Pecking: another measurement for filial attachment in group-reared domestic fowl chicks

This study examines different types of attachment measurements and pecking behavior in group-reared domestic chicks imprinted to a silent imprinting object. Immediately upon arriving from the hatchery, groups of heavy strain chicks were imprinted to a colored foam rubber ball for 72 h. The bond between these chicks and the imprinting object was then tested from 8 days until 5 weeks of age. Three days of imprinting sessions demonstrated that a strong attachment could be formed between the imprinted chicks and the imprinting object. In both experimental and control groups, the number of pecks was found to be significantly correlated with the latency measurements. It is, therefore, reasonable to consider the number of pecks of the imprinting object as another measurement for filial attachment.     

Reversibility revisited: Stimulus-dependent stability of filial preference in the chick

The reversibility of filial preferences was studied in domestic chicks, Gallus gallus domesticus, using an artificial as well as a more naturalistic imprinting stimulus. Day-old chicks were exposed for 2 h to either a rotating stuffed jungle fowl or a rotating red box. Two hours later their preferences were tested in a simultaneous choice test (test I), involving the box and the fowl. The next day, the chicks were trained and tested (test II) as before, except box-trained chicks were now exposed to the fowl and vice versa. In three different experimental groups, the relative attractiveness of the two stimuli (as measured in test I) was varied by altering the light intensity of the red box. In the chicks trained with the box and subsequently exposed to the fowl, there was a significant decrease in mean preference for the box from test I to test II. In contrast, the preference of the chicks trained with the stuffed fowl were not affected by subsequent exposure to the red box. This differential reversibility was also found when, in test I, the preference for the box was significantly higher than the preference for the fowl. These results support a model of the formation of filial preference as an interaction between acquired preference and a developing predisposition for a particular class of stimuli.     

MECHANISMS OF AVIAN IMPRINTING: A REVIEW

Filial imprinting is the process through which early social preferences become restricted to a particular object or class of objects. Evidence is presented showing that filial preferences are formed not only as a result of learning through exposure to an object, but also under the influence of visual and auditory predispositions. The development of these predispositions is dependent upon certain non-specific experience. There is little evidence for an endogenously affected sensitive period for imprinting. It is more likely that the end of sensitivity is a result of the imprinting process itself. Similarly, it is now firmly established that filial and sexual preferences are reversible. Evidence suggests, however, that the first stimulus to which the young animal is exposed may exert a greater influence on filial preferences than subsequent stimuli. The learning process of imprinting is often regarded as being different from conventional associative learning. However, the imprinting object itself can function as a reinforcer. Recent studies have attempted to test predictions from an interpretation of filial imprinting as a form of associative learning. The first results suggest that ‘blocking’ may occur in imprinting, whilst there is no evidence for ‘overshadowing’. Social interactions with siblings and parent(-surrogates) have been shown to affect the formation of filial and sexual preferences. The influence of these interactions is particularly prominent in sexual imprinting, making earlier claims about naive species-specific biases unlikely. Although auditory stimuli play an important role in the formation of social attachments, there is little evidence for auditory imprinting per se. Auditory preferences formed as a result of mere (pre- or postnatal) exposure are relatively weak and short-lasting. Exposure to visual stimuli during auditory training significantly improves auditory learning, possibly through a process of reinforcement. It is becoming increasingly clear that filial and sexual imprinting are two different (although perhaps analogous) processes. Different mechanisms are likely to underlie the two processes, although there is evidence to suggest that the same brain region is involved in recognition of familiar stimuli in both filial and sexual imprinting. There is little evidence for a direct role of hormones in the learning process of imprinting. Androgen metabolism may be a factor constraining the development of a predisposition in the chick. Research into the neural mechanism of filial imprinting in the chick has revealed that a restricted part of the forebrain (IMHV) is likely to be a site of memory storage. Changes in synapse morphology and in the number of NMDA receptors have been found, limited to this region, and correlated with the strength of preference.     

Pecking preferences and pre-dispositions in domestic chicks: implications for the development of environmental enrichment devices

Environmental enrichment is thought likely to benefit chickens and farmers in many ways; these include reduced fearfulness and feather pecking and improved productivity. Enrichment devices would intuitively be more effective if they reliably attracted and sustained appreciable interest but many fail to do so. This may reflect the fact that the choice of stimuli often reflects availability and human preconceptions rather than a critical consideration of the birds' preferences and pre-dispositions. We had previously identified string as a particularly attractive pecking stimulus for chicks and adult hens (Gallus gallus domesticus) of a laying strain (ISA Brown). In the present study we found that chicks of another laying strain (Lohmann Brown) also pecked sooner and more at a bunch of string than at chains or beads (Experiment 1). White or yellow strings were preferred to red, green or blue ones (Experiment 2) and white string elicited more pecking than did combinations of white and yellow or of all five colours (Experiment 3). Varying the length and width of the bunches of string exerted no detectable effects on pecking (Experiment 4) whereas incorporating small, shiny beads in the white string devices actually reduced pecking (Experiment 5). Virtually all the devices elicited progressively more interest with repeated presentation; this trend was particularly marked for white string. Collectively, the present findings demonstrate that young domestic chicks have clear and specific pecking preferences. Although the magnitude of response varied across experiments, white string consistently elicited the most interest. Our two main conclusions are: (i) white or yellow strings were particularly attractive stimuli that drew increasing interest, at least in the short term, and (ii) simple devices were preferred to more complex ones, or at least to those used here.     

Experimental analysis of systemogenesis process: c-fos gene expression in chicken brain during stimulation of development of a species-specific acceptor of action response

The aim of this study was to localize the areas of the chick brain involved in the development of filial predisposition to follow conspecifics. Expression of the immediate early gene c-fos mRNA was used to map such structures. One-day-old chicks were stimulated (primed) by placing for 90 min into a running wheel in darkness or by exposure to a loud nonspecific sound for 180 min. Brains of a part of these chicks taken 45 min after the beginning of priming were used for c-fos expression study. The remaining chicks were tested 24 hours after priming by simultaneous presentation of a "natural" object (stuffed fowl) and an "artifical" object (rotating red cube). Primed chicks demonstrated significant preference for the natural object as compared to untreated control chicks. In the forebrain of chicks primed both in the running wheel and by acoustic stimulation a significant c-fos expression was found in the medial part of the caudal neostriatum. Priming in the running wheel additionally induced c-fos expression in the lobus parolfactorius, while priming by acoustic stimulation produced high c-fos expression in the archistriatum. Both these areas are known to be involved in filial imprinting. The results suggest that these structures can be also involved in the development of filial predisposition in chicks.     

Predispositions and preferences. Effects on imprinting of lesions to the chick brain

Domestic chicks were exposed to a moving, stuffed jungle fowl or a rotating red box. The effects on the imprinting process of lesions to a restricted part of the hyperstriatum ventrale (IMHV) were studied in a series of experiments. Sham-operated control chicks developed a strong preference for the training object. Damage to IMHV impaired chicks' preferences for the training stimulus. However, the effect on chicks exposed to the red box was profound, whereas the effect on chicks exposed to the jungle fowl was relatively weak. The results suggest that information about a complex object, which in the experiments described resembled the chicks' own species, is stored in a different way from information derived from a relatively simple artificial object.     

Parental preference for red mouth of chicks in a songbird

Parental preferences during feeding and care-giving may select for ornamental traits in young, such as bright coloration. For chicks of coots, there is experimental evidence for this idea. We examined the hypothesis that bright yellow, orange and red mouths of chicks of songbirds have been favoured by feeding preferences in parents. In a field experiment, the orange–yellow mouths of great tit nestlings were dyed brightly red, and the feeding response of parents recorded. In nest boxes with extra daylight through a window, experimental chicks were on average given twice as much food (biomass) as control chicks (sham dyed). In normal nest boxes, the tendency was similar, but not significant. Thus, at least in good light, great tit parents prefer to feed young with red mouths, a preference for colourfulness that helps explain the evolution of bright gapes in chicks of songbirds (passerine birds).     

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.     

Secondary imprinting in the domestic chick blocked by previous exposure to a live hen

Two experiments were carried out to investigate secondary imprinting in the domestic chick, Gallus gallus. Experiment 1 established that, after chicks with very limited prior visual experience had been exposed to a moving cup during the first 3 days of life, the cup when stationary would suppress distress calling. The main experiment, experiment 2, then examined the effects of various forms of prior experience on the development of this kind of imprinting. Three groups of chicks were exposed on days 1 and 2 to either a live hen (Hen group), a moving object (Windmill group) or the same object when stationary (Control group). On days 4, 5 and 6 all subjects were given training of the kind used in experiment 1; imprinting to the cup was obtained in the Windmill and Control groups, but not in the Hen group. On days 11 and 12 retention tests were given; these showed good retention of primary imprinting, in that the cup was still effective in suppressing distress calling in the Control group and likewise the windmill in the Windmill group, but no effect of secondary imprinting was detected, in that the cup no longer affected the behaviour of the chicks in the Windmill group. These results indicated that primary imprinting with certain stimuli, as in the Hen group, can subsequently exert powerful interference with secondary imprinting and they supported a previous claim that, where secondary imprinting does occur, it is relatively unstable.     

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.     

卷蛾分索赤眼蜂雌蜂的颜色偏好性

为了确定卷蛾分索赤眼蜂Trichogrammatoidea bactrae Nagaraja 雌蜂的颜色偏好性, 在室内通过在培养皿底部黏贴彩纸的方法测定卷蛾分索赤眼蜂雌蜂对红、 黄、 黑、 紫、 绿、 白、 蓝7种颜色的行为趋性反应。结果表明, 卷蛾分索赤眼蜂雌蜂在红、 黄、 紫、 绿和蓝5种颜色上的滞留时间都极显著地高于对照（P<0.01）, 在黑和白2种颜色上的滞留时间与对照没有显著差异（P>0.05）; 对黄色的首次选择率极显著高于对照(P< 0.01), 对红、 紫、 绿和蓝色的首次选择率均显著高于对照(P＜0.05), 对黑色和白色的首次选择率与对照没有显著差异。当雌蜂分别在黄与红、 紫、 绿和蓝两两颜色之间选择时, 雌蜂在黄色彩纸上的滞留时间显著长于其他4种颜色。当雌蜂对红、 紫、 绿、 蓝和黄色5种颜色一起选择时, 在首次选择率、 滞留次数上5种颜色间都没有明显差异(P＞0.05); 但在红色和蓝色上的滞留时间显著长于紫色(P＜0.05), 在这3种颜色上的滞留时间与在黄色和绿色上的滞留时间均无显著差异(P＞0.05)。卷蛾分索赤眼蜂雌蜂在7种颜色卵卡上分别与透明纸（对照）上的米蛾卵的选择寄生时, 在黄色卵卡上的寄生卵量极显著多于对照(P＜0.01), 黑色卵卡上的寄生卵量极显著少于对照(P＜0.01), 其他5种颜色的卵卡上的寄生卵量与对照没有显著差异(P＞0.05)。结果说明, 卷蛾分索赤眼蜂雌蜂对黄色最为偏好, 其次偏好红、 紫、 绿和蓝色, 较不喜好白色和黑色。     

青斑蝶Tirumala limniace成虫颜色偏好性研究

【目的】明确青斑蝶Tirumala limniace雌、雄成虫对颜色的偏好性,可以为此蝴蝶人工饲养时的合理环境条件设定提供必要的理论支持。【方法】本研究通过在室内和野外对青斑蝶成虫对6种颜色(白色、红色、绿色、黄色、蓝色和紫色)光源的选择行为进行观察,明确青斑蝶成虫对不同颜色的偏好性。【结果】(1)在室内和在野外雌、雄成虫均优先选择黄色。(2)雌雄成虫在野外无其它光源干扰下对黄色的选择性强于室内。(3)单色光源照明和多种不同颜色光源同时存在下,成虫对颜色的偏好程度存在差异:当只有单色光源照明时,成虫对颜色的偏好程度为黄色>绿色=红色≥蓝色≥紫色;当不同颜色光源同时存在时,成虫对颜色的偏好程度则为黄色>蓝色≥绿色=红色≥紫色。(4)在不同颜色光源同时存在时,配对后的雌成虫表现出了比雄成虫更强的颜色选择性:雌成虫对黄色光源的选择性强于雄成虫,且其未产生颜色选择的成虫数量也少于雄成虫。【结论】青斑蝶成虫存在对黄色选择的颜色恒常性,野外成虫对颜色的选择性强于室内,雌、雄成虫对偏好颜色选择性存在性别差异。     

Colour categorization by domestic chicks

Spectral stimuli form a physical continuum, which humans divide into discrete non-overlapping regions or categories that are designated by colour names. Little is known about whether non-verbal animals form categories on stimulus continua, but work in psychology and artificial intelligence provides models for stimulus generalization and categorization. We compare predictions of such models to the way poultry chicks (Gallus gallus) generalize to novel stimuli following appetitive training to either one or two colours. If the two training colours are (to human eyes) red and greenish-yellow or green and blue, chicks prefer intermediates, i.e. orange rather than red or yellow and turquoise rather than green or blue. The level of preference for intermediate colours implies that the chicks interpolate between the training stimuli. However, they do not extrapolate beyond the limits set by the training stimuli, at least for red and yellow training colours. Similarly, chicks trained to red and blue generalize to purple, but they do not generalize across grey after training to the complementary colours yellow and blue. These results are consistent with a modified version of a Bayesian model of generalization from multiple examples that was proposed by Shepard and show similarities to human colour categorization.     

Effects of beta-chlornaltrexamine on separation distress in chicks

Intraventricular B-chlornaltrexamine (2 micrograms) increased distress vocalizations (DVs) in chicks, and reduced the ability of intraventricular morphine (.1-.5 micrograms), to inhibit DVs. Object imprinting was not blocked by central CNA, but systemic naloxone (10 mg/kg) did attenuate imprinting to a green but not a red object.     

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.     

Stimulus-Evoked Increase of Glutamate in the Mediorostral Neostriatum/Hyperstriatum Ventrale of Domestic Chick After Auditory Filial Imprinting: An In Vivo Microdialysis Study

Abstract: Imprinting in chicks is a form of juvenile learning that has been used to study the basic cellular mechanisms of learning and memory. The forebrain area mediorostral neostriatum/hyperstriatum ventrale (MNH) is a center for acoustic imprinting. Electrophysiological and pharmacological behavioral studies in the MNH have demonstrated that the glutamatergic system and the associated receptors are critically involved in auditory filial imprinting. Accordingly, we investigated the hypothesis that stimulus-evoked glutamate release may be altered after this learning process. Using an in vivo microdialysis technique, we observed a significantly higher increase of extracellular glutamate level in tone-imprinted chicks during exposure to the previously imprinted tone than in socially imprinted control chicks. In a further series of experiments, where we exposed animals from both experimental groups to handling distress, glutamate levels in MNH showed only a slight increase, whereas we observed a pronounced increase of extracellular glutamate in the lobus parolfactorius (LPO), the avian analogue of the basal ganglia. No difference of distress-evoked glutamate release was found in MNH and LPO between tone-imprinted and socially imprinted chicks. The tone-evoked enhanced glutamate response in tone-imprinted chicks suggests that during auditory imprinting glutamatergic synapses develop the potential to increase transmitter release in response to the imprinting stimulus.     

Intuitive physical reasoning about occluded objects by inexperienced chicks

Questions concerning the role of nature and nurture in higher cognition appear to be intractable if one restricts one's attention to development in humans. However, in other domains, such as sensory development, much information has been gained from controlled rearing studies with animals. Here, we used a similar experimental strategy to investigate intuitive reasoning about occluded objects. Newborn domestic chicks (Gallus gallus) were reared singly with a small object that became their social partner. They were then accustomed to rejoin such an imprinting object when it was made to move and disappear behind either one of two identical opaque screens. After disappearance of the imprinting object, chicks were faced with two screens of different slants, or of different height or different width, which may or may not have been compatible with the presence of the imprinting object hidden beneath/behind them. Chicks consistently chose the screen of slant/height/width compatible with the presence of the object beneath/behind it. Preventing chicks from touching and pecking at the imprinting object before testing did not affect the results, suggesting that intuitive reasoning about physical objects is largely independent of specific experience of interaction with objects and of objects' occluding events.     

Learning and discrimination of coloured papers in the walking blowfly,Lucilia cuprina

Summary A new training and testing paradigm for walking sheep blowflies, Lucilia cuprina, is described. A fly is trained by presenting it with a droplet of sugar solution on a patch of coloured paper. After having consumed the sugar droplet, the fly starts a systematic search. While searching, it is confronted with an array of colour marks consisting of four colours displayed on the test cardboard (Fig. 1). Colours used for training and test include blue, green, yellow, orange, red, white and black.Before training, naive flies are tested for their spontaneous colour preferences on the test array. Yellow is visited most frequently, green least frequently (Table 2). Spontaneous colour preferences do not simply depend on subjective brightness (Table 1).The flies trained to one of the colours prefer this colour significantly (Figs. 5 and 9–11). This behaviour reflects true learning rather than sensitisation (Figs. 6–7). The blue and yellow marks are learned easily and discriminated well (Figs. 5, 9, 11). White is also discriminated well, although the response frequencies are lower than to blue and yellow (Fig. 11). Green is discriminated from blue but weakly from yellow and orange (Figs. 5, 9, 10). Red is a stimulus as weak as black (Figs. 8, 9). These features of colour discrimination reflect the spectral loci of colours in the colour triangle (Fig. 14).The coloured papers seem to be discriminated mainly by the hue of colours (Fig. 12), but brightness may also be used to discriminate colour stimuli (Fig. 13).     

Novelty effects in a multimodal warning signal

The warning signals of toxic insects are often 'multimodal', combining bright coloration with sounds or odours (or both). Pyrazine (a common insect warning odour) can elicit an intrinsic avoidance in domestic chicks Gallus gallus domesticus, both against novel coloured food, and also against food colours that are specifically associated with aposematism, namely yellow and red. In three experiments, we investigated the role of novelty in this innate bias against yellow coloured food in the presence of pyrazine. Naive chicks were familiarized either to pyrazine odour or to coloured food before being tested for a bias against yellow (warningly coloured) food as opposed to green (nonwarningly coloured) food. In experiment 1, pyrazine novelty was shown to be vital for eliciting a bias against yellow food. However, experiment 2 suggested that colour novelty was not important: chicks familiarized with coloured crumbs still avoided yellow crumbs when pyrazine was presented. In a third experiment that gave chicks an even greater degree of pre-exposure to coloured crumbs, the bias against yellow food eventually waned, although pyrazine continued to elicit an aversion to yellow even after birds had had experience of up to 24 palatable yellow crumbs. Pyrazine novelty has been an important pressure in the evolution of multimodal warning signals, and can continue to promote the avoidance of warningly coloured food, even when it is relatively familiar. The implications for warning signals are discussed. Copyright 1999 The Association for the Study of Animal Behaviour.     

Mate choice and imprinting in birds studied by cross-fostering in the wild

Sexual-selection theories generally assume that mating preferences are heritable traits. However, there is substantial evidence that the rearing environment may be important for the development of mating preferences, indicating that they may be learnt, or modified by experience. The relative importance of such sexual imprinting across species remains largely unexplored. Here, we report results of a large-scale cross-fostering experiment in the wild in which nestling birds were raised by parents of a different species. We show that resulting sexual imprinting may have a negative effect on pairing success in one species (the great tit, Parus major), but not in two other species (the blue tit, P. caeruleus and the pied flycatcher, Ficedula hypoleuca). A remarkable variation thus seems to exist, even between species that are congeneric and have similar breeding ecologies. The cross-fostering resulted in heterospecific pairings between the two tit species (female blue tit breeding with male great tit), which has never, to our knowledge, been previously documented. However, the chicks fledging from these nests were all blue tit.