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    ABSTRACT: Decision-making under ambiguity in cognitive bias tasks is a promising new indicator of affective valence in animals. Rat studies support the hypothesis that animals in a negative affective state evaluate ambiguous cues negatively. Prior automated operant go/go judgement bias tasks have involved training rats that an auditory cue of one frequency predicts a Reward and a cue of a different frequency predicts a Punisher (RP task), and then measuring whether ambiguous cues of intermediate frequency are judged as predicting reward ('optimism') or punishment ('pessimism'). We investigated whether an automated Reward-Reward (RR) task yielded similar results to, and was faster to train than, RP tasks. We also introduced a new ambiguity test (simultaneous presentation of the two training cues) alongside the standard single ambiguous cue test. Half of the rats experienced an unpredictable housing treatment (UHT) designed to induce a negative state. Control rats were relatively 'pessimistic', whilst UHT rats were quicker, but no less accurate, in their responses in the RR test, and showed less anxiety-like behaviour in independent tests. A possible reason for these findings is that rats adapted to and were stimulated by UHT, whilst control rats in a predictable environment were more sensitive to novelty and change. Responses in the new ambiguity test correlated positively with those in single ambiguous cue tests, and may provide a measure of attention bias. The RR task was quicker to train than previous automated RP tasks. Together, they could be used to disentangle how reward and punishment processes underpin affect-induced cognitive biases.
    Behavioural Brain Research 08/2014; 274. DOI:10.1016/j.bbr.2014.07.048
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    ABSTRACT: The ability to discriminate between group-mates and non-group-mates likely underpins the occurrence of affiliative and aggressive behaviour towards 'in-group' and 'out-group' individuals. Here we present two experiments aimed at testing the ability of rats (Rattus norvegicus) to discriminate between cage-mate (CM: animals residing in the subject's home cage) and non-cage-mate (NCM) conspecifics. In experiment 1, rats were trained to discriminate between different exemplars of CM and NCM using a lever pressing task employing symmetrical reinforcement. Subjects did not reach performance criterion, but they did show some evidence of discrimination between the two types of stimuli. In experiment 2, we employed a digging task to determine if rats can discriminate between odour cues from CM and NCM presented simultaneously on two sand-filled bowls. Subjects reached performance criterion on the first pair of odours, and on three more different pairs of CM and NCM odours. The results of a reversal task, using a fifth pair of odours, indicate that the rats were using a common factor to discriminate between social cues from CM and NCM conspecifics, rather than learning each pair independently. Possible candidates include a group-specific odour cue, or the development of a CM/NCM category concept.
    Behavioural Processes 05/2014; 106. DOI:10.1016/j.beproc.2014.05.007
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    ABSTRACT: Intraspecific differences in retinal physiology have been demonstrated in several vertebrate taxa and are often subject to adaptive evolution. Nonetheless, such differences are currently unknown in birds, despite variations in habitat, behaviour and visual stimuli that might influence spectral sensitivity. The parrot Platycercus elegans is a species complex with extreme plumage colour differences between (and sometimes within) subspecies, making it an ideal candidate for intraspecific differences in spectral sensitivity. Here, the visual pigments of P. elegans were fully characterised through molecular sequencing of five visual opsin genes and measurement of their absorbance spectra using microspectrophotometry. Three of the genes, LWS, SW1 and SWS2, encode for proteins similar to those found in other birds; however, both the RH1 and RH2 pigments had polypeptides with carboxyl termini of different lengths and unusual properties that are unknown previously for any vertebrate visual pigment. Specifically, multiple RH2 transcripts and protein variants (short, medium and long) were identified for the first time that are generated by alternative splicing of downstream coding and non-coding exons. Our work provides the first complete characterisation of the visual pigments of a parrot, perhaps the most colourful order of birds, and moreover suggests more variability in avian eyes than hitherto considered.
    Journal of Experimental Biology 12/2013; 216(Pt 23):4454-4461. DOI:10.1242/jeb.094136
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    ABSTRACT: We developed a novel delay discounting task to investigate outcome impulsivity in pigs. As impulsivity can affect aggression, and might also relate to proactive and reactive coping styles, eight proactive (HR) and eight reactive (LR) pigs identified in a manual restraint test ("Backtest", after Bolhuis et al., 2003) were weaned and mixed in four pens of four unfamiliar pigs, so that each pen had two HR and two LR pigs, and aggression was scored in the nine hours after mixing. In the delay discounting task, each pig chose between two levers, one always delivering a small immediate reward, the other a large delayed reward with daily increasing delays, impulsive individuals being the ones discounting the value of the large reward quicker. Two novel strategies emerged: some pigs gradually switched their preference towards the small reward ('Switchers') as predicted, but others persistently preferred the large reward until they stopped making choices ('Omitters'). Outcome impulsivity itself was unrelated to these strategies, to urinary serotonin metabolite (5-HIAA) or dopamine metabolite (HVA) levels, aggression at weaning, or coping style. However, HVA was relatively higher in Omitters than Switchers, and positively correlated with behavioural measures of indecisiveness and frustration during choosing. The delay discounting task thus revealed two response strategies that seemed to be related to the activity of the dopamine system and might indicate a difference in execution, rather than outcome, impulsivity.
    Physiology & Behavior 08/2013; 120. DOI:10.1016/j.physbeh.2013.08.009
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    ABSTRACT: Contact zones between subspecies or closely related species offer valuable insights into speciation processes. A typical feature of such zones is the presence of clinal variation in multiple traits. The nature of these traits and the concordance among clines are expected to influence whether and how quickly speciation will proceed. Learned signals, such as vocalizations in species having vocal learning (e.g. humans, many birds, bats and cetaceans), can exhibit rapid change and may accelerate reproductive isolation between populations. Therefore, particularly strong concordance among clines in learned signals and population genetic structure may be expected, even among continuous populations in the early stages of speciation. However, empirical evidence for this pattern is often limited because differences in vocalisations between populations are driven by habitat differences or have evolved in allopatry. We tested for this pattern in a unique system where we may be able to separate effects of habitat and evolutionary history. We studied geographic variation in the vocalizations of the crimson rosella (Platycercus elegans) parrot species complex. Parrots are well known for their life-long vocal learning and cognitive abilities. We analysed contact calls across a ca 1300 km transect encompassing populations that differed in neutral genetic markers and plumage colour. We found steep clinal changes in two acoustic variables (fundamental frequency and peak frequency position). The positions of the two clines in vocal traits were concordant with a steep cline in microsatellite-based genetic variation, but were discordant with the steep clines in mtDNA, plumage and habitat. Our study provides new evidence that vocal variation, in a species with vocal learning, can coincide with areas of restricted gene flow across geographically continuous populations. Our results suggest that traits that evolve culturally can be strongly associated with reduced gene flow between populations, and therefore may promote speciation, even in the absence of other barriers.
    PLoS ONE 12/2012; 7(12):e50484. DOI:10.1371/journal.pone.0050484
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    ABSTRACT: It is well established that there is anxiety-related variation between observers in the very earliest, pre-attentive stage of visual processing of images such as emotionally expressive faces, often leading to enhanced attention to threat in a variety of disorders and traits. Whether there is also variation in early-stage affective (i.e. valenced) responses resulting from such images, however, is not yet known. The present study used the subliminal affective priming paradigm to investigate whether people varying in trait social anxiety also differ in their affective responses to very briefly presented, emotionally expressive face images. Participants (n = 67) completed a subliminal affective priming task, in which briefly presented and smiling, neutral and angry faces were shown for 10 ms durations (below objective and subjective thresholds for visual discrimination), and immediately followed by a randomly selected Chinese character mask (2000 ms). Ratings of participants' liking for each Chinese character indicated the degree of valenced affective response made to the unseen emotive images. Participants' ratings of their liking for the Chinese characters were significantly influenced by the type of face image preceding them, with smiling faces generating more positive ratings than neutral and angry ones (F(2,128) = 3.107, p<0.05). Self-reported social anxiety was positively correlated with ratings of smiling relative to neutral-face primed characters (Pearson's r = .323, p<0.01). Individual variation in self-reported mood awareness was not associated with ratings. Trait social anxiety is associated with individual variation in affective responding, even in response to the earliest, pre-attentive stage of visual image processing. However, the fact that these priming effects are limited to smiling and not angry (i.e. threatening) images leads us to propose that the pre-attentive processes involved in generating the subliminal affective priming effect may be different from those that generate attentional biases in anxious individuals.
    PLoS ONE 05/2012; 7(5):e37011. DOI:10.1371/journal.pone.0037011
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    ABSTRACT: The arms race between brood parasites and their hosts has led to many different host behaviours for avoiding parasitism. Some of these behaviours are social, and require the presence of conspecifics to work effectively: in response to alarm calls, some species engage in mobbing behaviour where neighbours join nest tenants in attacking and repelling an invading brood parasite. There are risks involved for the neighbours, but it has been demonstrated that social mobbing allows individuals to learn about the presence of brood parasites in the environment, suggesting that social learning is occurring. Here, I consider whether using social signals to alert naive individuals to the presence of brood parasites is a suitable strategy, compared with sitting tight on the nest in response to the signal (which should reduce the chances of being parasitized). I also compare the efficiency of these strategies with the case where individuals fail to change behaviour in response a brood parasite. Using an individual-based simulation model, I demonstrate that both mobbing and sitting tight are effective strategies in response to a signal, and that mobbing is more effective when the chances of being parasitized increase. These results are discussed and compared with known host-brood parasite relationships.
    Interface focus: a theme supplement of Journal of the Royal Society interface 04/2012; 2(2):217-25. DOI:10.1098/rsfs.2011.0080
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    ABSTRACT: The areas of wild land around the edges of agricultural fields are a vital resource for many species. These include insect pollinators, to whom field margins provide both nest sites and important resources (especially when adjacent crops are not in flower). Nesting pollinators travel relatively short distances from the nest to forage: most species of bee are known to travel less than two kilometres away. In order to ensure that these pollinators have sufficient areas of wild land within reach of their nests, agricultural landscapes need to be designed to accommodate the limited travelling distances of nesting pollinators. We used a spatially-explicit modelling approach to consider whether increasing the width of wild strips of land within the agricultural landscape will enhance the amount of wild resources available to a nesting pollinator, and if it would impact differently on pollinators with differing foraging strategies. This was done both by creating field structures with a randomised geography, and by using landscape data based upon the British agricultural landscape. These models demonstrate that enhancing field margins should lead to an increase in the availability of forage to pollinators that nest within the landscape. With the exception of species that only forage within a very short range of their nest (less than 125 m), a given amount of field margin manipulation should enhance the proportion of land available to a pollinator for foraging regardless of the distance over which it normally travels to find food. A fixed amount of field edge manipulation should therefore be equally beneficial for both longer-distance nesting foragers such as honeybees, and short-distance foragers such as solitary bees.
    PLoS ONE 10/2011; 6(10):e25971. DOI:10.1371/journal.pone.0025971
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    ABSTRACT: Although social behaviour can bring many benefits to an individual, there are also costs that may be incurred whenever the members of a social group interact. The formation of dominance hierarchies could offer a means of reducing some of the costs of social interaction, but individuals within the hierarchy may end up paying differing costs dependent upon their position within the hierarchy. These differing interaction costs may therefore influence the behaviour of the group, as subordinate individuals may experience very different benefits and costs to dominants when the group is conducting a given behaviour. Here, a state-dependent dynamic game is described which considers a pair of social foragers where there is a set dominance relationship within the pair. The model considers the case where the subordinate member of the pair pays an interference cost when it and the dominant individual conduct specific pairs of behaviours together. The model demonstrates that if the subordinate individual pays these energetic costs when it interacts with the dominant individual, this has effects upon the behaviour of both subordinate and the dominant individuals. Including interaction costs increases the amount of foraging behaviour both individuals conduct, with the behaviour of the pair being driven by the subordinate individual. The subordinate will tend to be the lighter individual for longer periods of time when interaction costs are imposed. This supports earlier suggestions that lighter individuals should act as the decision-maker within the pair, giving leadership-like behaviours that are based upon energetic state. Pre-existing properties of individuals such as their dominance will be less important for determining which individual makes the decisions for the pair. This suggests that, even with strict behavioural hierarchies, identifying which individual is the dominant one is not sufficient for identifying which one is the leader.
    PLoS Computational Biology 10/2011; 7(10):e1002252. DOI:10.1371/journal.pcbi.1002252
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    ABSTRACT: Functional explanations of behaviour often propose optimal strategies for organisms to follow. These 'best' strategies could be difficult to perform given biological constraints such as neural architecture and physiological constraints. Instead, simple heuristics or 'rules-of-thumb' that approximate these optimal strategies may instead be performed. From a modelling perspective, rules-of-thumb are also useful tools for considering how group behaviour is shaped by the behaviours of individuals. Using simple rules-of-thumb reduces the complexity of these models, but care needs to be taken to use rules that are biologically relevant. Here, we investigate the similarity between the outputs of a two-player dynamic foraging game (which generated optimal but complex solutions) and a computational simulation of the behaviours of the two members of a foraging pair, who instead followed a rule-of-thumb approximation of the game's output. The original game generated complex results, and we demonstrate here that the simulations following the much-simplified rules-of-thumb also generate complex results, suggesting that the rule-of-thumb was sufficient to make some of the model outcomes unpredictable. There was some agreement between both modelling techniques, but some differences arose - particularly when pair members were not identical in how they gained and lost energy. We argue that exploring how rules-of-thumb perform in comparison to their optimal counterparts is an important exercise for biologically validating the output of agent-based models of group behaviour.
    PLoS ONE 07/2011; 6(7):e22104. DOI:10.1371/journal.pone.0022104
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