Accurate visual memory of colors in controlling the pecking behavior of quail chicks

Animals are predisposed to memorize specific features of objects they encounter, and to link them with behavioral outputs in a selective manner. In this study, we examined whether chicks memorize objects by colors, and how they exploit the memorized color cues for selective pecking in 1- to 2-days-old quail chicks (Coturnix japonica). Ball-shaped beads painted in green (G), yellowish green (YG) and the intermediate color (YGG) were used. Repetitive presentation of a bead (interval: 4.5 min) resulted in gradually fewer pecks (habituation). Subsequent presentation of a different color caused proportionately more pecks (dishabituation); e.g., after habituation to the G bead, the YG bead caused a stronger dishabituation than the YGG bead did. The dishabituation appeared symmetric; e.g., the YG bead caused as strong dishabituation after the G-habituation, as was caused by the G bead after the YG-habituation. Number of pecks could thus reveal the memory-based color perception in chicks. Similar discrimination of beads by memorized color cues was found after one-trial passive avoidance training, where chicks learned to avoid a bitter-tasting object without any differential pre-training experiences. However, proportion of the chicks that discriminated between different colors became progressively smaller at test 15 min, 1 hr, and 24 hr post-training. On the other hand, proportion of chicks that distinguished beads by non-color cues remained unchanged. Chicks may primarily form an accurate memory of colors, but gradually change the link between the color memory and the pecking behavior.     

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.     

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.     

Predisposed visual memory of shapes in quail chicks

Newly hatched chicks will spontaneously peck at conspicuous objects in their field of view, and soon learn to distinguish between edible food particles and unpleasant tasting objects. To examine whether the selective pecking is based on the ability to memorize shapes, we analyzed pecking behavior of 1- to 2-days-old quail chicks (Coturnix japonica) by using ball- and triangle-shaped beads both painted in green. Repeated presentation of dry bead (either ball or triangle) resulted in a progressively fewer number of pecks (habituation). When chicks were tested by triangle after repeated presentation of ball, chicks showed a significant increase in the number of pecks at the triangle (dishabituation). On the other hand, when tested by ball after a series of triangle presentations, pecking frequency did not increase (no dishabituation). Chicks thus distinguished the triangle as a novel object after being habituated to ball, but did not respond to the ball after triangle. A similar asymmetry was found in one-trial passive avoidance task. Chicks were pre-trained by water-coated (neutral) triangle and then trained by methylanthranilate-coated (aversive) ball. In this case, most chicks learned to avoid the ball, and half of these successful learners pecked at the triangle; they distinguished triangle from ball. When chicks were pre-trained by neutral ball and trained by aversive triangle, on the other hand, most chicks did not distinguish the ball from triangle, and showed a generalized avoidance for both beads. Chicks may be innately predisposed to memorize a limited category of shapes such as ball, and associate them with selective avoidance.     

Is there social transmission of feather pecking in groups of laying hen chicks?

Feather pecking is an abnormal behaviour where laying hens peck the feathers of conspecifics, damaging the plumage or even injuring the skin. If it occurs in a flock, more and more birds show it within a short period of time. A possible mechanism is social transmission. Several studies have shown that laying hen chicks, Gallus gallus domesticus, are able to modify their own behaviour when observing the behaviour of other chicks, for example, when feeding and foraging. As there is good experimental evidence that feather pecking originates from foraging behaviour, we hypothesized that feather pecking could also be socially transmitted. To test this, we reared 16 groups of 30 chicks. After week 4, the birds were regrouped into 16 groups of 20 chicks into each of which we introduced either five chicks that showed high frequencies of feather pecking or, as controls, five chicks that had not developed feather pecking. We then determined the feather-pecking rate and the frequency of foraging, dustbathing, feeding, drinking, preening, moving, standing and resting of all birds in a group. Data from the introduced birds were analysed separately and excluded from the group data. Chicks in groups with introduced feather-pecking chicks had a significantly higher feather-pecking rate than chicks in the control groups. In addition, birds in groups with introduced feather peckers showed significantly lower foraging frequencies than those in the control groups, although the housing conditions were identical and there were no differences in either the number or the quality of the stimuli relevant to foraging behaviour. The study therefore suggests that feather pecking is socially transmitted in groups of laying hen chicks. Copyright 2000 The Association for the Study of Animal Behaviour.     

Effect of diet change on the behavior of chicks of an egg-laying strain

Injurious pecking has serious welfare consequences in flocks of hens kept for egg laying, especially when loose-housed. Frequent diet change is a significant risk for injurious pecking; how the mechanics of diet change influence pecking behavior is unknown. This study investigated the effect of diet change on the behavior of chicks from a laying strain. The study included a 3-week familiarity phase: 18 chick pairs received unflavored feed (Experiment 1); 18 pairs received orange oil-flavored (Experiment 2). All chicks participated in a dietary preference test (P); a diet change (DC); or a control group (C), 6 scenarios. All P chicks preferred unflavored feed. In Experiment 1, DC involved change from unflavored to orange-flavored; Experiment 2, orange- flavored to unflavored. Compared with controls, Experiment 2 DC chicks exhibited few behavioral differences; Experiment 1 DC chicks exhibited increased behavioral event rates on Days 1 and 7. They pecked significantly longer at their environment; by Day 7, they showed significantly more beak activity. There was little evidence of dietary neophobia. Change from more preferred to less preferred feed led to increased activity and redirected pecking behavior.     

Synaptic Vesicle Proteins and Acetylcholine Levels in Chick Forebrain Nuclei Are Altered by Passive Avoidance Training

In a search for biochemical markers of modified synaptic function following training of day-old chicks on a passive avoidance task, we have assayed two monoclonal antibodies to synaptic vesicle proteins (anti-p65 and anti-SV2) and one raised to postsynaptic densities (411B). We have also measured total acetylcholine (ACh) content. Measurements were made on three forebrain regions known to show metabolic and morphological change consequent on training--the lobus parolfactorius (LPO), paleostriatum augmentatum (PA), and medial hyperstriatum ventrale (MHV)--in the right and left hemispheres 2 and 24 h after training chicks on a passive avoidance task, in which they learn to avoid pecking a bead coated with methylanthranilate [methylanthranilate-trained (M-trained)]. Control chicks were trained on a water-coated bead [water-trained (W-trained)]. Twenty-four hours after training, 411B levels showed no differences between W-trained and M-trained chicks in any region. M-training reduced the titre of anti-p65 by 16% in the left PA and 15% in the left MHV and that of anti-SV2 by 19% in the left PA. M-trained chicks showed reduced total ACh content in the LPO by up to 40% and in the PA by up to 48% but had no change in ACh level in the MHV. The decreases in antibody titre were not seen in forebrains analysed 2 h after training, but tendencies toward increases in levels in the right PA and MHV were observed with all three antibodies. Significant differences between right and left hemispheric regions, independent of training, were observed for all the antibodies and for ACh content.(ABSTRACT TRUNCATED AT 250 WORDS)     

Met-enkephalin effects on associations formed in utero

Rat fetuses exposed to an odor stimulus and an aversive stimulus in utero showed an aversion to the odor when tested 16 days postnatally. Fetuses that also received 80 μg/kg Met-enkephalin showed a greater aversion to the odor stimulus than those subjects that did not receive the peptide. The difference between these groups was marginally significant. Control subjects did not show an aversion. But, subjects exposed to the odor and Met-enkephalin without the aversive stimulus, when tested, showed a significant preference for the odor over other control groups. These data show that associative learning in rat fetuses at 20 days of gestation may be enhanced by administration of Met-enkephalin.     

The developmental stage of chicken embryos modulates the impact of in ovo olfactory stimulation on food preferences

Like mammals, bird embryos are capable of chemosensory learning, but the ontogeny of their feeding preferences has not been examined. We tested if the timing of stimulation in chicken embryos modulates the impact of in ovo olfactory stimulation on later food preferences. We exposed chicken embryos to an olfactory stimulus for a 4-day period in the middle or toward the end of the incubation period. The chicks were tested for their preference between foods with and without the olfactory stimulus in 3-min choice tests and on a 24-h time scale. Regardless of the type of food (familiar or novel) or the duration of the test, the control chicks not exposed to the olfactory stimulus consistently showed significant preferences for non-odorized foods. Chicks that were exposed in ovo to the olfactory stimulus did not show a preference for odorized or non-odorized foods. Only those chicks that were exposed to the olfactory stimulus toward the end of the incubation period differed from the controls and incorporated a higher proportion of odorized food into their diets on a 24-h time scale. This result indicates that olfactory stimulation at the end of embryonic development has a stronger impact on later feeding preferences. Our findings contribute to the growing pool of recent data appreciating the impact of olfactory signals on behavior regulation in avian species.     

Familiarity interferes with filial imprinting

The present study was performed to investigate whether and how pre-exposure to an object affects subsequent filial imprinting to that object. In Experiment 1 junglefowl chicks (Gallus gallus spadiceus) were first exposed to either a red object alone (control group), or a red and a yellow object simultaneously (experimental group; phase 1). Subsequently, all chicks were exposed to the yellow object in the presence of a black and blue one (phase 2). At the end of phase 1, most experimental chicks had developed a preference for the red object over the yellow one. At the end of phase 2, preferences of experimental chicks were shifted away from the yellow object towards the novel black and blue object, relative to preferences of control chicks. This shows that pre-exposure may interfere with imprinting. Experiment 2 revealed that when control chicks were tested with the yellow object at the end of phase 1, filial responses were as strong as in experimental chicks. This shows that the yellow object had not acquired control over filial behaviour during phase 1, and also that the relatively impaired imprinting on that object in phase 2 was not due to reduced generalization from the red object. One possible explanation why pre-exposure may interfere with imprinting is that familiarity alters the level of attention attracted by an object, a mechanism suggested to underlie ‘latent inhibition’ in conditioning.     

An Easy Method to Test Odour Recognition in Songbird Hatchlings

We describe an easy method to test odour detection and recognition in 1‐d‐old zebra finch hatchlings (Taeniopygia guttata). Day‐old chicks beg in a stereotypical posture, which can be induced by directing gentle puffs of air from a plastic wash bottle near the face. We used this method to experimentally test whether begging duration of chicks was indicative of nest odour recognition. We manipulated the olfactory environment of 12 nests throughout incubation and hatching with either an artificial odour (orange oil) or with a neutral control (tap water). We then presented these two stimulus odours to 25 day‐old chicks and measured the duration of the first begging bout exhibited for each odour. Zebra finches hatched in a nest environment enriched with orange oil scent begged significantly longer when exposed to orange oil odour, compared to control hatchlings. Our simple testing procedure can be used to efficiently quantify odour recognition and/or preference in altricial songbirds at a very early developmental stage.     

Open-field responses of domestic chicks in the presence or absence of familiar cues

Male domestic chicks were reared in groups with distinctive cues (red crosses) placed on the walls of the home boxes from 2 to 7 d of age and their subsequent behaviour in an open field in the presence or absence of the familiar rearing cues was recorded.Ambulation, peeping, pecking, jumping and standing were all significantly higher and the latency to the first step, duration of freezing, sitting time, lying time and time spent with the eyes closed were significantly lower when the familiar rearing cues were present in the test situation. These observations are consistent with the suggestion that the birds are less fearful in the presence of familiar stimuli. The presence of the familiar cues also increased the time spent feeding and drinking. The preference effect was a specific one, for birds exposed to red crosses in early life preferred such stimuli to black circles in a subsequent two-choice situation, whereas chicks reared with black circles preferred them to red crosses. There was no generalisation of attachment from one stimulus to the other. As the chicks spent more time feeding and drinking in the presence of the familiar cues, it is assumed that fear was reduced. Thus, the results tend to confirm that the frequency of certain behaviour patterns is indicative of certain levels of fear.     

Visual memory of shapes in quail chicks: discrimination among 2-dimensional objects

Newly hatched chicks spontaneously peck at conspicuous objects, and soon learn to discriminate between edible food particles and inedible objects. To examine whether this discrimination is based on a chick's ability to memorize objects by shape cues, we analyzed the pecking behavior. One- to 3-day old quail chicks (Coturnix japonica) were presented with dry objects of different shapes (ball, disk, triangle and T-shape) of similar size (4 mm) and color (green). Habituation occurred after repeated presentation of any one of these objects (duration: 30 sec; interval: 4 min). When chicks showed significantly more pecks at a novel object (dishabituation), we assumed that chicks had memorized the habituated shapes and distinguished the novel object. Chicks did not show dishabituation between a ball and a disk. On the other hand, chicks discriminated a triangle or T-shape from the memorized image of disk, but did not memorize either triangle or T-shape by its shape. Similarly, chicks did not memorize the size of disks as a reference for subsequent pecking behavior. Chicks proved to have a limited ability to memorize shape and size cues for selective pecking behavior, in strong contrast to their accurate memorization of colors.     

Prenatal light exposure affects early feather-pecking behaviour in the domestic chick

Recently we proposed that early feather pecking is a form of social exploration. Social recognition, important for exploration, is a lateralized function in the domestic chick. Lateralization of functions can be influenced by light exposure late in embryonic development. Therefore, we investigated whether this light exposure affected early posthatching feather-pecking behaviour in domestic chicks, Gallus gallus domesticus. White leghorn embryos either were exposed to light or remained in darkness in the last week of incubation. After hatching, they were housed in groups of two light-exposed and two dark-incubated chicks. Light-exposed chicks showed more feather pecking than did their dark-incubated cagemates. Dark-incubated chicks preferred to direct feather pecks to unfamiliar peers than to familiar peers; light-exposed chicks showed no preference. These effects were present in the first week after hatching and remained at least another 3 weeks. These results support the hypothesis that early gentle feather pecking is part of the normal behavioural repertoire of young chicks and influences social exploration. We discuss a possible mechanism underlying these results. We also suggest that it may be worthwhile not to expose embryos to light during the last week of incubation when housing hatchlings in commercial conditions, where feather pecking is a serious problem.     

Ontogenetic and Imprinting-Induced Changes in Chick Brain Protein Metabolism and Muscarinic Receptor Binding Activity

Abstract— Over the 20-min period following exposure of young chicks to a flashing light as an imprinting stimulus there is an increased incorporation of [¹⁴C]leucine into an acidic (tubulin-enriched) protein fraction of the anterior dorsal forebrain in birds which have learnt the characteristics of the stimulus as compared with, either birds which have been exposed to an imprinting stimulus but learn poorly, or chicks kept in the dark. This brain region has been implicated in several studies as the locus for a number of biochemical modulations that accompany learning. The amount of [¹⁴C]leucine incorporated does not seem to be determined by precursor pool availability; it does, however, correlate with a well-validated measure of the extent to which birds have learnt to recognise the characteristics of the stimulus, as shown by a two-choice discrimination test. There is no change in the total content of tubulin dimer as assayed by colchicine binding under these conditions. Additionally, in birds which show evidence of learning, the binding of quinuclidinyl benzilate, an irreversible muscarinic ligand, is altered in both the posterior dorsal forebrain and midbrain regions. None of these effects could be simply the result of visual stimulation. The meaning of these changes is discussed.     

Ontogenetic development of conditioned food aversion in chickens

Development of conditioned food aversion (CFA) was studied in 25-, 35-, and 45-day Leghorn chicks. Food-deprived birds had 10-min access to normal food on days 1 and 2, and to green coloured food CS on day 3. Injection of LiCl (0.15 M, 3–4% body weight) administered 10 min after CS on day 3 served as the US. Control groups were injected with the same volume of saline. Retention was tested on day 4 in three 3-min presentations of normal food alternating with two presentations of green food. Number of pecks and amount of food consumed were measured. Significant neophobic rejection of green food was observed in 25- but not in 35- and 45-day-old control birds. Neophobia in the youngest chicks was further accentuated by CFA which could be observed in pure form in the 45-day-old experimental group. Comparison of pecking rate and food intake showed that CFA in younger birds was accomplished predominantly by inhibition of pecking, and in older birds also by reduction of peck volume. It is concluded that protection against poisoning in chickens shifts between 25 and 45 days of posthatching age from neophobia to CFA.     

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.     

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.     

Reducing feather pecking when raising laying hen chicks in aviary systems

Aviary systems for laying hens offer several advantages over battery cages. However, pecking the feathers of conspecifics remains a serious problem that negatively affects the welfare of the birds as well as the economy of a farm. From experimental studies with small groups, it has been shown that feather pecking and foraging behaviour are related and that both behaviour are influenced by early access to litter substrate. We, therefore, hypothesised, that feather pecking in aviaries can be reduced with an adequate management in the first 2 weeks of life.Each of seven pens on six commercial poultry farms, was divided into two identical compartments (matched pair design). In one of the compartments (experimental compartment) chicks were reared for the first 2 weeks of life with access to litter (wood shavings, in one case with additional straw), while the chicks in the other compartment (control) were kept on a plastic grid. Thereafter, all chicks had unrestricted access to litter and there were no differences between the two compartments neither in housing conditions nor in management procedures.Chicks in the experimental compartments spent significantly more time foraging (week 5), showed significantly less feather pecking (weeks 5 and 14) and significantly fewer birds had damaged tail feathers (weeks 5 and 14).The study demonstrates that in aviaries, under commercial conditions, early access to litter substrate has a significant effect on the development of feather pecking. In order to reduce feather pecking and to increase foraging behaviour, it is recommended that laying hen chicks raised in aviary systems do get access to litter from day 1 on.     

Evidence for olfactory learning in procellariiform seabird chicks

Burrow nesting procellariiform seabirds use olfactory cues for both foraging and nest recognition. As chicks, burrow nesters develop in the dark, but are exposed to both prey‐related and individual‐specific scents through contact with their parents. This exposure suggests that chicks may have the opportunity to learn odours while still in the nest. In this study, we examined whether exposure to odourants during development might influence olfactory search behaviour expressed later in life. To test this idea, we exposed eggs of thin‐billed prions Pachyptila belcheri to a rosy‐scented novel odour (phenyl ethyl alcohol, PEA) or a control (water) just before hatching; chicks were then tested with these odours in a simple wind tunnel. Prior to fledging, subjects who had received pre‐exposure to PEA displayed head sweeps nearly twice as frequently as control birds did when presented with PEA. This study demonstrates that under natural rearing conditions, procellariiforms learn odour characteristics of their rearing environment in the nest.