Effects of experience and object complexity on exploration in garden warblers (Sylvia borin)

Animals are increasingly exposed to novel environmental conditions because of human activities. An organism's ability to cope with these challenges is critical to its survival. The decision of whether or not an animal should investigate environmental changes involves a trade-off between the risk and the potential benefit of a novel resource. Novelty often elicits approach (exploratory) as well as avoidance behaviour in animals, and the extent of each of these behaviours may influence learning about the environment. We formulated and tested several hypotheses to determine how external factors (complexity of a novel stimulus) and internal factors (experience with objects) might influence exploration latencies to touch a novel object based on either cost or benefit considerations. Garden warblers were confronted with a simple and a complex novel object in their familiar cages, and latencies until first touch were measured. Additionally, corticosterone levels before and after the experiment were measured to assess whether differences in baseline levels might explain variations in latency to approach and whether presentation of novel objects is stressful. The complex object was touched significantly later than the simple object. Experienced birds investigated both types of objects later than inexperienced birds. Although birds showed marked approach–avoidance behaviour with the complex object, none of the objects elicited a corticosterone stress response. The results indicate that garden warblers consider both costs and benefits when exploring novel objects, and that the relative influence of costs and benefits varies with external and internal factors.     

The role of antennal hair plates in object-guided tactile orientation of the cockroach (Periplaneta americana)

The searching behavior of blinded cockroaches was examined under unrestrained conditions, in an arena, and on a treadmill. When cockroaches searching in a circular arena touched a stationary object (metal pole) with their antennae, they frequently approached the object more closely, and then climbed up it. Similar orientation behavior was observed in tethered animals in open loop conditions, walking on a Styrofoam ball. In these restrained cockroaches, a single antenna sufficed to distinguish the angular positions of an object, in the horizontal plane (0 degrees, 45 degrees, and 90 degrees). A group of mechanosensitive hairs on the basal segment of the antenna (scapal hair plate) appears to play a major role in antennal object detection in the horizontal plane, as gauged by shaving off these scapal hair plates. In unrestrained cockroaches, shaving the scapal hair plate increased the time needed to approach an object. Under tethered conditions, the ability to turn towards and to establish antennal contact with a test object was significantly impaired.     

Personality assessment in snow leopards (Uncia uncia)

Knowledge of individual personality is a useful tool in animal husbandry and can be used effectively to improve welfare. This study assessed personality in snow leopards (Uncia uncia) by examining their reactions to six novel objects and comparing them to personality assessments based on a survey completed by zookeepers. The objectives were to determine whether these methods could detect differences in personality, including age and sex differences, and to assess whether the two methods yielded comparable results. Both keeper assessments and novel object tests identified age, sex, and individual differences in snow leopards. Five dimensions of personality were found based on keepers' ratings: Active/Vigilant, Curious/Playful, Calm/Self-Assured, Timid/Anxious, and Friendly to Humans. The dimension Active/Vigilant was significantly positively correlated with the number of visits to the object, time spent locomoting, and time spent in exploratory behaviors. Curious/Playful was significantly positively correlated with the number of visits to the object, time spent locomoting, and time spent in exploratory behaviors. However, other dimensions (Calm/Self-Assured, Friendly to Humans, and Timid/Anxious) did not correlate with novel-object test variables and possible explanations for this are discussed. Thus, some of the traits and behaviors were correlated between assessment methods, showing the novel-object test to be useful in assessing an animal's personality should a keeper be unable to, or to support a keeper's assessment.     

Response to novel object in Wistar and wild-type (WWCPS) rats

Laboratory rats react to a novel object with a rapid redirection of exploratory behaviour towards the source of the novelty and a subsequent decline of this neotic preference with repeated object exposure. Studies with wild Rattus norvegicus have shown that a novel object in a familiar cage results in avoidance reactions (neophobia) in those animals, but it is also well established that the wild R. norvegicus show strong aversive reactivity to a variety of high intensity stimuli. In this study we aimed to create low-stress conditions enabling the comparison of spontaneous exploratory behaviour directed at a novel object in male (age = 80 days) “wild-type” WWCPS rats (n = 21; fourth generation bred in captivity) and Wistar (n = 24) rats. The study involved repeated placing of individual animals in the experimental chamber for 15 (6 min) trials on consecutive days. On the 11th day the novel object was introduced. Animals were tested in darkness and without human presence. Under these conditions neither WWCPS nor Wistar have shown behavioural signs of high emotional arousal, both lines have shown comparable general levels of experimental cage exploration and the positive new object exploratory reaction was observed only in Wistar rats.     

Exploration as an indicator of good welfare in beef bulls: An attempt to develop a test for on-farm assessment

Exploration appears to be carried out for its own purpose due to being self-rewarding. Furthermore, it increases environmental certainty with the effect of increased predictability and control of the environment. We, therefore, supposed that exploratory behaviour is associated with positive emotions and could be a valuable tool in the assessment of animal welfare from the positive side. Since assessing the level of exploration in daily life is time consuming, the goal of our study was to develop a feasible on-farm test to estimate the daily exploratory level of beef bulls. This was based on the hypothesis that bulls in barren environments (here: pens with fully slatted floors) will explore an unknown object more than those housed under more enriched conditions (here: littered pens). A novel object test at the feed rack over a 1-h period was carried out in 64 groups of fattening bulls on farms in Germany and Austria. In order to evaluate sensitivity of the test half of the barren pens in Germany were slightly enriched after these tests and bulls tested again after 1 week. During the first three quarters of the test hour, significantly more barren-housed bulls were occupied with the objects than animals in littered pens. This was predominantly due to more licking/chewing of the novel object. However, the differences in exploration levels were only slight. Moreover, the test was not sensitive to the slight short-term change in stimulation of the bulls following the simple enrichment. The motivational basis for increased exploration of the novel object by licking or chewing in bulls kept in barren conditions is not clear-cut. While it might reflect that these bulls gain less positive emotions from intrinsic exploration in their daily life, nutritional aspects may also play a role. While further investigations should aim to identify the motivational basis of occupation with the novel object and relate true daily exploration levels to test results, for the on-farm assessment of good animal welfare the test appears not to be promising in its current form.     

Puumala hantavirus infection alters the odour attractiveness of its reservoir host

The random-mixing assumptions of many parasite-transmission models are challenged if healthy individuals can alter their behaviour to reduce their risk of infection. Some pathogens reduce the attractiveness of their hosts’ excretions, for example, potentially altering contact rates and thus the predicted force of infection for pathogens transmissible by contact with excretions. For bank voles (Myodes glareolus), contact with contaminated urine is an important route of transmission for Puumala hantavirus (PUUV); however, it is not known whether PUUV infection changes the voles’ urinary odours or their attractiveness. Here, we use a Y-maze to test whether PUUV infection alters the attractiveness of male bank voles’ urine. We presented wild-caught PUUV-free male and female bank voles with PUUV-infected conspecific urine, uninfected urine and a water control, and measured the relative and absolute latency to first visit, number of visits, and total time bank voles spent investigating each treatment over 30 min. PUUV infection significantly altered the bank voles’ initial response to conspecific urine, with fewer visits and less time spent close to infected urine relative to uninfected urine, and less total time spent near the infected urine than the uninfected urine or control. These strong preferences weakened over the 30-min trial, however, partly due to a general decline in male activity, and there were no absolute differences between the treatments overall. This suggests that PUUV infection does change the attractiveness of bank vole urine to conspecifics, and we discuss the implications of these results for random-mixing assumptions.     

Comparisons of toxic baits for controlling the cockroach, Blattella germanica: attractiveness and feeding stimulation

Female German cockroaches, Blattella germanica (L.) (Dictyoptera: Blattellidae) and nymphs (stages 1, 3 and 6) were used to test the attractiveness and the feeding stimulation of several commercial gel formulations of insecticide food baits in laboratory binary choice tests. The four toxic gel baits tested were: 'Avert' (0.05% abamectin=avermectin B1), 'Drax' (33.3% boric acid), 'Goliath' (0.05% fipronil) and 'Maxforce' (2.15% hydramethylnon). The attractiveness of a bait was estimated by the number of cockroaches that chose it first, and by their tendency to change to it secondarily. Feeding stimulation was estimated by the mean duration of feeding bouts and total duration of food intake. Variability of these factors was assessed in relation to age of cockroaches and to age of baits. With a few exceptions, cockroach age did not affect performance in any of these tests. Attractiveness to B. germanica ranked 'Goliath' gel higher than 'Avert', 'Drax' and 'Maxforce' gels, whether or not the active ingredient fipronil was present in 'Goliath' gel. Feeding stimulation ranked 'Avert' and 'Maxforce' gels higher than 'Goliath' gel, and 'Drax' gel was inferior. With ageing up to 3 months, the feeding stimulation power of 'Goliath' and 'Maxforce' gels increased, whereas that of 'Avert' fluctuated. Assuming that all types of bait effectively kill cockroaches once ingested, these results illustrate the importance of behavioural observations on bait efficiency and show that ranking of these different food baits varies in relation to the parameter analysed.     

Detection of genetic variants affecting cattle behaviour and their impact on milk production: a genome‐wide association study

Behaviour traits of cattle have been reported to affect important production traits, such as meat quality and milk performance as well as reproduction and health. Genetic predisposition is, together with environmental stimuli, undoubtedly involved in the development of behaviour phenotypes. Underlying molecular mechanisms affecting behaviour in general and behaviour and productions traits in particular still have to be studied in detail. Therefore, we performed a genome‐wide association study in an F₂ Charolais × German Holstein cross‐breed population to identify genetic variants that affect behaviour‐related traits assessed in an open‐field and novel‐object test and analysed their putative impact on milk performance. Of 37 201 tested single nucleotide polymorphism (SNPs), four showed a genome‐wide and 37 a chromosome‐wide significant association with behaviour traits assessed in both tests. Nine of the SNPs that were associated with behaviour traits likewise showed a nominal significant association with milk performance traits. On chromosomes 14 and 29, six SNPs were identified to be associated with exploratory behaviour and inactivity during the novel‐object test as well as with milk yield traits. Least squares means for behaviour and milk performance traits for these SNPs revealed that genotypes associated with higher inactivity and less exploratory behaviour promote higher milk yields. Whether these results are due to molecular mechanisms simultaneously affecting behaviour and milk performance or due to a behaviour predisposition, which causes indirect effects on milk performance by influencing individual reactivity, needs further investigation.     

Exploratory behaviour of goldfish Carassius auratus

The exploratory locomotor behaviour of twenty-one naïve goldfish in a homogeneous open field during 20-hr periods was monitored and analysed into spatial and temporal characteristics. The fish did not randomly explore the entire field. Locomotion is organized spatially and temporally. The fish moves about in one specific area during a specific time after which it displaces the centre of its activity to another area. The time spent in earch area is a function of rate of activity, expressed by a linear equation and, during the initial 4 hr, the elapsed time in the novel environment (novelty content). The area-by-area exploration was never replaced by random exploration throughout the environment during the 20-hr periods of observation.     

The role of model female quality in the mate choice copying behaviour of sailfin mollies

Female mate choice copying is a socially mediated mate choice behaviour, in which a male's attractiveness to females increases if he was previously chosen by another female as a mate. Although copying has been demonstrated in numerous species, little is known about the specific benefits it confers to copying females. Here we demonstrate that the mate choice behaviour of female sailfin mollies (Poecilia latipinna) is influenced by the phenotypic quality of model females with whom males are observed consorting. Test females choosing between two males of similar body length were found to significantly increase time spent with previously non-preferred males after having observed them with a relatively high-quality female. Conversely, females were found to significantly decrease time spent with previously preferred males after having observed them with a relatively low-quality female. Female mate choice copying might be maintained by selection based on the heuristic value it provides females choosing between males whose quality differences are not easily distinguishable.     

The influence of olfaction on exploratory behaviour in the paradise fish (Macropodus opercularis L.)

We studied the role of olfaction on the exploratory behaviour of the paradise fish. Intact animals showed habituation of exploratory behaviour toward a heterospecific fish after five consecutive encounters. Fish with olfactory nerves destroyed spent significantly longer time with exploration compared to the control even at the fifth encounter. We suggest that olfactory inputs have a strong influence on exploratory behaviour.     

Effects of the protozoan parasite Sarcocystis rauschorum on open-field behaviour of its intermediate vertebrate host, Dicrostonyx richardsoni

Behaviour and activity levels were measured in varying lemmings experimentally infected with the heteroxenous parasite, Sarcocystis rauschorum to test the hypothesis that the parasite alters behaviour of this intermediate host and thereby increases probability of transmission to the definitive host, the snowy owl (Nyctea scandiaca). Measures of short-term activity levels on a running wheel indicated no effect of the parasite, either directly, or indirectly as a result of illness. We observed behaviour of infected lemmings placed in an "open field" (arena). Lemmings would increase their susceptibility to predators if they spent more time away from cover, used crypsis (stationary postures) less, spent more time exploring (especially in unfamiliar areas), or responded inappropriately to threats from predators. We found that only exploratory activity showed significant change after infection. The frequency of exploratory activity increased and became disassociated from the usual fear response. This may increase the lemmings' susceptibility to aerial predation. The mechanism for this effect is unknown, but neurological lesions have been observed. The examination of the modes of transmission of the S. rauschorum parasite within lemming populations and of a possible fecundity compensation strategy adopted by the lemmings, and their relevance to population control, are suggested as areas for future study.     

Bearded reedlings adjust their pair-bond behaviour in relation to the sex and attractiveness of unpaired conspecifics

An individual's investment in mating or keeping a pair bond intact may be influenced not only by the attractiveness of its current mate, but also by that of other potential mates. In this study, we investigated the effect of relative attractiveness on pair-bond behaviour in bearded reedlings, Panurus biarmicus. We showed that mate attractiveness, in terms of beard length in males and tail length in females, influenced courtship behaviour when the pair was kept isolated. In the presence of a conspecific, contact initiations within a pair increased. This increment was mainly related to the sex of the unpaired conspecific, however, and less to differences in attractiveness between the current partner and the unpaired conspecific. Female contact initiations towards potential extra mates were independent of male attractiveness, whereas male contact behaviour was significantly influenced by female attractiveness. However, females displayed more contact initiations to their current mate when they were less attractive than the unpaired females. Males decreased their overtures towards other females with increasing attractiveness of their current mates. Overall, our results suggested that, when there was a risk of losing their mate, bearded reedlings adjust their pair-bond investment mainly in response to the presence or absence of a competitor, and fine-tune investment to a lesser extent in response to the attractiveness of that potential competitor.     

The effect of preweaning and postweaning housing on the behaviour of the laboratory mouse (Mus musculus)

Standard housing for laboratory mice severely restricts natural behaviour and the control that the animal has over its environment. Providing the cage with objects is a method that has been used to both increase environmental complexity, promote the performance of natural behaviour and provide greater controllability for the animal. This method of furnishing cages has mostly been studied in adult animals, and little is known about the influence that the preweaning environment has on the behaviour of mice as adults. This study aimed to investigate the effects on mice behaviour of preweaning and postweaning housing environment. In this experiment, 64 pairs of animals of the strain C57BL/6J were used. Half of the animals were born and reared until weaning in standard cages and the other half in cages twice the size of the standard and furnished with nesting material, a cardboard tube, a PVC nest box and a wooden chewblock. After weaning, half the animals in each group were changed to the other type of cage, whereas the other half remained in the same environment; in both cases they were kept in single-sex pairs of littermates. Behaviour during the dark, active period was studied through video recordings. We found no main effects of preweaning environment on behaviour; however, mice moved from furnished to standard cages at weaning showed a decrease in inactive behaviour at four weeks of age. Mice housed after weaning in standard cages spent less time inactive, and more time engaging in activities like feeding and drinking, self-grooming and allogrooming. A sex difference was also found, in that females showed a greater performance of exploratory behaviour as well as a greater prevalence of stereotypies. The use of different objects and locations within the furnished cage was also analysed at both ages. Results show that at eight weeks of age mice spent more time at the top of the cage, and that the use of the nest box (although not for resting) increased between four and eight weeks. Mice were found to use the nest box as a nesting site/sleeping place only at age four weeks, whereas they always used the nesting material for sleeping.     

Progesterone can enhance consolidation and/or performance in spatial, object and working memory tasks in Long-Evans rats

Progesterone has a ubiquitous role in reproduction and fitness and may influence cognitive performance. We examined the effects of administration of progesterone (a regimen that facilitates sexual behaviour) on consolidation of complex information in Long-Evans rats, Rattus norvegicus, that may be relevant for social engagement. We also examined the effects of subcutaneous progesterone administration (4 mg/kg versus oil vehicle placebo) on memory of ovariectomized rats during various cognitive tasks. Ovariectomized rats that received progesterone, versus the vehicle, immediately post-training were better able to find a hidden platform in the water maze. In a recognition task, rats that received progesterone spent more time in the novel arm of the Y-maze task than rats that received the vehicle. Ovariectomized rats that received progesterone immediately after training spent significantly more time exploring a novel object (compared to a familiar object) than did vehicle-administered rats. When socially relevant stimuli (i.e. objects with the scent of familiar or novel conspecifics) were used in the social cognition task, ovariectomized rats that received progesterone spent more time exploring the object with the novel conspecifics' scent than did vehicle-administered rats. Pairing of progesterone, but not the vehicle, conditioned a place preference to the originally nonpreferred side of the conditioning chamber. We found no significant differences in motor activity measures in these tasks due to progesterone treatment. These results suggest that progesterone's effects to improve cognitive processes with nonsocial and socially relevant stimuli, as well as have reinforcing effects, may underlie some of its salient effects on reproduction-related behaviours.     

Behavioural development of conspecific odour preferences in bank voles,Clethrionomys glareolus

Biological odours of conspecifics are known to have strong influences on behavioural interaction in bank voles Clethrionomys glareolus. This experiment tested two hypotheses. (1) Olfactory cues from familiar and unfamiliar mature opposite-sex conspecifics differ in their attractiveness to males and females, and their behavioural reactions change with age. (2) A genetically based mechanism is involved in female recognition of kin.In a two-choice preference test, prepubertal males and females were more attracted to familiar than to unfamiliar odours of opposite-sex conspecifics, as manifested by more time spent sniffing familiar voles. As the young reached sexual maturity they shifted their odour preferences. Mature males and females preferred the novel odour of unrelated opposite-sex conspecifics to that of relatives. The results of experiments testing the second hypothesis indicate that females use a genetically based mechanism to recognise their kin. Young and mature females were able to recognise the odour of their biological but socially unknown fathers, and showed the same pattern of behaviour as females in previous experiments.The possible biological functions of kin recognition in bank voles are discussed.     

Effects of Anesthetic Ketamine on Anxiety-Like Behaviour in Rats

There is scarce information regarding the effects of anesthetic doses of the non-competitive N-methyl-d-aspartate receptor antagonist ketamine on anxiety. The current study evaluated the acute effects of intraperitoneally (i.p.) administered anesthetic ketamine (100 mg/kg) i.p. on anxiety in rats. For this purpose, the light/dark and the open field tests were utilized. The effects of anesthetic ketamine on motility were also examined using a motility cage. In the light/dark test, anesthetic ketamine, administered 24 h before testing reduced the number of transitions between the light and dark compartments and the time spent in the light compartment in the rats compared with their control cohorts. In addition, ketamine was found to exert a depressive effect on rats’ motility. In the open field test, animals treated with anesthetic ketamine 24 h before testing spent essentially no time in the central area of the apparatus, decreased horizontal ambulatory activity, and preserved to a certain extent their exploratory behaviour compared to their control counterparts. The results suggest that, in spite of its hypokinetic effect, a single anesthetic ketamine administration apparently induces an anxiety-like state, while largely preserving exploratory behaviour in the rat. These effects were time-dependent they since they were extinguished when testing was carried out 48 h after anesthetic ketamine administration.

     

The exploratory behaviour of rats in the hole-board apparatus: Is head-dipping a valid measure of neophilia?

The exploratory behaviour of laboratory rodents is of interest within a number of areas of behavioural pharmacology. However, how best to measure exploratory behaviour in rodents remains a contentious issue. Many unconditioned tests, such as the open field, potentially confound general locomotor activity with exploration. The hole-board apparatus appears to avoid this confound, as head-dipping into holes in the floor is assumed to be a valid measure of the subject's attraction towards novelty (neophilia). This study aimed to investigate whether head-dipping should be considered a valid measure of neophilia by comparing performance of adult male and female Lister hooded rats on the hole-board task (a) over repeated sessions and (b) when novel objects were absent or present underneath the holes. The results show that head-dipping initially decreased across repeated exposures, while time spent in the aversive central area increased. No change in head-dipping was seen in response to objects being placed underneath the holes. Rather than being a measure of neophilia, these results support the hypothesis that head-dipping represents an escape response, which declines as the subject becomes less fearful. These results are compared with previous studies of repeated exposure to other novel environments.     

The Significance of Ecological Factors for Exploration and Neophobia in Parrots

Exploratory behaviour plays an important role in most animals for gathering information about their environment. If it constitutes an adaptation to different environmental conditions exploratory behaviour should differ between species. This has been tested with several hypotheses.
Sixty-one parrot species (Psittacidae) from eight tribes with different diets and habitat preferences were investigated in aviaries. Two tests were carried out. First, a novel object (wooden ring) in the familiar aviary was presented on two test days in the exploration test. Latencies until first contact with the object and the duration of exploration were recorded. Secondly, in the neophobia test, novel objects were placed beside the feeding dish and latencies until first food intake were recorded.
The exploration and neophobia data were related to 12 (13) ecological variables using multiple regression analyses. Phylogenetic relationships were considered.
Species that inhabit complex habitats, such as forest edges, or that feed on buds or species from islands showed the shortest latencies in the exploration test. In contrast, long latencies were related to a diet including a great amount of seeds and/or flowers. The longest duration of exploration occurred in species eating nuts or originating from islands, whereas short durations were related to feeding on seeds. Neophobia was positively related to a diet consisting of insects, and negatively to a diet of leaves. There was no relationship between measures of exploration and neophobia. Exploration and neophobia seem to be tightly related to the ecology of a species.     

Genetic correlation between resting metabolic rate and exploratory behaviour in deer mice (Peromyscus maniculatus)

According to the 'pace-of-life' syndrome hypothesis, differences in resting metabolic rate (RMR) should be genetically associated with exploratory behaviour. A large number of studies reported significant heritability for both RMR and exploratory behaviour, but the genetic correlation between the two has yet to be documented. We used a quantitative genetic approach to decompose the phenotypic (co)variance of several metabolic and behavioural measures into components of additive genetic, common environment and permanent environment variance in captive deer mice. We found significant additive genetic variance for two mass-independent metabolic measures (RMR and the average metabolic rate throughout the respirometry run) and two behavioural measures (time spent in centre and distance moved in a novel environment). We also detected positive additive genetic correlation between mass-independent RMR and distance moved (r(A) = 0.78 ± 0.23). Our results suggest that RMR and exploratory behaviour are functionally integrated traits in deer mice, providing empirical support for one of the connections within the pace-of-life syndrome hypothesis.