Mediating factors in chick social pecking: 1. Effects of isolation,imprinting, and peer contact

Two experiments examined the influence of periods of social isolation on rates of intersubject pecking in pairs of domestic chicks. Of central interest was the effect of an imprinting condition wherein single birds were reared with either a red or green Styrofoam object. In the first experiment, imprinted subjects were given social pecking tests in the absence of the imprinting object, while in the second study the tests for social pecking were conducted in the presence of one of the imprinting objects. In the latter test the object was familiar to one bud of a pair, but was novel to the other animal. The results of both experiments showed that the rate of intersubject pecking of the imprinted subjects was intermediate to, and statistically different from both the low pecking rate of chicks reared in pairs and the high pecking rate of chicks reared in total isolation. The presence of a familiar object during tests in the second study seemed to serve as a secure base since chicks that had been reared with that object exhibited more social pecking than did chicks that were unfamiliar with the object. However, regardless of the provision of the object the social pecking of the imprinted chicks again fell on a point intermediate to the other groups. Hypotheses concerning a) intolerance of figureground movement and b) sheer stress of isolation were advanced to account for these results.     

Isolation-induced social pecking in domestic chicks: Contemporary correlates and sequelae

Aspects of the social pecking of domestic chicks in a free response situation were examined in a series of experiments. It was proposed that a dual pattern of responses involving (a) a preponderance of pecks at the head of a conspecific, and (b) heightened emotionality (as indicated by the emission of distress calls) might be taken to reflect an aggressive tendency, and evidence was reviewed that showed that a brief period of isolation was a sufficient condition to elicit such a pattern. The main questions asked in this paper are: (a) whether the gender of a chick had a material influence on the incidence of such social pecking, and (b) whether orders established in pecking exchanges reflected a unidimensional social ordering among chicks. In the 1st set of studies the effect of the sex of subject on rates of social pecking was tested. While an isolation procedure clearly produced the dual pattern of heightened head pecking and calling, reliable sex differences were not in evidence. In a 2nd study, concerned with the generality of social orders as observed in social pecking exchanges, the isolation procedure again was sufficient to produce apparently aggressive reactions but the resulting peck orders between chicks did not predict success in later tests when the same 2 subjects were pitted against one another in either approach or avoidance competitive tasks. These results are discussed in terms of the relationship of infant aggression to adult aggression, and suggestions are made for a revised view of the development of the aggressive behavior of fowl.     

Stimulus configuration and valence-enhanced pecking by neonatal chicks

Valence-enhanced pecking occurs when hatchling chicks preferentially peck at releasing stimuli that have been assigned increased releasing power or valence. Enhanced valence is assigned by another releasing stimulus—here a “pecking” arrow. Pecking movements by the arrow have two separate effects: the stimulation or release of pecking by chicks and releasing valence enhancement, that is, pecking directed to particular stimuli. Exposure of chicks to the moving arrow yields pecking significantly increased over that accruing to a stationary arrow. A distinctive stimulus placed on the floor at the tip of the arrow, or, most effective, attached to the tip of the moving arrow, produces enhanced valence: pecking by observing chicks that is selectively increased to matching stimuli.     

Preference for symmetry is experience dependent in newborn chicks (Gallus gallus)

Spontaneous pecking preferences toward symmetric or asymmetric stimuli were tested in newborn chicks (Gallus gallus). A preference for asymmetric patterns was found in na?ve chicks (either 24 or 48 hours old), although a preference for symmetry appeared at retest after chicks had experienced standard rearing conditions (Experiments 1 and 2). Only food-experienced chicks preferred symmetric patterns; food-deprived and hand-fed chicks did not show any preference (Experiment 3). A key factor that allowed for the emergence of a preference for symmetry may relate to the improving of pecking sensorimotor skills occurring during active food manipulation. Possible explanations are discussed for the late emergence of the preference for symmetry and for the preference for asymmetry found in na?ve chicks.     

Environmental variables and releasing-valence transfer in stimulus-directed pecking of chicks

Releasing valence transfer occurs when the power or valence to release responses is transferred from a primary releasing stimulus to a second, initially neutral, stimulus. In the exemplar, pecking responses by neonatal chicks are released and directed by a pointed object operated to make pecking movements. Stimuli attached to or pecked by the arrow thereby acquire enhanced releasing valence. Chicks peck at matching stimuli in preference to comparable but unenhanced stimuli. Research reported here shows that primary and transferred releasing valences are differentially affected by environmental variables. Specific findings were Pecking by chicks occurs only within a narrow range of ambient temperature. Outside of this range, pecking is low in frequency and insensitive to valence-enhanced release. Pecking by chicks appears to be finely tuned to arrow peck rates between 120 and 180 pecks per minute. Within this range, frequency of pecks by chicks is low whereas the percentage of pecks to the valence-enhanced stimulus is maximal. The maternal food call was a weak releaser of pecking by chicks with no valence-enhancing properties.     

Chick vocalization and emotional behavior influenced by apomorphine

For the purpose of studying the role of dopamine (DA) in the causation of vocalization and other behavior in domestic chicks, 5-day-old birds were injected with 1 mg/kg doses of apomorphine hydrochloride, and their behavior was recorded by methods of direct observation. The effects of the drug on birds with bilateral lesions of the intercollicular nucleus (a vocal area) and on birds pretreated with the DA antagonists pimozide and haloperidol were also examined. In intact chicks, apomorphine induced trills, facilitated twitters, and inhibited warbles. Pecking at conspicuous objects in the cage and locomotion were increased, whereas the duration of eye closure was reduced. In chicks with lesions there was no facilitation of trills, twitters, or pecking, whereas the other drug-induced behavioral effects were as in intact chicks. Dopamine antagonists blocked the trills and twitters facilitated by apomorphine but did not protect against the inhibition of warbles. It is concluded that trills, twitters, and pecking are produced by activation of dopaminergic mechanisms. It is hypothesized that some of the behavior induced by apomorphine, especially vocalization and pecking, are a consequence of altered states of attention induced by the drug.     

Mediating factors in chick social pecking: II. effects of exposure to movement

Unfamiliarity with movement may lead isolated chicks to respond aggressively when tested with live, moving conspecifics. This hypothesis was tested by comparing the number and locus of pecks by isolated chicks to a stationary or moving model of a chick, and to a live chick. In a second test of the hypothesis chicks that viewed a moving object during isolation and chicks isolated without movement were tested with a live con-specific and their number and locus of pecks were compared. Neither experiment supported the hypothesis. Movement alone failed to excite aggressive pecks and prior experience with movement failed to suppress aggressive pecks. In the first experiment imlated chicks treated the live testmate more aggressively than the moving or stationary model. In the second experiment aggressive pecking of chicks was unaffected by prior experience with movement, These fmdmgs do not support the hypothesis that movement excites aggression in isolated chicks. Nonetheless the results focus attention on the generally neglected problem of emotional and motivational states produced by isolation.     

Chicks prefer to peck at insect-like elongated stimuli moving in a direction orthogonal to their longer axis

Spontaneous preferences towards possible prey have been little investigated using targets in motion. Preferences of domestic chicks (Gallus gallus) to peck at video-images of stimuli representing live insects moving along their longer body axis (i.e. “forwards”) or along the shorter body axis (i.e. “sideways”) were investigated. Chicks presented with both types of stimulus displayed a significant preference for pecking at stimuli moving sideways. This preference was already present on day 1 post-hatching, and it strengthened on day 6 for those chicks that had experienced pecking at live insects. Head angles used to fixate the stimuli prior to pecking were also analysed and were consistent (i.e. 30°–35° and 60°–65°) with those reported for fixation of non-edible targets (larger stimuli at a distance). In a first control experiment the same video-presented stimuli were used but the insect’s legs were removed to reduce flickering. In a second control experiment, paper-printed images of the whole insect were used. In both cases, the sideways direction of movement was clearly preferred. Overall, our data show that chicks have a spontaneous preference to peck at video-images resembling live insects moving along their shorter body axis. Sideways movement may constitute a crucial signal attracting chicks’ attention and enhancing predatory responses possibly because of stronger stimulation of motion detectors.     

Form preferences in successive discrimination learning of young chicks.

In four experiments the effects of form and orientation pecking preferences of 1- and 3-day old Vantress X Arbor Acre chicks on successive discrimination learning were determined, using heat reinforcement. Major findings were as follows: (a) The young chick has both circle and verticle orientation pecking preferences that are present during at least the first 3 days after hatching; (b) when either of these preferred cues is the nonreinforced cue, the young chick has difficulty in learning not to respond to it but learns quickly not to respond to an unpreferred cue (e.g., triangle and horizontal oriented dots or bar); and (c) these pecking preferences can be modified by heat reinforcement, and the effects of this conditioning is evidenct in subsequent extinction and retention tests. The main conclusion from these experiments is that form and orientation preferences, like brightness and color preferences, are important developmental constraints on conditioning of the young chick.     

Comparability of a single-trial passive avoidance learning task in the young chick across different laboratories

The discrepant results noted by the La Trobe/Monash, Sussex, Open, and Berkeley University memory research groups employing the passive avoidance task (PAT) with the chick indicate that some of these differences may be due to differences in training procedures between the various groups. The procedures employed by each group were replicated as closely as possible and compared using the same strain of chick. Higher levels of pecking and lower training latencies were observed in the La Trobe/Monash chicks. Improved training latency was observed following a change in the day of experimentation with Sussex chicks, and in white light in Open chicks. A powerful reinforcing effect was observed when pretraining was conducted with a wet lure across multiple pretraining trials, indicating the importance of this difference between groups. The observed impact on chick performance of these variables indicates that differences between PAT procedures may have important interactive effects with aversive learning. The results of data gathered using different procedures may only be superficially comparable, and further investigation of the effects of these variables is indicated.     

Presence/absence-sample matching by pigeons: divergent retention functions may result from the similarity of behavior during the absence sample and the retention interval

Divergent choose-absence retention functions typically found in pigeons following presence/absence-sample matching have been attributed to the development of a single-code/default coding strategy. However, such effects may result from adventitious differential responding to the samples. In Experiment 1, retention functions were divergent only when differential sample responding could serve as the basis for comparison choice. In Experiment 2, when pecking did not occur during the retention interval, a choose-absence bias was found, but when pecking occurred during the retention interval, a choose-presence bias resulted. In Experiment 3, positive transfer was found when a stimulus associated with the absence of pecking replaced the absence sample but not when a stimulus associated with pecking replaced the presence sample. Thus, presence/absence-sample matching may not encourage the development of a single-code/default coding strategy in pigeons.     

Passive avoidance learning produces focal elevation of bursting activity in the chick brain: amnesia abolishes the increase

Presentation of a bright bead to day-old chicks (Gallus domesticus: Ross 1 Chunky Chicks) elicits spontaneous pecking. If the bead is coated with an aversive substance (e.g., methylanthranilate), they will avoid similar beads subsequently; if it is coated with water, they peck avidly on re-presentation. Formation of a memory for this one-trial passive avoidance task is unaffected by subconvulsive transcranial electroshock when applied 10 min after training in 60% of birds, whereas "immediate" post-training electroshock renders 63% of chicks amnesic. Memory formation and retention is associated with a large bilateral enhancement in trained over control chicks (320 and 350% in left and right hemispheres, respectively; p less than 0.001) of a particular spontaneous multi-unit activity firing pattern, that is, short-duration (15-40 ms) bursts of large-amplitude (greater than or equal to 200 microV, 450 microV max p-p), high-frequency (400-450 Hz) spiking in anesthetized chicks. This effect is observed in data lumped from 1-13 h after training and is restricted to the intermediate medial hyperstriatum ventrale. When chicks are rendered amnesic by electroshock immediately following training, there is a complete abolition of this increase in burst firing; in those chicks where this treatment fails to elicit amnesia, the increase in bursting is still observed. In birds in which the shock is delayed and memory formation occurs, the increase in bursting activity is maintained; however, if the delayed shock produces apparent amnesia, then the increase is once again abolished. The electroshock had no effect on bursting per se in untrained chicks. There was no significant difference in tonic spiking between the chicks. A marked increase in the occurrence of bursting epochs in the IMHV of anesthetized chicks following passive avoidance training is therefore closely associated with memory formation, but not with the nonspecific concomitants of the training procedure.     

Enhancement of long-term memory retention by Colostrinin in one-day-old chicks trained on a weak passive avoidance learning paradigm

Colostrinin (CLN) is a biologically active proline-rich polypeptide which has therapeutic potential for the alleviation of memory deficits in age-related dementias in a number of human conditions, particularly Alzheimer's disease. To examine the efficacy of CLN in other species, day-old domestic chicks were used as a model system to study its effects on retention of memory for a single one-trial learning paradigm--avoidance of a bitter-tasting substance (methylanthranilate, MeA). Birds were presented with a bead coated with either a dilute (10%) solution of MeA or a bead coated with 100% MeA. Those trained on 100% MeA avoided pecking at a similar but dry bead 24 h later, thereby demonstrating long-term memory whereas chicks trained on the 10% solution pecked the bead at 24 h, indicating lack of long term memory for the task. However, when CLN was injected (i.c.) into a region known to be important in memory formation, the mesopallium intermediomediale (IMM), prior to training with 10% MeA, chicks exhibited strong memory retention at 24 h, similar to those trained on 100% MeA. Control chicks trained on 10% MeA but injected i.c. with a 10% saline solution did not show improvement in memory retention. Intraperitoneal (i.p.) injections of CLN were as effective as the i.c. route. These data extend the known efficacy of CLN from mammals demonstrating its widespread efficacy as a cognitive enhancer.     

Pecking and feeding in chicks

Pecking is not controlled by nutritional state in a newly hatched chick, but by 3 days posthatch it normally is. This paper tests the hypothesis that first a chick must learn that pecking leads to ingestion; then, other (nonlearned) processes are automatically responsible for differential pecking rates to nutritive and nonnutritive items. The first experiment shows that experience pecking and ingesting sawdust is sufficient for differential pecking to develop. The second experiment shows that swallowing without pecking (using a force-feeding technique) does not lead to differential pecking. The third experiment shows that pecking without swallowing (using sand glued down to the floor) also does not lead to differential pecking. Finally, the last experiment shows that food deprivation affects pecking rate only after pecking and ingestion have been associated. These results support the above hypothesis that pecking followed by ingestion leads to the “attachment” of the pecking response to the feeding system. Motivational independence of pecking and the hunger system, the association of pecking with ingestion, and the development of behavior systems are discussed. It is concluded that the function of a behavior may not be related to its causes.     

Topographical variations in behavior during autoshaping, automaintenance, and omission training

Three pigeons were exposed to an autoshaping and automaintenance procedure while a computer-controlled tracking system continuously recorded the position of the bird's head as it moved freely in the experimental chamber. Although only 2 birds pecked the key during the conditional stimulus (red keylight), all 3 birds exhibited stable patterns of approaching the conditional stimulus and withdrawing from the intertrial stimulus (white keylight). Subsequent exposure to an omission procedure, in which pecks on the red key cancelled the presentation of food upon the termination of the red keylight, greatly reduced key pecking, but approaching and pecking in the vicinity of the conditional stimulus were maintained at high levels. When the omission contingency was removed key pecking increased. During all phases the birds withdrew from the area of the white key and engaged in repetitive back-and-forth or circuiting movements during this intertrial stimulus. The data document (a) the strong control the conditional stimulus in autoshaping and automaintenance exerts over approach to the key and pecking motions whether or not the conditional stimulus elicits key pecking at a high level; and (b) withdrawal from the vicinity of the key and the occurrence of stereotypic behavior during the intertrial interval.     

Sequence-Specific Impairment of Memory Formation by NCAM Antisense Oligonucleotides

The functional role of NCAM gene expression in memory formation was studied in the one-trial passive avoidance task in day-old chicks by pretraining injections of one of three different 18-mer end-protected oligonucleotides corresponding to positions 190-, 207-, and 332- of the NCAM Ig1 domain. Twenty-four-hour-old chicks were trained by pecking at a bitter-tasting bead and tested for avoidance 30 min, 3, 8, or 24 hr later. Memory retention was significantly reduced only in the group of animals injected with the NCAM antisense corresponding to position 207- (AS-ODN-207), and only if given twice, both immediately after hatching and 12 hr before training. This antisense was without effect on the general behavior of the chicks, training or acquisition, and did not produce observable neurotoxic damage. Under such conditions amnesia was evident by 3 hr after training and lasted until at least 24 hr after training. The two other tested oligonucleotides were without behavioral effect. To control for nonsequence-specific effects of AS-ODN-207, brains from injected and trained animals were processed for Western blotting and probed using anti-NCAM, anti-L1, and anti-actin antibodies. NCAM antisense corresponding to position 207- significantly reduced the level of NCAM, whereas the level of L1 and actin remained unchanged. These results confirm our earlier conclusion that NCAM is necessary for longer term memory retention.     

Autoshaping as a function of prior food presentations

Young chickens were given 1, 10, 100, or 1000 presentations of grain in a hopper. Subsequently, the key was illuminated before each presentation of grain to study autoshaping of the key-peck response. The number of keylight-grain pairings before a bird first pecked the lighted key was found to be a U-shaped function of the number of prior food-only presentations, with pecks occurring significantly sooner after 100 food-only trials than after any of the other values. Two of five chicks at the 100-trial value pecked on the first illumination of the key. Experiment II showed further that when a series of food-only trials (no keylight) preceded keylight-only trials (no food) 30% of the chicks pecked the illuminated key. Experiment III extended the generality of first-trial pecking to pigeons. After preliminary training with food-only, two of five pigeons pecked on the first illumination of a key. The results suggest a close relationship between autoshaping and pseudo-conditioning.     

Lateralization of a food search task in the domestic chick

Monocularly occluded Warren sex-linked domestic chicks (Gallus domesticus) were presented with a search test which required the differentiation of familiar food grains from unfamiliar pebbles bearing a strong resemblance to food. Male and female chicks using their left eye performed this discrimination significantly more poorly than did chicks using their right eye, both during learning and after asymptotic performance was reached. This left/right difference was superimposed upon a difference between the sexes, with females pecking fewer pebbles than males. Because of the complexity of the search test utilized, it is difficult to determine whether the left/right difference found is attributable to lateralization of visual discrimination ability or to other factors, such as lateralized differences in food search pattern or investigatory responses to novel pebbles. Nevertheless, asymmetric responses of this type provide additional evidence that lateralization of function in the brain, long thought to be a uniquely human phenomenon, is widespread among vertebrates.     

Affiliation and aggression as related to gender in domestic chicks (Gallus gallus).

Social discrimination in male and female domestic chicks (Gallus gallus) was investigated by using (a) latencies of approach response, (b) simultaneous free choice, and (c) intersubject aggressive-pecking tests. In approach-response tests, females showed shorter latencies when tested with cagemates than when tested with strangers, whereas males showed shorter latencies when tested with strangers than when tested with cagemates. In simultaneous-choice tests, females spent more time near a cagemate, whereas males spent more time near a strange chick. In aggressive-pecking tests, both sexes pecked more at strangers than at cagemates; aggressive pecking at strangers, however, was higher in males than in females. It is argued that gender effects in social discrimination can be accounted for in terms of stronger social attachment in females and aggressive responses in males.     

REINFORCER MAGNITUDE ATTENUATES: APOMORPHINE'S EFFECTS ON OPERANT PECKING

When given to pigeons, the direct‐acting dopamine agonist apomorphine elicits pecking. The response has been likened to foraging pecking because it bears remarkable similarity to foraging behavior, and it is enhanced by food deprivation. On the other hand, other data suggest the response is not related to foraging behavior and may even interfere with food ingestion. Although elicited pecking interferes with food capture, it may selectively alter procurement phases of feeding, which can be isolated in operant preparations. To explore the relation between operant and elicited pecking, we provided pigeons the opportunity to earn different reinforcer magnitudes during experimental sessions. During signaled components, each of 4 pigeons could earn 2‐, 4‐, or 8‐s access to grain for a single peck made at the end of a 5‐min interval. In general, responding increased as a function of reinforcer magnitude. Apomorphine increased pecking for 2 pigeons and decreased pecking for the other 2. In both cases, apomorphine was more potent under the component providing the smallest reinforcer magnitude. Analysis of the pattern of pecking across the interval indicated that behavior lost its temporal organization as dose increased. Because apomorphine‐induced pecking varied inversely with reinforcer magnitude, we conclude that elicited pecks are not functionally related to food procurement. The data are consistent with the literature on behavioral resistance to change and suggest that the effects of apomorphine may be modulated by prevailing stimulus—reinforcer relationships.