Some factors that influence the acquisition of complex, stereotyped, response sequences in pigeons

Two pigeons were required to peck six to nine illuminated response keys. A response on any one of the keys darkened that key. When each key had been darkened, a reinforcer was delivered. No specific sequence of key pecking was ever required. The keys were presented in various matrices: three by two, three by three, horizontal rows, and vertical columns. The keys either presented the same stimulus, white light; or each key presented a different stimulus, a color or form. The results indicated that although there were 720 to 362,880 different sequences that would produce reinforcement, each bird developed a particular, stereotyped sequence that dominated its behavior. Variability among the birds across phases yielded less than 60 sequences, .0001 to 6 percent of the possible sequences. The data suggest that a reinforcement contingency that includes “free choice” of response sequence will produce stereotypical response sequences that function as complex “units” of behavior.     

Transfer of matching-to-figure samples in the pigeon

Three pigeons were trained on a modified six-key matching-to-sample procedure. The third peck on the figure-sample key (which presented a bird, hand, face, beetle, rabbit, fish, flower, or red hue, as the sample) lighted only one comparison key. Every three additional pecks on the sample lighted another comparison key, up to a maximum of five keys. Pecks on keys of matching figures produced grain. Pecks on nonmatching keys (mismatches) turned off all lights on the comparison keys and repeated the trial. Three figures were used during acquisition. The birds learned to peck each sample until the matching comparison stimulus appeared on one of three comparison stimulus keys, and then to peck that key. Later, five novel stimuli, employed as both sample and comparison stimuli, and two additional matching keys were added. Each bird showed matching transfer to the novel samples. The data suggest that the birds may have learned the concept of figure matching rather than a series of two-component chains or discrete five-key discriminations.     

Auditory word discriminations in the pigeon.

Four pigeons were trained on a multiple variable-interval 30-s extinction schedule with various pairs of spoken English words presented as the discriminative stimuli. The birds typically produced discrimination indices of 70% to 90% accuracy. Discrimination accuracy was improved by shortening the interval between auditory stimulus presentations, and by increasing the number of syllables in the words.     

Combinations of response-dependent and response-independent schedule-correlated stimulus presentation in an observing procedure

Pigeons pecked a response key on a variable-interval (VI) schedule, in which responses produced food every 40 s, on average. These VI periods, or components, alternated in irregular fashion with extinction components in which food was unavailable. Pecks on a second (observing) key briefly produced exteroceptive stimuli (houselight flashes) correlated with the component schedule currently in effect. Across conditions within a phase, the dependency between observing and presentation of the stimuli was decreased systematically while the density of stimulus presentation was held constant. Across phases, the proportion of session time spent in the VI component was adjusted from 0.5 to 0.25, and then to 0.75. Results indicate that rate of observing decreased as the dependency between responses and stimulus presentations was decreased. Further, discriminative control by the schedule-correlated stimuli was systematically weakened as dependency was decreased. Increasing the proportion of session time spent in VI decreased the rate of observing. This effect was additive with the manipulation of the dependency between observing and presentation of the stimuli. Overall, these results show that conditioned reinforcers function similarly to unconditioned reinforcers with respect to response-consequence dependencies, and that stimulus control is enhanced under conditions in which the relevant stimuli are produced by an organism's behavior.     

Development of complex, stereotyped behavior in pigeons

A pigeon's peck on one key moved a light down one position in a 5×5 matrix of lights, while a peck on another key moved the light across one position. Reinforcement depended upon the occurrence of four pecks on each key (moving the matrix light from the top left to the bottom right), and a fifth peck on either key ended a trial without food. Though there were 70 different sequences that led to reinforcement, each of 12 pigeons developed a particular, stereotyped sequence which dominated its behavior (Experiment 1). Extinction produced substantial increases in sequence variability (Experiment 2). Removal of the matrix cues disrupted performance, though it partially recovered with extended training (Experiment 3). The pigeons did not master a contingency which required a different sequence on the current trial than on the previous one (Experiment 4), though they were able to learn to emit sequences which began with either left-left or left-right response patterns (Experiment 5). The experiments suggest that contingencies of reinforcement may contribute to the creation of complex units of behavior, and that stereotypy may be a likely consequence of contingent reinforcement.     

Discriminated timeout avoidance in pigeons: the roles of added stimuli

Two experiments examined pigeons' postponement of a signaled extinction period, or timeout (TO), from an ongoing schedule of response-dependent food delivery. A concurrent-operant procedure was used in which responses on one (food) key produced food according to a variable-interval schedule and responses on a second (postponement) key delayed the next scheduled TO according to a response-TO (R-TO) interval. A series of response-independent stimulus changes on the food key temporally partitioned the R-TO into three equal segments (S1, S2, and S3). Postponement responses, in addition to postponing TO, also reinstated S1, the stimulus correlated with the greatest temporal distance from TO. In Experiment 1, the R-TO interval was manipulated systematically across blocks of sessions (conditions) at a given ratio of R-TO:TO duration. This R-TO:TO ratio was manipulated across blocks of conditions (phases). Postponement response rates varied inversely with R-TO interval in each phase. Changes in the R-TO:TO ratio did not produce consistent differences except at the 1:10 ratio for some pigeons, where it disrupted postponement responding in some conditions. Most of the postponement responses occurred in the presence of S2 and S3, the stimuli most proximal to TO, whereas most of the food-key responses occurred in S1. In Experiment 2, the R-TO contingencies were systematically manipulated in the presence of the time-correlated stimuli. In one set of conditions, the R-TO contingencies were made either ineffective or less effective in the presence of one or more stimuli. Postponement responses typically shifted to stimuli in the presence of which responses were relatively more effective. Postponement responses decreased markedly when the added stimuli were removed, and then recovered when the stimuli were reinstated. Results from both experiments indicate that the added stimuli in a discriminated TO-avoidance procedure serve predominately discriminative functions, delineating periods during which behavior is maximally effective. The results parallel those obtained in shock-avoidance procedures, providing further evidence that TO functions as an aversive stimulus.     

Binocular depth perception in the pigeon.

By means of a discrete-trial simultaneous discrimination procedure, pigeons were trained to respond differentially to visual arrays that were identical except that one of them contained a circle displaced in depth when viewed stereoscopically. Performance was severely disrupted when one eye was occluded. The monocular deficit was peculiar to the depth task, inasmuch as no such decrement was seen on a pattern discrimination. The results imply that presence of the displaced circle was discriminated on the basis of a binocular cue. It was also found that pigeons could discriminate the direction of the displacement. Discrimination of depth was independent of the global form and still occurred when elements of the array were randomly displaced in depth. Performance was not disrupted when the absolute convergence angle of the depth stimulus was changed. The cue that consistently accounted for the behavior seen was the detection of the relative angles of convergence--that is, the retinal disparity of the two planes in depth. Thus, despite the lateral position of the eyes of the pigeon, a small binocular field mediates the binocular discrimination of near objects in depth.     

Memory for recent behavior in the pigeon

Variations of the symbolic delayed-matching-to-sample procedure were used to study a pigeon's memory for a small number of pecks. In the first experiment a choice of a left or right sidekey after a delay or retention interval was reinforced if a bird had not pecked at all or had pecked exactly once, before the delay, respectively. In the second experiment a choice of a red or green sidekey, regardless of its position, was reinforced if a bird had not pecked at all or had pecked exactly twice, respectively. In the first experiment a bird could orient toward the correct choice during the delay, whereas it could not in the second experiment. In a third experiment a feature-probing method was used to study a pigeon's memory for a number of pecks in the context of certain other pecks. The results showed that a pigeon can remember a small number of pecks for one-half to one minute or more and that the percent correct is a decreasing function of the log retention interval. When a second number of pecks is different from the first number, memory for the first number lasts only a few seconds. When a second number is the same, memory lasts considerably longer. The more recent number of pecks is remembered better. The results are interpreted in terms of a theory which holds that a reinforcer, in general, may act on a subjects' memory for recent behavior to generate patterns of behavior.     

Seasonal variation in pigeon body weight and delayed matching-to-sample performance

The weights of 5 pigeons with free access to food, monitored over 3 calendar years in the laboratory, were found to fluctuate with season. All pigeons were at their heaviest in the winter and were lightest in the summer. Five different pigeons performed a standard delayed matching-to-sample task for 44 weeks from January to November. Their weights were held at 85% of their summer free-feeding weights, making their predicted deprivation level higher in the winter relative to predicted winter free-feeding weights. Slopes of forgetting functions fit to weekly response totals for each pigeon were shallower in winter, showing an improvement in accuracy with longer delays. Thus, delayed matching-to-sample performance may have been affected by the practice of maintaining the pigeons at a constant body weight throughout the calendar year.     

Interresponse-time analysis of stimulus control in multiple schedules

Interresponse-time distributions were recorded in two components of multiple variable-interval schedules that were varied over several conditions. Values of the exponent for power functions relating ratios of interresponse times emitted per opportunity to ratios of reinforcers obtained in the two components varied with interresponse-time class interval. The exponent (sensitivity to reinforcement) afforded a measure of stimulus control exerted by the discriminative stimuli. Exponents were near zero for short interresponse times, consistent with previous conclusions that responses following short interresponse times are controlled by response-produced or proprioceptive stimuli. Values of exponents increased with longer interresponse times, indicating strong control by exteroceptive stimuli over responses following interresponse times of approximately one second or longer.     

Concurrent random-interval schedules and the matching law

In Experiment I, a group of eight pigeons performed on concurrent random-interval schedules constructed by holding probability equal and varying cycle time to produce ratios of reinforcer densities of 1:1, 3:1, and 5:1 for key pecking. Schedules for a second group of seven were constructed with equal cycle times and unequal probabilities. Both groups deviated from simple matching, but the two forms of the schedules appeared to produce no consistent patterns of deviation. The data were found to be consistent with those obtained in concurrent variable-interval situations. The parameters of the matching equation in the form of Y=k X^a were estimated; the value of k was unity and a was 0.84. In Experiment II, six pigeons were exposed to two conc RI RI schedules in which one component increasingly approximated an FI schedule. The value of k was not 1.0. Concurrent RI RI schedules were shown to represent a continuum from conc FI VI to conc VI VI schedules. The use of the exponential equation in testing “matching laws” suggests that a<1 will continue to be observed, and this will set limits on the form of new laws and the assumed or rational values of the component variables in these laws.     

Delayed stimulus control: recall for single and relational stimuli.

In a discrete-trial symbolic matching-to-sample procedure, pigeons' left-key responses were reinforced following presentation of one center-key sample, and right-key responses were reinforced following presentation of another. Recallability was measured by the difference between log ratios of left to right responses following each sample. In Experiment 1, samples were successively presented same or different wavelengths in the relational discrimination, or individual wavelengths in the single discrimination. The rate at which recallability decreased with increasing delay since sample presentation was the same for single and relational discriminations, but the initial level of performance differed, indicating that the relational discrimination was more difficult. In Experiment 2, recall functions for easy and difficult discriminations between individual wavelengths also differed in levels of initial performance but not in rate of decrement of recallability over time. Recall for stimuli differing in complexity may therefore reflect differences in discrimination difficulty.     

Response Stereotypy in Humans Maintained by Response-Contingent Events

The present study examined stereotyped behaviors developed during human performances that were generated by response-dependent intermittent schedules of reinforcement. Thirty university students were assigned to either fixed-interval 30-s or fixed-ratio 30-s schedules in which points or monetary reinforcers were produced only by presses on the number keys of a 41-key computer keyboard. Behavior patterns developed by all subjects were classified into four categories: optimal, random, unique, and general stereotypes. The general stereotypes category was further subdivided into five idiosyncratic types: connection, order, shift, repeat, and restriction. Analysis of the data demonstrated the role of contiguity: Whatever behavior happened to precede reinforcers was repeated even though reinforcers did not depend on that behavior. These findings support the argument that much of idiosyncratic and stereotyped human behavior is produced and maintained by contingencies of reinforcement, rather than schedule-induced or adjunctive behavior.     

Categories in the pigeon brain: A reverse engineering approach

Pigeons are well known for their visual capabilities as well as their ability to categorize visual stimuli at both the basic and superordinate level. We adopt a reverse engineering approach to study categorization learning: Instead of training pigeons on predefined categories, we simply present stimuli and analyze neural output in search of categorical clustering on a solely neural level. We presented artificial stimuli, pictorial and grating stimuli, to pigeons without the need of any differential behavioral responding while recording from the nidopallium frontolaterale (NFL), a higher visual area in the avian brain. The pictorial stimuli differed in color and shape; the gratings differed in spatial frequency and amplitude. We computed representational dissimilarity matrices to reveal categorical clustering based on both neural data and pecking behavior. Based on neural output of the NFL, pictorial and grating stimuli were differentially represented in the brain. Pecking behavior showed a similar pattern, but to a lesser extent. A further subclustering within pictorial stimuli according to color and shape, and within gratings according to frequency and amplitude, was not present. Our study gives proof‐of‐concept that this reverse engineering approach—namely reading out categorical information from neural data—can be quite helpful in understanding the neural underpinnings of categorization learning.     

The associative relation underlying autoshaping in the pigeon

Fifteen pigeons were exposed to either response-independent or response-dependent schedules of water reinforcement, whereby water was injected directly into the unrestrained pigeons' mandibles. Key-contact responses were released by a lighted key correlated with water, but not by a lighted key uncorrelated with water. A negative response-reinforcer contingency suppressed autoshaped key-contact responses, resulting in responding directed away from the lighted key. In all pigeons, water injected directly into the mandibles elicited a consummatory fixed-action pattern of “mumbling” and swallowing. The lighted key correlated with water released a broader set of both appetitive and consummatory responses: approach to the lighted key, “bowing”, “rooting”, “mumbling”, and swallowing. Key-contact responses were “rooting” and “mumbling” motions of the beak on the surface of the key. Views of autoshaping based on stimulus substitution or stimulus surrogation do not fully explain the origin of autoshaped responses not previously elicited by the reinforcer. The present findings are consonant with views of conditioning that emphasize the large degree of biological pre-organization in conditioned response patterns, and the importance of associative factors in the control of such patterns.     

The differentiation of response numerosities in the pigeon

Two experiments examined how pigeons differentiate response patterns along the dimension of number. In Experiment 1, 5 pigeons received food after pecking the left key at least N times and then switching to the right key (Mechner's Fixed Consecutive Number schedule). Parameter N varied across conditions from 4 to 32. Results showed that run length on the left key followed a normal distribution whose mean and standard deviation increased linearly with N; the coefficient of variation approached a constant value (the scalar property). In Experiment 2, 4 pigeons received food with probability p for pecking the left key exactly four times and then switching. If that did not happen, the pigeons still could receive food by returning to the left key and pecking it for a total of at least 16 times and then switching. Parameter p varied across conditions from 1.0 to .25. Results showed that when p= 1.0 or p=.5, pigeons learned two response numerosities within the same condition. When p=.25, each pigeon adapted to the schedule differently. Two of them emitted first runs well described by a mixture of two normal distributions, one with mean close to 4 and the other with mean close to 16 pecks. A mathematical model for the differentiation of response numerosity in Fixed Consecutive Number schedules is proposed.     

Speed analyses of stimulus equivalence.

The functional substitutability of stimuli in equivalence classes was examined through analyses of the speed of college students'' accurate responding. After training subjects to respond to 18 conditional relations, subjects'' accuracy and speed of accurate responding were compared across trial types (baseline, symmetry, transitivity, and combined transitivity and symmetry) and nodal distance (one- through five-node transitive and combined transitive and symmetric relations). Differences in accuracy across nodal distance and trial type were significant only on the first tests of equivalence, whereas differences in speed were significant even after extended testing. Response speed was inversely related to the number of nodes on which the tested relations were based. Significant differences in response speed were also found across trial types, except between transitivity and combined trials. To determine the generality of these comparisons, three groups of subjects were included: An instructed group was given an instruction that specified the interchangeability of stimuli related through training; a queried group was queried about the basis for test-trial responding: and a standard group was neither instructed nor queried. There were no significant differences among groups. These results suggest the use of response speed and response accuracy to measure the strength of matching relations.     

Increasing the variability of response sequences in pigeons by adjusting the frequency of switching between two keys.

Three experiments compared the amounts of behavioral variability generated with two reinforcement rules. In Experiments 1 and 2 pigeons received food whenever they generated a sequence of eight pecks, distributed over two keys, provided that the sequence contained a certain number of change-overs between the keys. Although no variability was required-the birds could obtain all reinforcers by repeating the same sequence-the pigeons emitted a large number of different sequences. In Experiment 3 pigeons received food whenever they generated a sequence that had not occurred during the last 25 trials. After prolonged training, the birds showed more sequence variability than in the first two experiments. The analysis of the internal structure of the response sequences revealed that, in general, (a) the location of the first peck was highly stereotyped; (b) as the trial advanced, the probability of switching to the initially preferred key decreased whereas the probability of switching to the other key increased; and (c) a first-order Markov chain model with transition probabilities given by a logistic function accounted well for the internal structure of the birds' response sequences. These findings suggest that, to a large extent, the variability of response sequences is an indirect effect of adjustments in changeover frequency.     

Discriminative stimulus control and the effects of concurrent operants.

Discriminative-stimulus-control functions were investigated in a concurrent operant context. Variable-interval reinforcement schedules were arranged for pigeons on two response keys. One key, illuminated with a white vertical line on green background (position irrelevant), was programmed with a given schedule value across groups. For different groups of pigeons, the alternative key, illuminated with green alone, was programmed with twice, the same, or half the reinforcement frequency of the other key. Stimulus-control gradients were collected from both keys as line orientation was varied. On the green-plus-line alternative, flattest gradients were observed when twice the reinforcement frequency was concurrently programmed and the steepest were observed when equal values were concurrently programmed. Also examined were the effects of a programmed changeover delay, important in maintaining the independence of concurrent operants. The changeover delay was found to have relatively minor effects upon stimulus control, despite its typical and marked effects upon steady-state responding.