Effects of social influence on neonatal approach responses of Gallus chicks

Although most laboratory studies of imprinting rear and test subjects individually, in the natural setting birds are imprinted in groups. In order to approximate and evaluate naturally occuring conditions, the present study compared the approach responses of chicks reared and trained alone to those reared and trained in pairs to an audio-visual imprinting stimulus. The responses of both groups were virtually identical during the presence of the imprinting stimulus. However, under conditions of experimental extinction, the paired trained birds (now tested alone) were more resistant to extinction than their isolated counterparts, thus suggesting that birds with prior social experience are more strongly imprinted than has been previously reported.     

Embryonic Pituitary Adrenal Axis, Behavior Development and Domestication in Birds

ACTH and corticosterone exert opposite effects on the approachand imprinting behavior of newly hatched ducklings. Wild mallardand domesticated Pekin ducklings differ in the early posthatchperiod in both plasma corticosterone levels and approach/avoidancebehaviors. Injection of Pekin duckling embryos with pituitary—adrenocorticalhormones alters both later adrenal function and certain aspectsof posthatch behavior. These birds have behavioral and hormonalcharacteristics which resemble those of wild mallards. The hypothesisthat behavioral differences in wild and domesticated ducklingsresult from a higher level of pituitary adrenal function inthe wild embryo is explored. Although adrenocortical functionchanges during domestication in many species, evidence thatthe hormonal changes mediate the concomitant changes in approachand avoidance behavior remains inconclusive. Factors which causeadrenal function and early behaviors to differ in wild and domesticatedgenotypes must be sought in the gene action during embryonicdevelopment. Since imprinting behavior is modulated by pituitary—adrenalhormones, any factor which affects post—hatch adrenalfunction may potentially affect imprinting. Later behavior developmentin the adult is strongly dependent on neonatal experiences;and, therefore, hormonal modulation of early imprinting behaviormay constitute an important determinant of adult social behavior.     

Getting better all the time: improving preference scores reflect increases in the strength of filial imprinting

In the investigation of the neural mechanisms of filial imprinting, neurochemical measures are often correlated with preference score (PS): approach activity to the training stimulus/total approach in a test. In a previous study, domestic chicks, Gallus gallus domesticus, that had a PS under 65% were classed as 'poor learners' and those with a PS greater than 65% were called 'good learners'. We tested the effects of continued imprinting training in chicks from these two categories. After exposure to an imprinting stimulus for 30 min on day 1 after hatching, preferences were tested and then the chicks were exposed to the same stimulus for either 90 min, 3 h or 4 h on the next day, followed by a second preference test. In all these experiments there was a significant improvement in mean PS in the 'poor learners' between the first and second test, such that these chicks acquired a significant mean preference for the training stimulus. There was no such improvement in chicks that did not receive further training on day 2. When absolute approach was analysed, there was no significant difference between 'poor learners' and 'good learners' at the second test, after 4 h of retraining. Overall, mean preference scores increased with length of training. These results suggest that 'poor learners' are better characterized as 'slow learners', and that their initially low PS is not caused by, for example, a lack of motivation to express a preference. Preference scores reflect the strength of learning during imprinting. Copyright 2000 The Association for the Study of Animal Behaviour.     

Imprinting in ducklings: variability among individuals and responses

Duck embryos show a highly variable heart rate, which remains variable, though to a lesser degree, upon hatching. After hatching, behavioral responses of ducklings in an imprinting paradigm are also variable both between ducklings, and between particular responses (behavioral inter se and behavioral vs physiological). Nonetheless, responses, when viewed multi-dimensionally, do appear to cluster, suggesting that there exist a variety of discrete developmental ‘types’, even among an inbred strain of ducklings.     

Neurotransmitter Release from the Medial Hyperstriatum Ventrale of the Chick Forebrain Accompanying Filial Imprinting Behavior,Measured by In Vivo Microdialysis

The imprinting behavior of chicks was quantified as a preference score (correct response ratio) achieved in a running wheel apparatus. A total of 249 chicks were exposed to an imprinting stimulus and tested for stimulus-approaching behavior. The chicks were then classified as good learners (imprinted), poor learners (non-imprinted) and a gray-zone group, those were 46%, 31% and 23% of the total chicks respectively. Using the classified chicks, the acetylcholine (ACh) and glutamate releases from the medial hyperstriatum ventrale (MHV) of the chick forebrains were determined by in vivo microdialysis. The non-imprinted chicks were used as yoked controls. Increases of ACh and glutamate released were observed in the imprinted chicks during exposure to the imprinting stimulus, whereas there were no changes in the release of these neurotransmitters in the non-imprinted chicks during the imprinting exposure. These results might be indicated that cholinergic and glutamatergic synapses which are newly formed as functioning synapses with imprinting stimulus in the MHV are involved in the performance of imprinting behavior.     

Neurochemical studies on imprinting behavior in chick and duckling

The effect of lesiens in the parts of medial hyperstriatum ventrale (MHV) or lateral neostriatum (LN) on impriting behavior was examined. Bilateral lesions in MHV severely impaired imprinting behavior in ducklings, but lesions in LN did not impair such behavior. On the other hand, the birds lesioned in MHV still had brightness discriminative learning ability with perfect performance of visuomotor coordination. The neurotransmitter candidates controlling imprinting behavior in MHV were investigated using blockers of neurotransmission. 6-Hydroxydopamine (6-OHDA) and haloperidole injected into MHV did not have specific effect on imprinting behavior although catecholamine contents in MHV was clearly decreased and locomotor activity was strongly suppressed. Atropine injection into MHV caused significant impairment of following behavior, but locomotor activity was not affected. The injection of atropine into other region did not have any effect. -Bungarotoxin showed no effect on either imprinting or locomotor activity. Kainic acid injection into MHV caused the decrease of glutamate content in the MHV region and it reached maximum at 24 hours after injection. However, imprinting behavior was already impaired at 3 hours after injection, but at 24 hours it had been recovered. A same effect was observed by glutamate injection. The effect of protein synthesis inhibitor on the aquisition process of imprinting was investigated. The aquisition of imprinting was significantly impaired by cycloheximide (CHX) injection into MHV within 2 hours after the first exposure to the imprinting stimulus. When CHX was given 24 hours after, imprinting behavior was not affected.     

Distress call alternation in peking ducklings (Anas platyrhynchos)

Peking ducklings (Anas platyrhynchos) were found to alternate their distress calls with those of conspecifics. This alternation response consisted of two components: a strong tendency to suppress vocalizing during the conspecific's calls and a weaker tendency to increase vocalizing (above the level given by birds tested in auditory isolation) following those calls. Similar results were obtained whether pairs of ducklings were tested or whether individual ducklings were tested with tape-recorded calls. Further, visual contact with the conspecific, or a mirror image, although not necessary for eliciting an alternation response, did have a significant facilitative effect on this response. This latter effect may have been due to the general arousal-reducing properties of the visual stimulus. Finally, in contrast to the strong response of ducklings to conspecific distress calls, they displayed a much weaker tendency to alternate their calls with (a) clicking sounds and 7.0-kHz tones with the same temporal patterning and intensity as the taped distress calls, (b) 3.8-kHz tones that additionally matched the dominant frequency of the taped calls, and (c) synthetic distress calls that also approximated the frequency modulation of the taped calls. These results indicate that the distress call alternation response is not simply an artifact of a more general orienting response in which ducklings inhibit their own vocalizations whenever they hear any salient auditory stimulus. Rather, the response appears to be quite specifically triggered by the sound of the sibling calls. The present results stand in contrast to the common assertion that young birds largely ignore the distress calls of siblings. These results are, however, consistent with recent literature emphasizing the importance of sibling interactions in the analysis of mallard early social attachments.     

Development of visual species identification in ducklings: What is the role of imprinting?

The phenomenon of imprinting (a rapid form of exposure learning) is widely presumed to underlie the formation of normal, species-typical social preferences in precocial birds. To determine whether this is in fact the case, 24-h-old domestic and semi-wild mallard ducklings (Anas platyrhynchos) were allowed to follow a stuffed hen of one of three sympatric waterfowl species for 20 min. The models used were mallard, redhead (Aythya americana), and pintail (Anas acuta) hens. When later tested for their preference for the familiar hen in simultaneous choice tests with one of the other two stuffed models, only those birds trained with the Mallard and tested with the Mallard and Pintail models (designated the Mallard versus Pintail group) showed a preference for the familiar model. (That preference was shown by both domestic and semi-wild ducklings.) In none of the other three groups (Mallard versus Redhead, Redhead versus Mallard, and Pintail versus Mallard) was the imprinting procedure effective in producing a preference for the familiar model. When other ducklings were similarly trained with either the Mallard model, a red-and-white-striped box, or a green styrofoam ball, a preference for the familiar model was found in all four groups (Mallard versus Red Box, Red Box versus Mallard, Red Box versus Green Ball, and Green Ball versus Red Box). Increasing the length of the training period from 20 min to 2 h and to 24 h did not produce a preference for the familiar Mallard over the unfamiliar Redhead. These results raise some doubt that imprinting as currently conceived is the behavioural mechanism of visual species identification as it occurs in nature.     

Stimulus Movement,Hen Behaviour and Filial Imprinting in Japanese Quail (Coturnix coturnix japonica)

A living hen is a very attractive imprinting stimulus compared to artificial stimuli. The present study examined to what extent this attractiveness is influenced by the behaviour of the hen. The effectiveness of a living quail hen as an imprinting stimulus was compared with that of a moving stuffed hen, a non-moving stuffed hen and an empty cage. Naive quail chicks were exposed to one of these stimuli for 2 h. Both during and after this period chicks exposed to the living hen showed the strongest attachment to the stimulus, next came chicks exposed to a moving stuffed hen, while chicks exposed to a non-moving stuffed hen showed no indication of any attachment, i.e. they behaved in the same way as chicks exposed to an empty cage. Also, among chicks exposed to the living hen a correlation was present between the strength of the attachment to the hen and the behaviour of this hen during the exposure. In particular pecking and behaviour directed at the chick seemed to stimulate the development of a filial bond. Together, these findings indicate that a living hen is attractive not just because she moved, but also because of specific qualities of this movement. The behaviour of the living hen differed in various ways from the movement of the moving stuffed hen. The influence of these differences on the filial imprinting process is discussed and compared with similar findings for sexual imprinting and song learning.     

Development of spatial memory in occlusion-experienced domestic chicks

At around day 11 of life, domestic chicks show a tendency to move out of sight of their mother before returning and regaining social and visual contact. We conducted a series of experiments to investigate the role of this voluntary ‘out-of-sight’ behaviour on the development of spatial memory in young chicks. We compared the behaviour of chicks that were reared in environments that provided opportunities to move out of sight of an imprinting stimulus (occlusion-experienced chicks) with the behaviour of chicks that were given minimal occlusion experience (controls). As in natural conditions, out-of-sight behaviour peaked on day 11. When chicks were released into larger pens at 14 days of age, occlusion-experienced chicks walked more than control chicks, but otherwise showed similar degrees of dispersal. Occlusion-experienced chicks tended to show better (although not significant, P=0.09) retrieval of a visually displaced imprinting stimulus than control chicks. Time spent out of sight in the rearing pens was negatively related to the number of orientation errors in a detour test. Occlusion-experienced chicks also tended to make fewer orientation errors in the first trial (P=0.07) and in subsequent trials (P=0.05). In contrast, experimentally manipulating the amount of time that chicks were out of sight of an imprinting stimulus (by confining the chicks) had no effect on their performance in displacement or detour tests. The results presented here suggest that active experience of occlusion around day 11 improved egocentric orientation towards an out-of-sight goal, supporting the hypothesis that enrichment-induced behavioural changes are dependent on the interaction with objects.     

Aging and error processing: age related increase in the variability of the error-negativity is not accompanied by increase in response variability

Background

Several studies report an amplitude reduction of the error negativity (Ne or ERN), an event-related potential occurring after erroneous responses, in older participants. In earlier studies it was shown that the Ne can be explained by a single independent component. In the present study we aimed to investigate whether the Ne reduction usually found in older subjects is due to an altered component structure, i.e., a true alteration in response monitoring in older subjects.

Methodology/Principal Findings

Two age groups conducted two tasks with different stimulus response mappings and task difficulty. Both groups received fully balanced speed or accuracy instructions and an individually adapted deadline in both tasks. Event-related potentials, Independent Component analysis of EEG-data and between trial variability of the Ne were combined with analysis of error rates, coefficients of variation of RT-data and ex-Gaussian fittings to reaction times. The Ne was examined by means of ICA and PCA, yielding a prominent independent component on error trials, the Ne-IC. The Ne-IC was smaller in the older than the younger subjects for both speed and accuracy instructions. Also, the Ne-IC contributed to a much lesser extent to the Ne in older than in younger subjects. RT distribution parameters were not related to Ne/ERP-variability.

Conclusions/Significance

The results show a genuine reduction as well as a different component structure of the Ne in older compared to young subjects. This reduction is not reflected in behaviour, apart from a general slowing of older participants. Also, the Ne decline in the elderly is not due to speed accuracy trade-off. Hence, the results indicate that older subjects can compensate the reduction in control reflected in the reduced Ne, at least in simple tasks that induce reaction slips.     

雏鸡听觉印记学习

雏鸡听觉印记学习是研究新生个体在早期发育阶段学习与记忆形成的神经机制的良好实验模型.听觉刺激对印记学习有很好的强化作用,作用效果依赖于频率、强度和持续时间等声音特征.从声音的选择、听觉刺激所引起的分子水平和组织水平改变等方面综述雏鸡听觉印记学习的研究进展.     

Neuronal plasticity and multisensory integration in filial imprinting

Many organisms sample their environment through multiple sensory systems and the integration of multisensory information enhances learning. However, the mechanisms underlying multisensory memory formation and their similarity to unisensory mechanisms remain unclear. Filial imprinting is one example in which experience is multisensory, and the mechanisms of unisensory neuronal plasticity are well established. We investigated the storage of audiovisual information through experience by comparing the activity of neurons in the intermediate and medial mesopallium of imprinted and naïve domestic chicks (Gallus gallus domesticus) in response to an audiovisual imprinting stimulus and novel object and their auditory and visual components. We find that imprinting enhanced the mean response magnitude of neurons to unisensory but not multisensory stimuli. Furthermore, imprinting enhanced responses to incongruent audiovisual stimuli comprised of mismatched auditory and visual components. Our results suggest that the effects of imprinting on the unisensory and multisensory responsiveness of IMM neurons differ and that IMM neurons may function to detect unexpected deviations from the audiovisual imprinting stimulus.     

The role of frequency modulation in controlling the response of mallard ducklings (Anas platyrhynchos) to conspecific distress calls

Three experiments examined the response of mallard ducklings to conspecific distress calls. In experiment 1, a synthetic call was constructed from a single distress note, by recording it on a digital disc and then using a software routine to regularly repeat this stored note with the average period (267 ms) of the original call. Ducklings were then tested for their tendency to inhibit their own distress vocalizations in response to either this synthetic call, the original call, or a constant-frequency tone mimic; they showed a significantly stronger inhibitory response to the synthetic call than to the tone mimic, but an even stronger response to the original call. The former result indicated that the synthetic call was an effective stimulus, and suggested that some aspect of the frequency modulation found in distress notes is required to evoke the normal duckling inhibitory response. The latter result further suggested that the acoustic variability found in the original call, but not the synthetic call, may be of importance for controlling duckling behaviour. Experiment 2 demonstrated that several other minor differences between the synthetic call and original call (number of notes per call and the digital recording of the synthetic call) could not account for the difference in duckling response to these two calls. Finally, in experiment 3, two additional synthetic stimuli were constructed from the digitized note, by first excising either the initial (front-chop) or terminal (rear-chop) frequency modulation found in each distress note, then creating amplitude envelopes for these new notes similar to that of the unaltered note, and finally repeating these notes to form calls with the same note period as the original call. The ducklings tested with the synthetic normal-note call and synthetic front-chop call showed a significantly stronger inhibitory response than the ducklings tested with the synthetic rear-chop call and the tone mimic. These results indicate that the terminal descending frequency sweep is an important feature of distress notes for triggering the response of ducklings to conspecific distress calls.     

A statistical framework for genome-wide scanning and testing of imprinted quantitative trait loci

Non-equivalent expression of alleles at a locus results in genomic imprinting. In this article, a statistical framework for genome-wide scanning and testing of imprinted quantitative trait loci (iQTL) underlying complex traits is developed based on experimental crosses of inbred line species in backcross populations. The joint likelihood function is composed of four component likelihood functions with each of them derived from one of four backcross families. The proposed approach models genomic imprinting effect as a probability measure with which one can test the degree of imprinting. Simulation results show that the model is robust for identifying iQTL with various degree of imprinting ranging from no imprinting, partial imprinting to complete imprinting. Under various simulation scenarios, the proposed model shows consistent parameter estimation with reasonable precision and high power in testing iQTL. When a QTL shows Mendelian effect, the proposed model also outperforms traditional Mendelian model. Extension to incorporate maternal effect is also given. The developed model, built within the maximum likelihood framework and implemented with the EM algorithm, provides a quantitative framework for testing and estimating iQTL involved in the genetic control of complex traits.     

Developing preferences for live female models of the same or other species in white peking ducklings

The preferences of white Peking ducklings (Anas platyrhynchos) for live models was evaluated for the first 7 days of life. The models were all females: a rouen duck (Anas platyrhynchos), a white Peking duck, and a blue Andalusian chicken (Gallus gallus). In three different experiments the orientation of the models was toward the subjects, away from the subjects, and toward the subjects but with fences around the models (to control for model aggressiveness). In all cases the subjects preferred the rouen duck and not the white Peking model or the chicken. In addition, when the models were facing away from the subjects there was a significant peak of preference that appeared on day 4. The latter effect was replicated three times and is therefore considered a reliable phenomenon.     

Bayesian mapping of genome-wide epistatic imprinted loci for quantitative traits

Genomic imprinting, an epigenetic phenomenon of parent-of-origin-specific gene expression, has been widely observed in plants, animals, and humans. To detect imprinting genes influencing quantitative traits, the least squares and maximum likelihood approaches for fitting a single quantitative trait locus (QTL) and Bayesian methods for simultaneously modeling multiple QTL have been adopted, respectively, in various studies. However, most of these studies have only estimated imprinting main effects and thus ignored imprinting epistatic effects. In the presence of extremely complex genomic imprinting architectures, we introduce a Bayesian model selection method to analyze the multiple interacting imprinted QTL (iQTL) model. This approach will greatly enhance the computational efficiency through setting the upper bound of the number of QTLs and performing selective sampling for QTL parameters. The imprinting types of detected main-effect QTLs can be estimated from the Bayes factor statistic formulated by the posterior probabilities for the genetic effects being compared. The performance of the proposed method is demonstrated by several simulation experiments. Moreover, this method is applied to dissect the imprinting genetic architecture for body weight in mouse and fruit weight in tomato. Matlab code for implementing this approach will be available from the authors upon request.     

The effect of egg size variability on thermoregulation of Mallard (Anas platyrhynchos) offspring and its implications for survival

Summary There is a range of egg size phenotypes in Mallards (Anas platyrhynchos) that has a large genetic component. It was hypothesized that egg size variation could play an important role in survival of newly hatched ducklings during their first few days out of the nest when they are most susceptible to thermal stress and starvation. Precocial young must be physiologically capable of maintaining homeothermy in order to spend adequate time foraging. Duckling size at hatching was highly correlated with egg mass, and those hatching from heavier eggs were able to maintain homeothermy at colder environmental temperatures than those from lighter eggs. Heavy ducklings had significantly lower mass-specific cooling rates, but lower critical temperature did not vary significantly among ducklings of different size. Although insulation and energy reserves were not proportionally greater in larger ducklings, those hatching from heavier eggs can survive starvation longer than those from lighter eggs. The relative cold tolerance of young from light and heavy eggs will affect the ratio of time spent foraging to time spent being brooded by the female parent. Although there is no direct evidence that selection is acting on egg size, variation in this trait within a population could be maintained by fluctuating environmental conditions at hatch.     

Threat assessment by domestic ducklings using visual signals: implications for animal-machine interactions

There are increasing opportunities for robots to work amongst animals in agricultural systems. One potential application is in herding animals prior to catching or transportation. An effective herding robot must be able to interrupt ongoing animal behaviour without causing panic or flight reactions. The extent to which different approaching stimuli interrupted the feeding behaviour of pairs of domestic ducklings was examined to assess their suitability as herding stimuli. Experiment 1 assessed the responses of ducklings to an approaching human (the most likely current herding stimulus), a vertical cylinder (a stimulus with the minimum features of a simple robot) and a model fox (simulating the more natural features of a predator). Interruption of feeding was greatest in response to the human and least to the cylinder. Experiment 2 found no significant difference in the extent to which feeding was interrupted when facial appearance and head orientation of the model fox were manipulated. Experiment 3 examined whether biologically realistic features of the model fox affected duckling response. At a distance of 7 m, photographs of the model fox interrupted feeding behaviour as much as the model itself. However, at a distance of 1 m, the model fox caused significantly more feeding interruption than an intact photograph. A fragmented photograph caused the least interruption of feeding behaviour.     

Bilateral imprinting in the chicks of precocial birds in response to monocular presentation of an object

The paper deals with lateralization of memory of imprinted object in chickens of precocial birds after monocular imprinting. Significant preference is shown of the imprinted stimulus in situation of choice from "learned" and "unlearned" eye in 24 hours after imprinting. Lesion of the ventral hyperstriatum of "learned" hemisphere in 28-30 hours after learning retained the preference of the imprinted stimulus. Analogous lesions made directly after learning, eliminated this preference. Conclusion is made about bilateral imprinting at monocular stimulus presentation.