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bioRxiv2018

Inflation versus filling-in: why we feel we see more than we actually do in peripheral vision

Do we perceive fine details in the visual periphery? Here, we propose that phenomenology in the visual periphery can be characterized by an inflated sense of perceptual capacity, as observers overestimate the quality of their perceptual inputs. Distinct from the well-known perceptual phenomenon of \"filling-in\" where perceptual content is generated or completed endogenously, inflation can be characterized by incorrect introspection at the subjective level. The perceptual content itself may be absent or weak (i.e., not necessarily filled-in), and yet such content is mistakenly regarded by the system as rich. Behaviorally, this can be reflected by metacognitive deficits in the degree to which confidence judgments track task accuracy, and decisional biases for observers to think particular items are present, even when they are not. In two experiments using paradigms which exploit unique attributes of peripheral vision (crowding and summary statistics), we provide evidence that both types of deficits are present in peripheral vision, as observers reports are marked by overconfidence in discrimination judgments and high numbers of false alarms in detection judgments. We discuss potential mechanisms which may be the cause of inflation and propose future experiments to further explore this unique sensory phenomenon.

animal behavior and cognition↗

bioRxiv2018

The variability of song variability in wild and domesticated zebra finches Taeniopygia guttata

Birdsong is a classic example of a learned social behavior. Like many traits of interest, however, song production is also influenced by genetic factors and understanding the relative contributions of genetic and environmental influences remains a major research goal. In this study we take advantage of genetic variation among captive zebra finch populations to examine variation in a population-level song trait: song variability. We find that zebra finch populations differ in levels of song variability. Domesticated T. g. castanotis populations displayed higher song diversity than more recently wild-derived populations of both zebra finch subspecies T. g. castanotis and T. g. guttata, the Timor zebra finch. To determine whether these differences could have a genetic basis, we cross-fostered domesticated T. g. castanotis and Timor zebra finches to Bengalese finches Lonchura striata domestica. Following cross-fostering, domesticated T. g. castanotis maintained a higher level of song diversity than T. g. guttata. We suggest that the high song variability of domesticated zebra finches may be a consequence of reduced purifying selection acting on song traits. Intraspecific differences in the mechanisms underlying song variability therefore represent an untapped opportunity for probing the mechanisms of song learning and production.

animal behavior and cognition↗

bioRxiv2018

Continuous flash suppression and monocular pattern masking impact subjective awareness similarly

Peters & Lau (2015) found that when criterion bias is controlled for, there is no evidence for unconscious visual perception in normal observers, in the sense that they cannot directly discriminate a target above chance without knowing it. One criticism of that study is that the visual suppression method used, forward and backward masking (FBM), may be too blunt in the way it interferes with visual processing to allow for unconscious forced-choice discrimination. To investigate this question we compared FBM directly to continuous flash suppression (CFS) in a two-interval forced choice task. Although CFS is popular, and may be thought of as a more powerful visual suppression technique, we found no difference in the degree of perceptual impairment between the two suppression types. To the extent that CFS impairs perception, both objective discrimination and subjective awareness are impaired to similar degrees under FBM. This pattern was consistently observed across 3 experiments in which various experimental parameters were varied. These findings provide evidence for an ongoing debate about unconscious perception: normal observers cannot perform forced-choice discrimination tasks unconsciously.

animal behavior and cognition↗

bioRxiv2018

Hawkmoth flight in the unsteady wakes of flowers

Flying animals maneuver and hover through environments where wind gusts and flower wakes produce unsteady flow. Although both flight maneuvers and aerodynamic mechanisms have been studied independently, little is known about how these interact in an environment where flow is already unsteady. Moths forage from flowers by hovering in the flowers wake. We investigate hawkmoths tracking a 3D-printed robotic flower in a wind tunnel. We visualize the flow in the wake and around the wings and compare tracking performance to previous experiments in a still air flight chamber. Like in still air, moths flying in the flower wake exhibit near perfect tracking at low frequencies where natural flowers move. However, tracking in the flower wake results in a larger overshoot between 2-5 Hz. System identification of flower tracking reveals that moths also display reduced-order dynamics in wind, compared to still air. Smoke visualization of the flower wake shows that the dominant vortex shedding corresponds to the same frequency band as the increased overshoot. Despite these large effects on tracking dynamics in wind, the leading edge vortex (LEV) remains bound to the wing throughout the wingstroke and does not burst. The LEV also maintains the same qualitative structure seen in steady air. Persistence of a stable LEV during decreased flower tracking demonstrates the interplay between hovering and maneuvering.\n\nSummary statementWe examined how moths maneuver in the wake of flowers and discover that flower tracking dynamics are simplified compared to still air, while the leading edge vortex does not burst and extends continuously across the wings and thorax.

animal behavior and cognition↗

bioRxiv2018

Uncertainty and exploration

In order to discover the most rewarding actions, agents must collect information about their environment, potentially foregoing reward. The optimal solution to this \"explore-exploit\" dilemma is often computationally challenging, but principled algorithmic approximations exist. These approximations utilize uncertainty about action values in different ways. Some random exploration algorithms scale the level of choice stochasticity with the level of uncertainty. Other directed exploration algorithms add a \"bonus\" to action values with high uncertainty. Random exploration algorithms are sensitive to total uncertainty across actions, whereas directed exploration algorithms are sensitive to relative uncertainty. This paper reports a multi-armed bandit experiment in which total and relative uncertainty were orthogonally manipulated. We found that humans employ both exploration strategies, and that these strategies are independently controlled by different uncertainty computations.

animal behavior and cognition↗

bioRxiv2018

Differences In Neural Activity, But Not Behavior, Across Social Contexts In Guppies, Poecilia reticulata

Animals are continually faced with the challenge of producing context-appropriate social behaviors. In many instances, animals produce unrelated behaviors across contexts. However, in some instances the same behaviors are produced across different social contexts, albeit in response to distinct stimuli and with distinct purposes. We took advantage of behavioral similarities across mating and aggression contexts in guppies, Poecilia reticulata, to understand how patterns of neural induction differ across social contexts when behaviors are nonetheless shared across contexts. While these is growing interest in understanding behavioral mechanisms in guppies, resources are sparse. As part of this study, we developed a neuroanatomical atlas of the guppy brain as a research community resource. Using this atlas, we found that neural activity in the preoptic area reflected social context, whereas individual differences in behavioral motivation paralleled activity in the posterior tuberculum and ventral telencephalon (teleost homologs of the ventral tegmental area and lateral septum, respectively). Our findings suggest independent coding of social salience versus behavioral motivation when behavioral repertoires are shared across social contexts.\n\nSummary statementActivity in distinct brain regions reflects behavioral context versus social motivation in a in which behavioral repertoires are shared across social contexts (Trinidadian guppies, Poecilia reticulata).

animal behavior and cognition↗

bioRxiv2018

Protein pheromone MUP20/Darcin is a vector and target of indirect genetic effects in mice.

Social behavior in animals is an adaptive process influenced by environmental factors and direct and indirect genetic effects. Indirect genetic effects (IGEs) include mechanisms by which individuals of particular genotypes can influence the behavioral phenotypes and genotypes (via modulated patterns of gene expression) of other individuals with different genotypes. In groups of adult mice, IGEs can be unidirectional, from one genotype to the other, or bidirectional, resulting in a homogenization of the behavioral phenotypes within the group. Critically, it has been theorized that IGEs constitute a large fitness target on which deleterious mutations can have pleiotropic effects, meaning that individuals carrying certain behavior-altering mutations can impose the fitness costs of those mutations on others comprising the broader social genome. Experimental data involving a mouse model support the existence of these IGE-amplified fitness losses; however, the underlying biological mechanisms that facilitate these remain unknown. In a mouse model of IGEs, we demonstrate that the Major Urinary Protein 20 pheromone, also called Darcin, produced by mice lacking the adhesion protein Neuroligin-3 acts as a vector to deleteriously modify the social behavior of wild-type mice. Additionally, we showed that lack of social interest on the part of Neuroligin-3 knockout mice is independent of their environment. These findings reveal a new role for mammalian pheromones in mediating the externalization of social deficits from one individual to others comprising the population through IGEs.\n\nAuthor SummaryIndirect genetic effects (IGEs) are mechanisms by which individuals of particular genotypes can influence the behavioral phenotype of individuals of different genotypes, sometimes disruptively, in instances where one member of the population carriers a deleterious behavior altering variant. Although disruptive IGEs have been demonstrated in mice, its underlying molecular and genetic mechanisms remain unknown. Using an IGEs mouse model, we demonstrated that the pheromone protein Major Urinary Protein 20, also named Darcin, is as a vector and target of social epistasis a specific type of IGEs. This finding reveals a new function for mammalian pheromones in mediating social epistasis to degrade group social behavior.

animal behavior and cognition↗

bioRxiv2018

Inverted Face with Upright Body:Evidence for Face Inversion Effect in Japanese Macaques (Macaca fuscata) under the Preferential Looking Procedure

Young Japanese macaques (Macaca fuscata) looked at the photographs that contained faces of the two species of macaques (M. fuscata and M. mulatta) for longer time when they were presented at upright orientation than inverted orientation, and that species discrimination was deteriorated when the photographs were presented at inverted orientation [16]. The present experiment further explored the factor affecting the inversion effect observed in the previous experiments. Five young laboratory-born Japanese macaques looked at the photographs by pressing the lever under the conjugate schedule of sensory reinforcement, that is, successive preferential looking procedure. Photographs of macaques were scrambled and made up two types of the bizarre photographs: upright face with upright body (orientation consistent) and inverted face with upright body (orientation inconsistent). These two types of photographs were presented at upright and inverted orientations. The monkeys looked longer when the face was upright than inverted. Orientation of body and background had no effect on looking time duration. The present results strongly suggest that inversion effect found in macaques is face-specific.

animal behavior and cognition↗

bioRxiv2018

Food hardness and stone tool weight in wild primate nut-cracking

This study presents data on average stone tool weights and the hardness of foods processed by the three known stone-tool-using primate species: Burmese long-tailed macaques (Macaca fascicularis aurea), bearded capuchins (Sapajus libidinosus) and Western chimpanzees (Pan troglodytes verus). Each of these primates uses stone hammers to crack open nuts in the wild, making them suitable for inter-species behavioural comparison. This work draws on published results to identify a distinct difference in the tool weight/food hardness curve between chimpanzees and the two monkey taxa, with the latter reaching an asymptote in mean tool weight of just over 1 kg regardless of increasing food hardness. In contrast, chimpanzees rapidly increase their tool weight in response to increasing hardness, selecting average masses over 5 kg to process the hardest nuts. Species overlap in their preference for tools of 0.8-1 kg for opening foods of hardness 2-3 kN, suggesting that this conjunction may represent a primate stone-tool-use optimum.

animal behavior and cognition↗

bioRxiv2018

Familiarity and Face-Inversion Effect in Japanese Macaques (Macaca fuscata) during the Preferential Looking Task

Four young laboratory-born Japanese macaques (Macaca fuscata) looked at the photographs of familiar and unfamiliar persons presented at upright and inverted orientations by pressing the lever under the conjugate schedule of sensory reinforcement (successive preferential looking procedure). Three types of photographs were prepared: photographs with persons taken in front view, those taken in back, and those without persons. The monkeys looked longer when the face was upright than inverted only for the pictures containing unfamiliar person with front view. The other types of photographs did not cause inversion effect. Familiarity weakened the face-specific inversion effect in monkeys. This difference may be due to in part the lower preference for familiar faces and the difference in processing mode between familiar and unfamiliar faces.

animal behavior and cognition↗

bioRxiv2018

Information sharing for a coordination game in fluctuating environments

Collective action dilemmas pervade the social and biological sciences - from human decision-making to bacterial quorum sensing. In these scenarios, individuals sense cues from the environment to adopt a suitable phenotype or change in behavior. However, when cues include signals from other individuals, then the appropriate behavior of each individual is linked. Here, we develop a framework to quantify the influence of information sharing on individual behavior in the context of two player coordination games. In this framework, the environment stochastically switches between two states, and the state determines which one of two actions players must coordinate on. Given a stochastically switching environment, we then consider two versions of the game that differ in the way players acquire information. In the first model, players independently sense private environmental cues, but do not communicate with each other. We find there are two types of strategies that emerge as Nash equilibria and fitness maximizers - players prefer to commit to one particular action when private information is poor, or prefer to employ phenotypic plasticity when it is good. The second model adds an additional layer of communication, where players share social cues as well. When the quality of social information is high, we find the socially optimal strategy is a novel \"majority logic\" strategy that bases decision-making on social cues. Our game-theoretic approach offers a principled way of investigating the role of communication in group decision-making under uncertain conditions.

animal behavior and cognition↗

bioRxiv2018

Making honey bees lie: experimental dissociation of flight experience and dance communication.

Honey bees use their dance to communicate flight distance and direction of a food source to their nest mates in the hive. How bees transpose flight information to generate a corresponding walking (dance) behavior is still unknown. We now present a detailed study of the changes in dance duration of individual bees after shifting feeder distance. Our experiments indicated that most bees needed two or more foraging trips to the new position before showing an updated dance duration. In addition, only a few bees significantly changed dance duration immediately, whereas most bees first produced intermediary durations. Double shift experiments showed that under certain conditions bees do not update dance duration but continued to perform dance duration for the previously visited feeder position. We propose that generation of dance information involves two memory contents one for newly acquired and one for previously stored distance information.\n\nOne Sentence SummaryGeneration of dance information is temporally separated from immediate flight experience and involves two different memory contents.

animal behavior and cognition↗

bioRxiv2018

Spacing of Cue-approach Training Leads to Better Maintenance of Behavioral Change

The maintenance of behavioral change over the long term is essential to achieve public health goals such as combatting obesity and drug use. Previous work by our group has demonstrated a reliable shift in preferences for appetitive foods following a novel non-reinforced training paradigm. In the current studies, we tested whether distributing training trials over two consecutive days would affect preferences immediately after training as well as over time at a one-month follow-up. In four studies, three different designs and an additional pre-registered replication of one sample, we found that spacing of cue-approach training induced a shift in food choice preferences over one month. The spacing and massing schedule employed governed the long-term changes in choice behavior. Applying spacing strategies to training paradigms that target automatic processes could prove a useful tool for the long-term maintenance of health improvement goals with the development of real-world behavioral change paradigms that incorporate distributed practice principles.

animal behavior and cognition↗

bioRxiv2018

Adaptive planning in human search

How do people plan ahead when searching for rewards? We investigate planning in a foraging task in which participants search for rewards on an infinite two-dimensional grid. Our results show that their search is best-described by a model which searches at least 3 steps ahead. Furthermore, participants do not seem to update their beliefs during planning, but rather treat their initial beliefs as given, a strategy similar to a heuristic called root-sampling. This planning algorithm corresponds well with participants behavior in test problems with restricted movement and varying degrees of information, outperforming more complex models. These results enrich our understanding of adaptive planning in complex environments.

animal behavior and cognition↗

bioRxiv2018

Value-based decisions involve sequential sampling from memory

Deciding between two equally appealing options can take considerable time. This observation has puzzled economists and philosophers, because more deliberation only delays the reward. Here we show that this seemingly irrational behavior is explained by the constructive use of memory. Using functional brain imaging in humans, we show that how long it takes to decide between two familiar food items is related to activity in the hippocampus, within specific regions shown to be associated with the retrieval of long-term memories. Moreover, we show that value is partially constructed during deliberation to resolve preference, and this constructive process changes behavior and brain responses. These results render memory as a supplier of evidence in value-based decisions, resolving a central paradox of choice.

animal behavior and cognition↗

bioRxiv2018

Functional Social Structure in Baboons: Modelling Interactions Between Social and Environmental Structure in Group-Level Foraging

In mobile social groups, cohesion is thought to be driven by patterns of attraction at both the individual and group level. In long-lived species with high group stability and repeated interactions, such as baboons, individual-to-individual attractions have the potential to play a large role in group cohesion and overall movement patterns. In previous work, we used GPS mapping of a group of baboons in De Hoop, Western Cape, South Africa, to demonstrate the influence of such attractions on movement patterns. We also demonstrated the existence of emergent group-level structures, which arose as a consequence of individual social influence. Specifically, we found a core-periphery structure, in which a subset of influential animals exerted an influence on each other and those animals in the periphery, while those in the periphery were influenced by the core but did not exert any influence over others. Here, we use agent-based modelling of baboon groups to investigate whether this group-level structure has any functional consequences for foraging behaviour. By varying individual level attractions, we produced baboon groups that contained influence structures that varied from more to less centralized. Our results suggest that varying centrality affects both the ability of the group to detect resource structure in the environment, as well as the ability of the group to exploit these resources. Our models predict that foraging groups with more centralized social structures will show a reduction in detection and an increase in exploitation of resources in their environment, and will produce more extreme foraging outcomes. More generally, our results highlight the link between social and environmental structure on functional outcomes for mobile social groups of animals.

animal behavior and cognition↗

bioRxiv2018

First data-driven approach to using individual cattle weights to estimate mean adult dairy cattle weight in the UK

IntroductionKnowledge of accurate weights of cattle is crucial for effective dosing of individual animals with medicine and for reporting antimicrobial usage metrics, amongst other uses. The most common weight for dairy cattle presented in current literature is 600 kg, but this is not evidenced by data. For the first time, we provide an evidence-based estimate of the average weight of UK dairy cattle to better inform decisions by farmers, veterinarians and the scientific community.\n\nMethodsWe collected data for 2,747 dairy cattle from 20 farms in the UK, 19 using Lely Automatic Milking Systems with weigh floors and 1 using a crush with weigh scales. These data covered farms with different breed types, including Holstein, Friesian, Holstein-Friesian and Jersey, as well as farms with dual purpose breeds and cross-breeds. Data were used to calculate a mean weight for dairy cattle by breed, and a UK-specific mean weight was generated by scaling to UK-specific breed proportions. Trends in weight by lactation number, DIM and production level were also explored using individual cattle-level data.\n\nResultsMean weight for adult dairy cattle included in this study was 617 kg (standard deviation (sd) 85.6 kg). Mean weight varied across breeds, with a range of 466 kg (sd=56.0 kg, Jersey) to 636 kg (sd=84.1, Holsteins). When scaled to UK breed proportions, the estimated mean UK dairy cattle weight was 620 kg. Overall, first-lactation heifers weighed 9% less than cows. Mean weight declined for the first 30 days post-calving, before steadily increasing. For cattle at peak production, mean weight increased with production level.\n\nConclusionsThis study is the first to calculate a mean weight of adult dairy cattle in the UK based on on-farm data. Overall mean weight was higher than that most often proposed in the literature (600 kg). Evidence-informed weights are crucial as the UK works to better monitor and report metrics to monitor antimicrobial use and are useful to farmers and veterinarians to inform dosing decisions.

animal behavior and cognition↗

bioRxiv2018

VideoTagger: User-Friendly Software for Annotating Video Experiments of Any Duration

BackgroundScientific insight is often sought by recording and analyzing large quantities of video. While easy access to cameras has increased the quantity of collected videos, the rate at which they can be analyzed remains a major limitation. Often, bench scientists struggle with the most basic problem that there is currently no user-friendly, flexible, and open source software tool with which to watch and annotate these videos.\n\nResultsWe have created the VideoTagger tool to address these and many of the other associated challenges of video analysis. VideoTagger allows non-programming users to efficiently explore, annotate, and visualize large quantities of video data, within their existing experimental protocols. Further, it is built to accept programmed plugins written in Python, to enable seamless integration with other sophisticated computer-aided analyses.\n\nWe tested VideoTagger ourselves, and have a growing base of users in other scientific disciplines. Capitalising on the unique features of VideoTagger to play back infinite lengths of video footage at various speeds, we annotated 39h of a Drosophila melanogaster lifespan video, at approximately 10-15x faster than real-time. We then used these labels to train a machine-learning plugin, which we used to annotate an additional 538h of footage automatically. In this way, we found that flies fall over spontaneously with increasing frequency as they age, and also spend longer durations struggling to right themselves. Ageing in flies is typically defined by length of life. We propose that this new mobility measure of ageing could help the discovery of mechanisms in biogerontology, refining our definition of what healthy ageing means in this extremely small, but widely used, invertebrate.\n\nConclusionsWe show how VideoTagger is sufficiently flexible for studying lengthy and/or numerous video experiments, thus directly improving scientists productivity across varied domains.

animal behavior and cognition↗