Sleep in the wild: the importance of individual effects and environmental conditions on sleep behaviour in wild boar.

Sleep serves vital physiological functions, yet how sleep in wild animals is influenced by environmental conditions is poorly understood. Here we use high-resolution biologgers to investigate sleep in wild animals over ecologically relevant time scales and quantify variability between individuals under changing conditions. We developed a robust classification for accelerometer data and measured multiple dimensions of sleep in the wild boar (Sus scrofa) over an annual cycle. In support of the hypothesis that environmental conditions determine thermoregulatory challenges, which regulate sleep, we show that sleep quantity, efficiency and quality are reduced on warmer days, sleep is less fragmented in longer and more humid days, while greater snow cover and rainfall promote sleep quality. Importantly, this longest and most detailed analysis of sleep in wild animals to date reveals large inter- and intra-individual variation. Specifically, short-sleepers sleep up to 46% less than long-sleepers but do not compensate for their short sleep through greater plasticity or quality, suggesting they may pay higher costs of sleep deprivation. Given the major role of sleep in health, our results suggest that global warming and the associated increase in extreme climatic events are likely to negatively impact sleep, and consequently health, in wildlife, particularly in nocturnal animals.

Neonate personality affects early-life resource acquisition in a large social mammal.

Although it is widely acknowledged that animal personality plays a key role in ecology, current debate focuses on the exact role of personality in mediating life-history trade-offs. Crucial for our understanding is the relationship between personality and resource acquisition, which is poorly understood, especially during early stages of development. Here we studied how among-individual differences in behavior develop over the first 6 months of life, and their potential association with resource acquisition in a free-ranging population of fallow deer (Dama dama). We related neonate physiological (heart rate) and behavioral (latency to leave at release) anti-predator responses to human handling to the proportion of time fawns spent scanning during their first summer and autumn of life. We then investigated whether there was a trade-off between scanning time and foraging time in these juveniles, and how it developed over their first 6 months of life. We found that neonates with longer latencies at capture (i.e., risk-takers) spent less time scanning their environment, but that this relationship was only present when fawns were 3-6 months old during autumn, and not when fawns were only 1-2 months old during summer. We also found that time spent scanning was negatively related to time spent foraging and that this relationship became stronger over time, as fawns gradually switch from a nutrition rich (milk) to a nutrition poor (grass) diet. Our results highlight a potential mechanistic pathway in which neonate personality may drive differences in early-life resource acquisition of a large social mammal.

The effects of stimulus distribution form during trace conditioning.

Three experiments examined the effect of distribution form of the trace interval on trace conditioning. In Experiments 1 and 2, two groups of rats were conditioned to a fixed-duration conditioned stimulus (CS) in a trace interval procedure; rats in Group Fix received a fixed-duration trace interval, whereas for rats in Group Var the trace interval was of variable duration. Responding during the CS was higher in Group Var than in Group Fix, whereas during the trace interval this difference in responding reversed-Group Fix showed higher response rates than Group Var. Experiment 3 examined whether the greater response rate observed during the CS in Group Var was due to a performance effect or the acquisition of greater associative strength by the CS. Following trace conditioning, the rats from Experiment 1 underwent a second phase of delay conditioning with the same CS; a 5-s auditory stimulus was presented in compound with the last 5 s of the 15-s CS, and the unconditioned stimulus (US) was delivered at the offset of the CSs. On test with the auditory stimulus alone, subjects in Group Var showed lower response rates during the auditory stimulus than subjects in Group Fix. We interpreted these findings as evidence that the superior responding in Group Var during the CS was a result of it acquiring greater associative strength than in Group Fix.

Lateralization during lateral display and its relationship with antler size and symmetry in fallow deer (Dama dama).

Individuals often display a preference for one side of their body during aggressive encounters. This may be a lateralized preference for using one structure of a bilateral trait during display or physical attack, or for keeping the opponent in one visual field. Alternatively, it may be the case that behavioural lateralization and the degree of symmetry expressed by bilateral structures are correlated forms of developmental instability. We examined whether there was an association between lateralization during a lateral display and different measurements of antler size and symmetry (beam length, beam circumference, brow tine length and coronet circumference). Three models addressed different structural measures: the right antler, the larger antler and antler symmetry. Results showed that beam length was negatively associated with behavioural lateralization irrespective of structural measure. A second analysis using a composite score of the four antler measurements, one for each structural measure, showed that only antler symmetry was negatively associated with lateralization during lateral display. Therefore, our second prediction was supported. We discuss these findings in relation to predator detection capability and stress reduction in prey species such as the fallow deer.

Weapon damage is associated with contest dynamics but not mating success in fallow deer (Dama dama).

Antlers function as primary weapons during fights for many species of ungulate. We examined the association between antler damage and (i) contest dynamics: the behavioural tactics used during fighting including fight duration, and (ii) mating success, fighting rate and dominance. Structural damage of the antlers was associated with contest dynamics: damage was negatively associated with jump clash attacks by individuals with damaged antlers, whereas opponents were more likely to physically displace individuals with damaged antlers during fighting. We found a positive association between dominance and damage indicating that high-ranking individuals were likely to have breaks to their antlers. We found no evidence that damage was associated with either mating success or the number of fights individuals engaged in. Our study provides a new perspective on understanding the association between contest dynamics and weapon structure, while also showing that damage has limited fitness consequences for individuals.

Blocking by fixed and variable stimuli: Effects of stimulus distribution on blocking.

An experiment with rats compared the ability of fixed and variable duration cues to produce blocking. Rats in group B (Blocking) were trained that both fixed- (F) and variable- (V) duration cues would be followed by food delivery. In a subsequent training stage F and V continued to be reinforced, but F was accompanied by X, and V by Y. In the test phase responding to X and Y was examined. Control group O (Overshadowing) received identical treatment, except that F and V were nonreinforced in the first training stage. In group B there was evidence for blocking, but only of X, which had been conditioned in compound with the fixed-duration F; there was no evidence for blocking of Y, which had been conditioned in compound with the variable-duration V. It is suggested that this result may occur because fixed cues reach a higher, more stable asymptote of associative strength than do their variable equivalents.

Learning about the CS during latent inhibition: Preexposure enhances temporal control.

In 3 experiments, rats were given nonreinforced preexposure to an auditory stimulus, after which this stimulus and a second, novel cue were paired with food. Lower rates of conditioned responding were observed to the preexposed stimulus across the 3 experiments, indicative of latent inhibition. The degree to which animals used these cues to time the occurrence of food delivery was also examined. Paradoxically, the response slopes-indicating the rate of increase in responding over the course of the conditioned stimulus-were greater for the preexposed than for the novel cues, consistent with the suggestion that the preexposed stimulus exerted greater temporal control. Moreover, this was the case irrespective of whether the duration of the cue during preexposure differed from that during conditioning. These results suggest that although conditioned stimulus preexposure retards conditioning, it may enhance timing. The findings are discussed in terms of current models of conditioning and timing.

Effects of dorsal hippocampal damage on conditioning and conditioned-response timing: A pooled analysis.

Behavioral findings suggest that the dorsal hippocampus (DHPC) plays a role in timing of appetitive conditioned responding. The present article explored the relationship between the extent of DHPC damage and timing ability, in a pooled analysis of three published studies from our laboratory. Initial analyses of variance confirmed our previous reports that DHPC damage reduced peak time (a measure of timing accuracy). However, the spread (a measure of timing precision) was unchanged, such that the coefficient of variation (spread/peak time) was significantly larger in DHPC-lesioned animals. This implies that, in addition to the well-established effect of DHPC lesions on timing accuracy, DHPC damage produced a deficit in precision of timing. To complement this analysis, different generalized linear mixed-effects models (GLMMs) were performed on the combined dataset, to examine which combinations of the different behavioral measures of timing were the best predictors of the degree of hippocampal damage. The results from the GLMM analysis suggested that the greater the DHPC damage, the greater the absolute difference between the observed peak time and reinforced duration. Nevertheless, this systematic relationship between damage and performance was not specific to the temporal domain: paradoxically the greater the damage the greater the magnitude of peak responding. We discuss these lesion effects in terms of scalar timing theory.

Overshadowing by fixed- and variable-duration stimuli.

Two experiments investigated the effect of the temporal distribution form of a stimulus on its ability to produce an overshadowing effect. The overshadowing stimuli were either of the same duration on every trial, or of a variable duration drawn from an exponential distribution with the same mean duration as that of the fixed stimulus. Both experiments provided evidence that a variable-duration stimulus was less effective than a fixed-duration cue at overshadowing conditioning to a target conditioned stimulus (CS); moreover, this effect was independent of whether the overshadowed CS was fixed or variable. The findings presented here are consistent with the idea that the strength of the association between CS and unconditioned stimulus (US) is, in part, determined by the temporal distribution form of the CS. These results are discussed in terms of time-accumulation and trial-based theories of conditioning and timing.

SSCC TD: a serial and simultaneous configural-cue compound stimuli representation for temporal difference learning.

This paper presents a novel representational framework for the Temporal Difference (TD) model of learning, which allows the computation of configural stimuli--cumulative compounds of stimuli that generate perceptual emergents known as configural cues. This Simultaneous and Serial Configural-cue Compound Stimuli Temporal Difference model (SSCC TD) can model both simultaneous and serial stimulus compounds, as well as compounds including the experimental context. This modification significantly broadens the range of phenomena which the TD paradigm can explain, and allows it to predict phenomena which traditional TD solutions cannot, particularly effects that depend on compound stimuli functioning as a whole, such as pattern learning and serial structural discriminations, and context-related effects.

Information gathering during contests: the relationship between lateralisation and contestant behaviour during fallow deer fights.

One class of model relating to animal contest behaviour assumes that individuals gather information concerning their opponents' competitive ability; these models argue that such a process allows contestants to avoid engaging in dangerous fighting behaviour with a superior opponent. The brain hemispheres of vertebrates are lateralised in that they are specialised for processing different type of information. Within the context of the current study, we might expect that lateralisation would play a role in facilitating the assessment of opponent quality; nevertheless, the degree of lateralisation shown by individuals can vary suggesting that contest behaviour might also vary based on the ability to process information about competitor quality. The current study tests this hypothesis by predicting that the duration that individuals engage in fighting and the rate of aggressive contest actions should decrease as lateralisation increases. There was a positive relationship between two laterality indices and the duration spent in antler contact; thus lateralised individuals experienced greater time costs. Further, lateralised individuals also experienced a greater disparity in contest actions: there was a negative relationship between lateralisation and the difference in the mean number of backward pushes achieved during fights. When only opponent signal rate was considered there was no effect of lateralisation, therefore, there is support for a mutual assessment process. These results suggest that information gathering via lateral displays may be disadvantageous to lateralised individuals during escalated fighting.

Dissociations in the effect of delay on object recognition: evidence for an associative model of recognition memory.

Rats were administered 3 versions of an object recognition task: In the spontaneous object recognition task (SOR) animals discriminated between a familiar object and a novel object; in the temporal order task they discriminated between 2 familiar objects, 1 of which had been presented more recently than the other; and, in the object-in-place task, they discriminated among 4 previously presented objects, 2 of which were presented in the same locations as in preexposure and 2 in different but familiar locations. In each task animals were tested at 2 delays (5 min and 2 hr) between the sample and test phases in the SOR and object-in-place task, and between the 2 sample phases in the temporal order task. Performance in the SOR was poorer with the longer delay, whereas in the temporal order task performance improved with delay. There was no effect of delay on object-in-place performance. In addition the performance of animals with neurotoxic lesions of the dorsal hippocampus was selectively impaired in the object-in-place task at the longer delay. These findings are interpreted within the framework of Wagner's (1981) model of memory.

The effect of stimulus distribution form on the acquisition and rate of conditioned responding: implications for theory.

In four experiments rats were conditioned to an auditory conditioned stimulus (conditioned stimulus; CS) that was paired with food, and learning about the CS was compared across two conditions in which the mean duration of the CS was equated. In one, the CS was of a single, fixed duration on every trial, and in the other the CS duration was drawn from an exponential distribution, and hence changed from trial to trial. Higher rates of conditioned responding to the fixed than to the variable stimulus were observed, in both between- (Experiment 1) and within-subject designs (Experiments 2 and 3). Moreover, this difference was maintained when stimuli trained with fixed or variable durations were tested under identical conditions (i.e., with equal numbers of fixed and variable duration trials)-suggesting that the difference could not be attributed to performance effects (Experiment 3). In order to estimate the speed of acquisition of conditioned responding, the scaled cumulative distribution of a Weibull function was fitted to the trial-by-trial response rates for each rat. In the within-subject experiments specific differences in the pattern of acquisition to fixed and variable CS were shown; a somewhat different pattern was found when intertrial interval (ITI) was manipulated (Experiment 4). The implications of these findings for theories of conditioning and timing are discussed.

Dorsal hippocampal involvement in conditioned-response timing and maintenance of temporal information in the absence of the CS.

Involvement of the dorsal hippocampus (DHPC) in conditioned-response timing and maintaining temporal information across time gaps was examined in an appetitive Pavlovian conditioning task, in which rats with sham and DHPC lesions were first conditioned to a 15-s visual cue. After acquisition, the subjects received a series of non-reinforced test trials, on which the visual cue was extended (45 s) and gaps of different duration, 0.5, 2.5, and 7.5 s, interrupted the early portion of the cue. Dorsal hippocampal-lesioned subjects underestimated the target duration of 15 s and showed broader response distributions than the control subjects on the no-gap trials in the first few blocks of test, but the accuracy and precision of their timing reached the level of that of the control subjects by the last block. On the gap trials, the DHPC-lesioned subjects showed greater rightward shifts in response distributions than the control subjects. We discussed these lesion effects in terms of temporal versus non-temporal processing (response inhibition, generalisation decrement, and inhibitory conditioning).

Vocal rate as an assessment process during fallow deer contests.

Two types of model propose that strategic decisions during contests are determined either by (i) a mutual-assessment process or (ii) a self-assessment process. Vocal signals are thought to convey information about the competitive abilities of individuals, the ultimate function of which is a reduction in costs associated with fighting consistent with the principle of mutual assessment. Nevertheless, the limited evidence that male ungulates engage in mutual assessment of vocal rates during dyadic contests has been questioned. Therefore, we examined the vocal rates of winners and losers during escalated dyadic contests between male fallow deer in order to further inform on this issue. Our results showed that winners and losers did not differ in vocal rate. The best model fit that accounted for individual vocal rates included a preponderance of factors related to the opponent indicating that contestants were attending to their opponent during fights. Vocal rate was, therefore, dependent on estimates of opponent quality without reference to self, supporting an 'opponent-only' rather than a mutual assessment process.

Some appetitive procedures for examining associative learning in the mouse: Implications for psychopathology.

There are few demonstrations of basic associative learning phenomena using appetitive procedures in mice. This article describes procedures for obtaining four associative learning phenomena in mice, using an appetitive conditioning procedure in which the reinforcer was delivery of a sucrose pellet, and the conditioned response head entry into the food tray. Experiment 1 demonstrated latent inhibition in a within-subjects procedure. Experiment 2 demonstrates both overshadowing and blocking, and Experiment 3 Pavlovian conditioned inhibition, which was evaluated by both summation and retardation tests. These procedures all have potential relevance to current translational research questions. The specific advantages of using appetitive tasks are discussed.

Overshadowing and stimulus duration.

In 3 experiments, the authors investigated the effect of stimulus duration on overshadowing. Experiments 1 and 2 examined responding to a target conditioned stimulus (CS1) when it was conditioned in compound with a coterminating overshadowing stimulus (CS2) that was longer, shorter, or of the same duration (the long, short, and matched groups, respectively). Equal overshadowing of conditioning to CS1 was obtained in all 3 conditions relative to a control group conditioned to the light alone. There were, however, differences in responding to CS2 as a function of its absolute duration. Experiment 3 examined the contribution of the food-food interval/CS onset-food interval ratio to these findings. In Experiments 1 and 2, the ratio differed for the overshadowing CS but not for the target CS. In Experiment 3, this arrangement was reversed, but the pattern of results remained the same. The implications of these findings for trial-based and real-time models of conditioning are discussed.

Is controllability of imagery related to canoe-slalom performance?

This study investigated the relationship of controllability of mental imagery with canoe-slalom performance. Controllability of mental imagery was assessed by an objective test of mental rotation, the Mental Rotations Test. This test was administered to both elite (n = 19) and intermediate (n = 12) athletes. Predictive validity of the controllability test was supported by a significant correlation between test scores and race rank-order for the elite canoeing group (rs = 0.42, p<.05); however, it did not distinguish elite from intermediate groups (t29 = 0.98, p>.05). Researchers should attempt to evaluate vividness of imagery, controllability of imagery, and accuracy of reference to understand more fully the nature of athletes' imagery.