinae alternate stereotypical discrete phases of foraging and reproduction. Such phasic cycles are thought to be adaptive because they minimize the amount of foraging and the related costs, and at the same time enhance the colony-level ability to rely on patchily distributed food sources. In order to investigate these hypotheses, we use here a simple computational approach to study the population dynamics of two species of virtual ant colonies that differ quantitatively in their foraging investment. One species, which we refer to as \"phasic\", forages only half of the time, mirroring the phasic activity of some army ants; the other \"non-phasic\" species forages instead all the time. We show that, when foraging costs are relatively high, populations of phasic colonies grow on average faster than non-phasic populations, outcompeting them in mixed populations. Interestingly, such tendency becomes more consistent as food becomes more difficult to find but locally abundant. According to our results, reducing the foraging investment, for example by adopting a phasic lifestyle, can result in a reproductive advantage, but only in specific conditions. We thus suggest phasic colony cycles to have emerged together with the doryline specialization in feeding on the brood of other eusocial insects, a resource that is hard to obtain but highly abundant if available.

animal behavior and cognition↗

bioRxiv2017

Colour Preferences of UK Garden Birds at Supplementary Seed Feeders

Supplementary feeding of garden birds has benefits for both bird populations and human wellbeing. Birds have excellent colour vision, and show preferences for food items of particular colours, but research into colour preferences associated with artificial feeders is limited to hummingbirds. Here, we investigated the colour preferences of common UK garden birds foraging at seed-dispensing artificial feeders containing identical food. We presented birds simultaneously with an array of eight differently coloured feeders, and recorded the number of visits made to each colour over 370 30-minute observation periods in the winter of 2014/15. In addition, we surveyed visitors to a garden centre and science festival to determine the colour preferences of likely purchasers of seed feeders. Our results suggest that silver and green feeders were visited by higher numbers of individuals of several common garden bird species, while red and yellow feeders received fewer visits. In contrast, people preferred red, yellow, blue and green feeders.

animal behavior and cognition↗

bioRxiv2017

Colour vision models: a practical guide, some simulations, and colourvision R package.

O_LIHuman colour vision differs from the vision of other animals. The most obvious differences are the number and type of photoreceptors in the retina. E.g., while humans are insensitive to ultraviolet (UV) light, most non-mammal vertebrates and insects have a colour vision that spans into the UV. The development of colour vision models allowed appraisals of colour vision independent of the human experience. These models are now widespread in ecology and evolution fields. Here I present a guide to colour vision modelling, run a series of simulations, and provide a R package - colourvision - to facilitate the use of colour vision models.\nC_LIO_LII present the mathematical steps for calculation of the most commonly used colour vision models: Chittka (1992) colour hexagon, Endler & Mielke (2005) model, and Vorobyev & Osorio (1998) linear and log-linear receptor noise limited models (RNL). These models are then tested using identical simulated and real data. These comprise of reflectance spectra generated by a logistic function against an achromatic background, achromatic reflectance against an achromatic background, achromatic reflectance against a chromatic background, and real flower reflectance data against a natural background reflectance.\nC_LIO_LIWhen the specific requirements of each model are met, between model results are, overall, qualitatively and quantitatively similar. However, under many common scenarios of colour measurements, models may generate spurious values and/or considerably different predictions. Models that log-transform data and use relative photoreceptor outputs are prone to generate unrealistic results when the stimulus photon catch is smaller than the background photon catch. Moreover, models may generate unrealistic results when the background is chromatic (e.g. leaf reflectance) and the stimulus is an achromatic low reflectance spectrum.\nC_LIO_LIColour vision models are a valuable tool in several ecology and evolution subfields. Nonetheless, knowledge of model assumptions, careful analysis of model outputs, and basic knowledge of calculation behind each model are crucial for appropriate model application, and generation of meaningful and reproducible results. Other aspects of vision not incorporated into these models should be considered when drawing conclusion from model results.\nC_LI

animal behavior and cognition↗

bioRxiv2017

Information seeking and simulation: Roles of attention in guiding a goal-directed behavior

We usually actively seek out the information we need. However, it is still debated whether information seeking in decision situation is a purposeful behavior or a random process. We investigated this issue using the decision task involving multiple goal-directed event sequences, in which a contextual cue specifies an associated target and touch to the target delivers the reward. We found that the gaze followed the sequence of contextual cue to the associated target, which was eventually chosen. This fixation sequence from contextual cue to the associated target could be observed even when there were multiple goals and when the focus was shifted from one goal to another. To causally investigate the effects of sequential simulation, we directly manipulated the processing of the contextual cues and found its influence on the final choice of target. Furthermore, past episodes of the sequences influenced both final choices of targets and initial gaze to contextual cues. We interpret the results as suggesting that the internal process of simulating goal-directed event sequence drives information-seeking behavior such as attention/gaze in decision situations.

animal behavior and cognition↗

bioRxiv2017

Exploration and recency as the main proximate causes of probability matching: a reinforcement learning analysis

Research has not yet reached a consensus on why humans match probabilities instead of maximise in a probability learning task. The most influential explanation is that they search for patterns in the random sequence of outcomes. Other explanations, such as expectation matching, are plausible, but do not consider how reinforcement learning shapes peoples choices.\n\nWe aimed to quantify how human performance in a probability learning task is affected by pattern search and reinforcement learning. We collected behavioural data from 84 young adult participants who performed a probability learning task wherein the majority outcome was rewarded with 0.7 probability, and analysed the data using a reinforcement learning model that searches for patterns. Model simulations indicated that pattern search, exploration, recency (discounting early experiences), and forgetting may impair performance.\n\nOur analysis estimated that 85% (95% HDI [76, 94]) of participants searched for patterns and believed that each trial outcome depended on one or two previous ones. The estimated impact of pattern search on performance was, however, only 6%, while those of exploration and recency were 19% and 13% respectively. This suggests that probability matching is caused by uncertainty about how outcomes are generated, which leads to pattern search, exploration, and recency.

animal behavior and cognition↗

bioRxiv2017

Mapping the unknown: The spatially correlated multi-armed bandit

We introduce the spatially correlated multi-armed bandit as a task coupling function learning with the exploration-exploitation trade-off. Participants interacted with bi-variate reward functions on a two-dimensional grid, with the goal of either gaining the largest average score or finding the largest payoff. By providing an opportunity to learn the underlying reward function through spatial correlations, we model to what extent people form beliefs about unexplored payoffs and how that guides search behavior. Participants adapted to assigned payoff conditions, performed better in smooth than in rough environments, and--surprisingly--sometimes performed equally well in short as in long search horizons. Our modeling results indicate a preference for local search options, which when accounted for, still suggests participants were best-described as forming local inferences about unexplored regions, combined with a search strategy that directly traded off between exploiting high expected rewards and exploring to reduce uncertainty about the spatial structure of rewards.

animal behavior and cognition↗

bioRxiv2017

Optimal Response Vigor and Choice Under Non-stationary Outcome Values

Within a rational framework a decision-maker selects actions based on the reward-maximisation principle which stipulates they acquire outcomes with the highest values at the lowest cost. Action selection can be divided into two dimensions: selecting an action from several alternatives, and choosing its vigor, i.e., how fast the selected action should be executed. Both of these dimensions depend on the values of the outcomes, and these values are often affected as more outcomes are consumed, and so are the actions. Despite this, previous works have addressed the computational substrates of optimal actions only in the specific condition that the values of outcomes are constant, and it is still unknown what the optimal actions are when the values of outcomes are non-stationary. Here, based on an optimal control framework, we derive a computational model for optimal actions under non-stationary outcome values. The results imply that even when the values of outcomes are changing, the optimal response rate is constant rather than decreasing. This finding shows that, in contrast to previous theories, the commonly observed changes in the actions cannot be purely attributed to the changes in the outcome values. We then prove that this observation can be explained based on the uncertainty about temporal horizons; e.g., in the case of experimental protocols, the session duration. We further show that when multiple outcomes are available, the model explains probability matching as well as maximisation choice strategies. The model provides, therefore, a quantitative analysis of optimal actions and explicit predictions for future testing.

animal behavior and cognition↗

bioRxiv2017

Flexibility to contingency changes distinguishes habitual and goal-directed strategies in humans

Decision-making in the real world presents the challenge of requiring flexible yet prompt behavior, a balance that has been characterized in terms of a trade-off between a slower, prospective goal-directed model-based (MB) strategy and a fast, retrospective habitual model-free (MF) strategy. Theory predicts that flexibility to changes in both reward values and transition contingencies can determine the relative influence of the two systems in reinforcement learning, but few studies have manipulated the latter. Therefore, we developed a novel two-level contingency change task in which transition contingencies between states change every few trials; MB and MF control predict different responses following these contingency changes, allowing their relative influence to be inferred. Additionally, we manipulated the rate of contingency changes in order to determine whether contingency change volatility would play a role in shifting subjects between a MB and MF strategy. We found that human subjects employed a hybrid MB/MF strategy on the task, corroborating the parallel contribution of MB and MF systems in reinforcement learning. Further, subjects did not remain at one level of MB/MF behavior but rather displayed a shift towards more MB behavior over the first two blocks that was not attributable to the rate of contingency changes but rather to the extent of training. We demonstrate that flexibility to contingency changes can distinguish MB and MF strategies, with human subjects utilizing a hybrid strategy that shifts towards more MB behavior over blocks, consequently corresponding to a higher payoff.\n\nAuthor SummaryTo make good decisions, we must learn to associate actions with their true outcomes. Flexibility to changes in action/outcome relationships, therefore, is essential for optimal decision-making. For example, actions can lead to outcomes that change in value - one day, your favorite food is poorly made and thus less pleasant. Alternatively, changes can occur in terms of contingencies - ordering a dish of one kind and instead receiving another. How we respond to such changes is indicative of our decision-making strategy; habitual learners will continue to choose their favorite food even if the quality has gone down, whereas goal-directed learners will soon learn it is better to choose another dish. A popular paradigm probes the effect of value changes on decision making, but the effect of contingency changes is still unexplored. Therefore, we developed a novel task to study the latter. We find that humans used a mixed habitual/goal-directed strategy in which they became more goal-directed over the course of the task, and also earned more rewards with increasing goal-directed behavior. This shows that flexibility to contingency changes is adaptive for learning from rewards, and indicates that flexibility to contingency changes can reveal which decision-making strategy is used.

animal behavior and cognition↗

bioRxiv2017

The erratic and contingent progression of research on territoriality: a case study.

Our understanding of animal mating systems has changed dramatically with the advent of molecular methods to determine individuals reproductive success. But why are older behavioral descriptions and newer genetic descriptions of mating systems often seemingly inconsistent? We argue that a potentially important reason for such inconsistencies is a research trajectory rooted in early studies that were equivocal and overreaching, followed by studies that accepted earlier conclusions at face value and assumed, rather than tested, key ideas about animal mating systems. We illustrate our argument using Anolis lizards, whose social behavior has been studied for nearly a century. A dominant view emerging from this behavioral research was that anoles display strict territorial polygyny, where females mate with just the one male in whose territory they reside. However, all genetic evidence suggests that females frequently mate with multiple males. We trace this mismatch to early studies that concluded that anoles are territorial based on limited data. Subsequent research assumed territoriality implicitly or explicitly, resulting in studies that were unlikely to uncover or consider important any evidence of anoles departures from strict territorial polygyny. Thus, descriptions of anole behavior were largely led away from predicting a pattern of female multiple mating. We end by considering the broader implications of such erratic trajectories for the study of animal mating systems, and posit that precise definitions, renewed attention to natural history, and explicitly questioning assumptions made while collecting behavioral observations will allow us to move towards a fuller understanding of animal mating systems.

animal behavior and cognition↗

bioRxiv2017

Individual differences in responsivity to social rewards: Insights from two eye-tracking tasks

Humans generally prefer social over nonsocial stimuli from an early age. Reduced preference for social rewards has been observed in individuals with autism spectrum conditions (ASC). This preference has typically been noted in separate tasks that measure orienting toward and engaging with social stimuli. In this experiment, we used two eye-tracking tasks to index both of these aspects of social preference in in 77 typical adults. We used two measures, global effect and preferential looking time. The global effect task measures saccadic deviation toward a social stimulus (related to orienting), while the preferential looking task records gaze duration bias toward social stimuli (relating to engaging). Social rewards were found to elicit greater saccadic deviation and greater gaze duration bias, suggesting that they have both greater salience and higher value compared to nonsocial rewards. Trait empathy was positively correlated with the measure of relative value of social rewards, but not with their salience. This study thus elucidates the relationship of empathy with social reward processing.

animal behavior and cognition↗

bioRxiv2017

Hypoxia has lasting effects on fast startle behavior of a tropical fish, (Haemulon plumieri)

Summary statementThis study describes for the first time long-lasting behavioral effects of hypoxia on a tropical fish, the white grunt (Haemulon plumieri) from Puerto Rico.\n\nAnthropogenic activities and climate change have resulted in an increase in hypoxia in nearshore ecosystems worldwide. The San Juan Bay Estuary System in Puerto Rico is one such ecosystem that has undergone an increase in hypoxic events over the past few years. We collected white grunts (Haemulon plumieri) from one of the estuary lagoons to study the effects of hypoxia on fast startle responses (fast-starts). We hypothesized that exposure to hypoxia would significantly decrease the frequency of fast-starts evoked by an abrupt sound stimulus. After an exposure to an oxygen concentration of 2.5 mg L-1 (40% of air saturation), there is a significant reduction in the frequency of fast-starts that is maintained for at least 24 h after the exposure. Exposure to a random sequence of oxygen levels of 5.0, 4.3 and 3.7 mg L-1 (80, 70, and 60% of air saturation) did not show a significant effect until one hour after exposure. We speculate that the lasting effect of hypoxia on fast-starts, thought to be involved in escape, will result in a greater susceptibility of the white grunt to predation. We have identified the Mauthner cell, known to initiate fast-starts, to allow future studies on how low oxygen levels impact a single cell and its circuit, the behavior it initiates and ultimately how changes in the behavior affect population and ecosystem levels.

animal behavior and cognition↗