Behavioral control by an imprinted stimulus

Newly hatched ducklings were exposed to imprinting procedures and subsequently trained to peck a key by presenting the imprinting stimulus as the reinforcing (response contingent) event. It was found that the key peck was learned only when imprinting procedures were initiated during the first 6 to 8 hr after hatch. Additional studies revealed that: (1) the duckling's distress vocalizations were reduced in the presence of the imprinting stimulus and enhanced in its absence; (2) when the ducklings had constant access to the imprinted stimulus (via a key peck), pecking responses occurred in bursts and relatively few distress vocalizations occurred; (3) the initial effect of extinction procedures was an increase in key peck rate. When, however, repeated key pecks failed to produce the imprinted stimulus, distress vocalization ensued and peck rate declined; (4) both the presentation of an unfamiliar mechanical figure and delivery of electrical shock enhanced distress vocalization and key pecks; (5) for some ducklings, certain familiar objects in the environment influenced distress calls in a manner comparable to the imprinted stimulus in that distress calls increased when these objects were removed.     

The control of feeding behavior by an imprinted stimulus

Imprinted ducklings were trained to peck a pole using brief presentations of the imprinted stimulus as the response-contingent (reinforcing) event. Subjects were then permitted to spend extended periods with continuous access to food and the imprinted stimulus (via a pole peck). For other (control) subjects the experimental situation was restricted to either responding for the stimulus, or feeding in the absence of the stimulus. For subjects in the control conditions, both activities occurred in cyclic fashion. When, however, there was continuous opportunity to respond for the stimulus and food was available, the tendency to respond was related to the tendency to feed. Other experiments showed that independent presentations of the stimulus could initiate feeding in imprinted ducklings with no prior pairing of the stimulus with food and with no prior pole-peck training. The most consistent control over feeding, however, was exhibited by ducklings that were imprinted and also accustomed to periodic removals of the stimulus. It is concluded that in ducklings, imprinting procedures are sufficient to endow an arbitrary stimulus with the capacity to release feeding behavior.     

Reinforcement by an imprinting stimulus versus water on simple schedules in ducklings

Ducklings (5 to 28 days old) were trained to peck a pole on fixed-ratio, fixed-interval, and multiple schedules using brief presentation of an imprinting stimulus as the response-contingent event. Other ducklings of the same age were trained similarly except that reinforcement consisted of access to water. With water reinforcement the typical fixed-ratio (“break-run”), fixed-interval (“scallop”), and multiple schedule response patterns were readily established and consistently maintained. With the imprinting stimulus these schedule effects were inconsistent in some subjects and virtually nonexistent in others, despite extended training. Schedule control with the imprinting stimulus was not improved by the use of a reinforcement signaling procedure which enhances responding reinforced by electrical brain stimulation on intermittent schedules. However, the overall rates of responding and the extinction functions generated after reinforcement with water versus the imprinting stimulus were comparable. These findings imply that control by temporal and discriminative stimuli may be relatively weak when a young organism's behavior is reinforced by presentation of an imprinting stimulus.     

Schedules of electric shock presentation in the behavioral control of imprinted ducklings

The behavioral effects of various schedules of electric shock presentation were investigated during and after the imprinting of Peking ducklings to moving stimuli. The behavior of following a moving imprinted stimulus was differentially controlled by a multiple schedule of punishment and avoidance that respectively suppressed and maintained following behavior. Pole-pecking, reinforced by presentations of the imprinted stimulus, was suppressed by response-produced shock (punishment); various schedules of response-independent shock and delayed punishment had an overall minimal effect. The delivery of response-independent shock in the presence of one of two stimuli, both during and after imprinting, resulted in a marked reduction in choice of the stimulus paired with shock. The experiments provide no support for a differentiation of imprinting from learning on the basis of the behavioral effects of aversive stimuli. Instead, as is the case with other organisms, the schedule under which shock is delivered to imprinted ducklings appears to be an important determinant of the temporal patterning of subsequent behavior.     

Isolation-induced aggression in newly hatched ducklings.

Newly hatched Khaki Campbell ducklings (Anas platyrhynchos domesticus) were initially housed in pairs and subsequently transferred to isolated housing conditions. Subjects living with another bird displayed filial behavior and little aggression upon encountering another duckling in a test arena. In contrast, ducklings housed in isolation exhibited aggressive pecking in addition to filial behavior when another duckling was subsequently encountered. In Experiment 2, ducklings were housed with an imprinting object (i.e., an object that elicits attachment behavior), but they were otherwise isolated from other birds. These subjects displayed little aggression when they were subsequently reunited with a conspecific, indicating that the aggression-precluding effects of social housing are not limited to the particular social stimulus with which the ducklings are housed. In Experiment 3, ducklings were again housed with an imprinting object, but this time the object was behind glass, thereby precluding tactile contact with it. Since these birds also exhibited little aggression when reunited with a conspecific, it is apparent that visual stimulation from an imprinting object is sufficient in itself to preclude subsequent aggression. These findings suggest that stimulation that elicits attachment behavior is the critical factor mediating isolation-induced aggression.     

Auditory basis of maternal attachment in ducklings (Anas platyrhynchos) under simulated naturalistic imprinting conditions

The presence of broodmates during the imprinting process results in peer imprinting that interrupts a visually mediated maternal attachment. We sought to determine the conditions in which group-trained mallard ducklings (Anas platyrhynchos) acquire a maternal attachment. At 48 hr of age, ducklings were allowed to follow a vocal, stuffed mallard hen individually or in groups of 4. Individual ducklings showed a preference for the silent, familiar mallard over an unfamiliar pintail. Ducklings trained in groups did not show a preference. When the mallard maternal call was present during testing, group-trained ducklings overwhelmingly responded to it regardless of whether it came from the familiar mallard or an unfamiliar pintail. Training ducklings in groups, which simulates the natural social context of imprinting, results in peer imprinting. Thus, early in development the young become visually imprinted to each other, and the maternal call mediates attachment to the mother.     

Social interaction with siblings is necessary for visual imprinting of species-specific maternal preferences in ducklings (Anas platyrhynchos)

This study was conducted to examine the influence of various social rearing experiences on the development of imprinted visual maternal preferences in domestic mallard ducklings (Anas platyrhynchos) during the first 3 days of postnatal life. Twenty-four-hour-old ducklings were allowed to follow a stuffed mallard hen for 30 min. This experience resulted in a visual preference for the familiar mallard hen over an unfamiliar stuffed redhead (Aythya americana) hen in simultaneous choice tests at 48 hr and 72 hr only if the ducklings were reared in conditions allowing unrestricted social interaction with siblings, as would normally occur in nature. No visual preference for the familiar mallard model was found at 48 hr or at 72 hr if ducklings were reared in social isolation but allowed to see another duckling, reared with one duckling, or reared in a group of ducklings but denied the opportunity for direct social interaction. These results demonstrate the importance of normal social experience in the development of the visual imprinting of filial behavior in ducklings. Imprinting studies have traditionally employed isolation rearing and ignored the precocial bird's natural social context. Thus, the present findings raise doubts about the appropriateness of the usual methods of studying imprinting in the laboratory for an understanding of the process of filial imprinting in nature.     

REINFORCEMENT OF DISTRESS VOCALIZATION BY PRESENTATION OF AN IMPRINTED STIMULUS

For one group of ducklings distress calls were reinforced using brief presentation of an imprinted stimulus as the response contingent (reinforcing) event. For a second group of ducklings presentation of an imprinted stimulus was contingent upon a more traditional operant (the pecking response). Regardless of which response was being reinforced, the same pattern ensued. Bursts of responding were separated by periods of little or no responding. When subsequently subjected to repeated presentation and withdrawal of the imprinted stimulus under conditions in which neither response was being reinforced, ducklings with a history of reinforcement for distress calls emitted many more calls than ducklings with a history of reinforcement for pole pecks.     

Preferences among stimulus matches in the pigeon

Three pigeons were trained to peck two to five illuminated response keys. A peck to any of the keys changed the stimulus on the key. When all keys showed the same stimulus (i.e., a stimulus match), an additional key was illuminated with white light. A peck on this key produced three-second access to grain, a three-second intertrial interval, and the next trial. For most sessions, no particular stimulus match was required. Although there were often several stimuli available, each bird preferred a particular stimulus match. With up to 12 stimuli available, birds matched a particular stimulus 60% to 100% of the time.     

Visually guided capture of a moving stimulus by the pigeon (Columba livia)

Although the pigeon is a popular model for studying visual perception, relatively little is known about its perception of motion. Three experiments examined the pigeons’ ability to capture a moving stimulus. In Experiment 1, the effect of manipulating stimulus speed and the length of the stimulus was examined using a simple rightward linear motion. This revealed a clear effect of length on capture and speed on errors. Errors were mostly anticipatory and there appeared to be two processes contributing to response locations: anticipatory peck bias and lag time. Using the same birds as Experiment 1, Experiment 2 assessed transfer of tracking and capture to novel linear motions. The birds were able to capture other motion directions, but they displayed a strong rightward peck bias, indicating that they had learned to peck to the right of the stimulus in Experiment 1. Experiment 3 used the same task as Experiment 2 but with naïve birds. These birds did not show the rightward bias in pecking and instead pecked more evenly around the stimulus. The combined results indicate that the pigeon can engage in anticipatory tracking and capture of a moving stimulus, and that motion properties and training experience influence capture.     

Local contrast and maintained generalization.

Pigeons received variable-interval reinforcement for key pecking during presentations of horizontal and vertical line-orientation stimuli, while pecks during five intermediate orientations were extinguished. Lowest peck rates were observed during presentations of negative stimuli adjacent to the positive orientations while peck rate during 45 degrees (the intermediate negative orientation) was relatively high, i.e., there were negative contrast shoulders. When peck rates were manipulated in the positive orientations, peck rate in neithboring orientations changed in the opposite direction. Contrast shoulders faded after prolonged training. A second type of contrast, local contrast, was correlated with similarity of preceding stimulus and different average peck rates during different stages of the discrimination process. The data suggest that sequential local contrast accompanying the formation of a discrimination contributes to the form of generalization gradients. Blough's model of stimulus control predicts the changes in gradient form described here, but may not accurately depict the underlying process responsible for gradient form.     

Nature reversed: Diminishing intensity of the maternal call affects approach behaviour of chicks

Skinner (1974) suggested that ducklings could be taught to move away from an imprinting object provided the natural consequences of approach behaviour was reversed. We constructed an apparatus in which approach behaviour toward an audio-visual stimulus caused reduced amplitude of a maternal call, whereas movement away caused an increase in amplitude. Chicks (Experiment 1) trained in this apparatus approached the training stimulus, but to a lesser degree than chicks exposed to calls of constant amplitude. Highly active experimental chicks kept a larger distance to the stimulus than passive chicks. Thus the reversal of the natural consequences of approaching influenced filial behaviour. However, a training period of three hours did not influence approach behaviour in the direction predicted by Skinner (1974). A second group of chicks (Experiment 2) which could increase the call amplitude by moving in one direction without being able to localize the sound source was more influenced by the amplitude gradient than Experiment I-chicks.     

The development of stimulus control with and without a lighted key

In two experiments, the effect of an illuminated response key on the acquisition of stimulus control by an airflow stimulus was assessed. In the first experiment, pigeons were given nondifferential training with airflow emerging from behind the response key in one of three conditions of illumination: trained to peck a lighted key, trained to peck an unlighted key with a houselight present, trained to peck a key in total darkness. After 10 days of training on a variable-interval schedule of reinforcement, all subjects were given a generalization test on airflow velocity. The gradients for subjects trained in the dark were sharp, while those for subjects trained in lighted conditions were shallow. In the second experiment, the effect of an irrelevant keylight on the acquisition of an airflow velocity discrimination was assessed. Two groups of pigeons were trained to discriminate two airflow velocities. One group was trained with a lighted response key and the other was trained to peck the response key in total darkness. The dark-trained subjects acquired the discrimination more rapidly. The results demonstrate that the acquisition of stimulus control by airflow with either a differential or nondifferential training procedure can be overshadowed by keylight.     

Aggressive reactions of ducklings to a nonliving target object

This study asked whether ducklings' forceful pecks at a nonliving target object could be validly identified as aggressive. Previously isolated ducklings were exposed to a small cylindrical object that could serve as a target for aggressive pecks and as an object for attachment. After initially attempting to flee from the target, they vigorously pecked at it and also showed signs of the formation of a social (imprinting) attachment. In all important respects this pattern of behavior was identical to the pattern of escape, aggressive pecks, and attachment seen when a previously isolated duckling first encounters a conspecific. Social housing, a manipulation which attenuates aggression against live targets in ducklings and other species, reduced pecking at the nonliving target object. Early aversive stimulation, which enhances aggression against live targets, increased pecking at the object. These findings support the use of nonliving targets in the study of aggression in ducklings.     

Short-term remembering of discriminative stimuli in pigeons.

Pigeons learned to peck the left or right of two white keys depending on whether a red or a green stimulus was displayed on a third key. The opportunity to peck the white keys was then dealyed for zero to six seconds after the red or green (to-be-remembered) stimulus. On half the trials, the feeder operated during the delay to interrupt behavior that might mediate discriminated responding. No events were scheduled on the remaining trials. In a later condition, the pigeons had the opportunity to peck the white keys during the delay. In general, accuracy decreased as delay increased in all conditions, but performance was least accurate following feeder operations and most accurate when pecking was allowed during the delay. The procedures may be analogous to varying the opportunity for rehearsal in studies of human short-term memory.     

Naturalistic psychophysics: thresholds of ducklings (Anas platyrynchos) and chicks (Gallus gallus) to tones that resemble mallard calls

Neonatal ducklings and chickens were tested for responsiveness to a pulsing pure tone that was as similar as possible to the mallard maternal alarm call. It is known that ducklings momentarily cease vocalizing when they hear the alarm call and that chicks do the same when they hear pure tones. The duration of peep suppression can thus be used as a measure of whether subjects of either species heard the stimulus. Chicks might not be as sensitive as ducklings to a mallard alarm call because the signal is less significant to them. An adaptive or staircase procedure was used to estimate absolute thresholds, and group psychometric functions were reconstructed for each species from the trial-by-trial data. Ducklings had lower thresholds than chickens as well as steeper psychometric functions to this stimulus. The results suggest that more sensitive and consistent behavioral responses can be elicited by naturalistic sounds than by more arbitrary acoustic stimuli.     

Stimulus control and the response-reinforcement contingency

Pigeons were trained under a schedule in which reinforcement was made available at varying periods of time after a prior reinforcement. The first key peck after a reinforcer was available began a timer and a second key peck, which exceeded a specified minimal time interval, produced the reinforcer. It was shown that a contingency which contains a minimal interresponse time does not necessarily weaken stimulus control by an exteroceptive stimulus.     

Redundant information in an observing-response procedure

In three observing-response experiments relevant to the information hypothesis of conditioned reinforcement, the basic procedure was one in which an observing response produced one stimulus on trials that terminated in non-contingent reinforcement and another stimulus on trials that terminated in a brief timeout. In Experiment I, the observing response consisted of a single peck or a short fixed-ratio schedule (FR 3 or FR 6), depending on the type of trial. If the single peck produced the negative stimulus and the fixed ratio produced the positive stimulus, observing responses were maintained. If the single peck produced the positive stimulus and the fixed-ratio produced the negative stimulus, observing responses were not maintained on negative trials. In the second experiment, the response key was either white or dark at the beginning of a trial, indicating whether it was a positive or negative trial. Observing responses continued to be maintained on positive trials but not on negative trials. In Experiment III, only positive or negative trials were scheduled for several sessions. Observing responses extinguished regardless of whether positive or negative trials were scheduled. The results do not support the hypothesis that making the stimuli produced by observing responses redundant will reduce observing responses.     

Delayed temporal discrimination in pigeons: A comparison of two procedures

A within-subjects comparison was made of pigeons' performance on two temporal discrimination procedures that were signaled by differently colored keylight samples. During stimulus trials, a peck on the key displaying a slanted line was reinforced following short keylight samples, and a peck on the key displaying a horizontal line was reinforced following long keylight samples, regardless of the location of the stimuli on those two choice keys. During position trials, a peck on the left key was reinforced following short keylight samples and a peck on the right key was reinforced following long keylight samples, regardless of which line stimulus appeared on the correct key. Thus, on stimulus trials, the correct choice key could not be discriminated prior to the presentation of the test stimuli, whereas on position trials, the correct choice key could be discriminated during the presentation of the sample stimulus. During Phase 1, with a 0-s delay between sample and choice stimuli, discrimination learning was faster on position trials than on stimulus trials for all 4 birds. During Phase 2, 0-, 0.5-, and 1.0-s delays produced differential loss of stimulus control under the two tasks for 2 birds. Response patterns during the delay intervals provided some evidence for differential mediation of the two delayed discriminations. These between-task differences suggest that the same processes may not mediate performance in each.