Collateral behavior of the pigeon during conditioned suppression of key pecking

Ethological recording procedures measured collateral behavior in pigeons whose key-pecking performance was suppressed during a tone that ended with unavoidable electric shock. Independent recordings of gross behavior were made by two observers throughout 60-sec intervals immediately before, during, and after tone presentation. Results indicated significant reductions in the frequency of collateral movements and an increase in the time between successive movements during tone presentations. These effects were observed in all subjects, despite differences in the sequential patterns of behavior. Only partial recovery of the behavior evidenced before tone presentation was found during a 60-sec interval following shock. It was concluded that conditioned suppression procedures caused the bird to “freeze” during tone presentation and in this fashion produced a general inhibitory effect on ongoing overt activity, including key pecking.     

Two types of pigeon key pecking: suppression of long- but not short-duration key pecks by duration-dependent shock

The key pecking of eight pigeons was maintained on a variable-interval 1-minute schedule of food reinforcement. Sometimes, all responses between 35 and 50 milliseconds in duration produced a shock; sometimes, all responses between 10 and 25 milliseconds produced a shock; sometimes, shocks were produced by pecks without regard to duration (nondifferential punishment), and sometimes shocks were delivered independently of responding. Punishment of 35- to 50-millisecond responses selectively suppressed those responses, while punishment of 10- to 25-millisecond responses and nondifferential punishment suppressed responding overall but did not suppress responses of particular duration. Punishment of 35- to 50-millisecond responses suppressed key pecking slightly less than did nondifferential punishment. Punishment of 10- to 25-millisecond responses and response-independent shock produced roughly equal amounts of suppression, substantially less than the other punishment procedures. The data support the view that there are at least two kinds of key peck, identifiable on the basis of duration, one of which (short duration) is insensitive to its consequences.     

On the directionality of key pecking during signals for appetitive and aversive events

Pigeons were exposed to a signal paired with either blackout or blackout plus shock and to another signal paired with food superimposed on a baseline of concurrent variable-interval reinforcement of pecks on two keys. The signals were changes of color of one of the two keys. The rate of pecking both keys during the signal paired with blackout or blackout plus shock was lower than the baseline rate of pecking (a conditioned emotional response), but the decrease in pecking was greater on the signal key. When the intensity of shock was increased, the rate of pecking did not decrease further on the signal key but did decrease on the other key. Rate of pecking during the signal paired with food increased sharply on the signal key (an autoshaping effect) and decreased sharply on the other key. These results support a view that there are two effects of the interaction between classical and instrumental conditioning, a stimulus-directed effect and a generalemotional effect.     

Conditioned suppression, punishment, and aversion

Three experiments were conducted to assess the aversive properties of a visual stimulus in the presence of which one group of birds received response-contingent shock (discriminated punishment) while a yoked group of birds received non-contingent shocks (conditioned suppression). In Experiment 1, presentation of the visual stimulus contingent on key pecking reduced the response rate (conditioned punishment effect) for birds under the conditioned suppression procedure but did not reduce the response rate of birds under the discriminative punishment procedure. Non-contingent shocks also produced greater suppression of responding maintained by positive reinforcement in the presence of a visual stimulus than did response-contingent shocks. In Experiment 2, a greater shock intensity (2 mA) was used. All the differences between the two groups found in Experiment 1 were also found in Experiment 2. Experiment 3 demonstrated that response-contingent shock did not result in a conditioned punishment effect even when positive reinforcers were unavailable during the discriminative punishment schedule. The exteroceptive stimulus that was paired with shock in the conditioned suppression procedure acquired the ability to punish behavior. The exteroceptive stimulus in the discriminative punishment schedule did not acquire this ability.     

Positive and negative conditioned suppression in the pigeon: Effects of the locus and modality of the CS

In Experiment I, four pigeons were exposed to trials in which a 12-sec key light illumination was followed by free food. These trials were superimposed upon a baseline of key pecking for food reinforcement on a variable-interval schedule. When the signal for food was on the operant key, response rate was substantially higher during the signal than during the baseline procedure. When the signal was on a second, signal key, operant responding was suppressed during the signal and substantial pecking of the signal key occurred. The sum of signal key and operant key pecks far exceeded the operant baseline rate of responding. An explanation of opposite results obtained with rats and pigeons as subjects in experiments of this type was suggested in terms of the spatial relation between the signal for free food and the operant target which usually characterizes these experiments. Experiment II assessed the importance of signal location when shock rather than food was the US. Suppression of operant key pecking was unaffected by signal location. Experiment III assessed the relative effectiveness of visual and auditory stimuli (clicks) as signals for food and shock, and found that all combinations of signal and US were equally effective in suppressing operant key responding. The three experiments together suggested that the identification of important effects of species—typical behavior in one experimental situation does not imply that there will be like effects in similar situations.     

Punishment-specific effects of pentobarbital: dependency on the type of punisher.

Pigeons were trained to peck a key under a multiple random-interval 1-minute, random-interval 6-minute schedule of food presentation. Subsequently, over three phases, additions were made during the random-interval 1-minute component as follows: pecks during the component occasionally were punished by timeout presentation (Phase 1), timeouts were presented independently of responding during the component (Phase 2), pecks during the component occasionally were punished by electric-shock presentation (Phase 3). In Phases 1 and 3, response-dependent timeout and shock suppressed responding and established equivalent rates in both components of the multiple schedule. Intermediate doses of pentobarbital increased responding suppressed by electric-shock punishment but had little or no effect on responding suppressed by timeout punishment. Response-independent presentation of timeouts did not result in suppression of responding (thus showing that response-dependent timeout acted as a punisher), and pentobarbital did not reliably increase unpunished responding. Pentobarbital's selective "punishment-attenuating" properties depend on the nature of the punisher.     

Stimulus factors in aversive controls: conditioned suppression after equal training to two stimuli

Pigeons trained to peck a key for food were periodically presented with tones ending with electrical shock until tone presentation consistently suppressed ongoing pecks. Shock was then discontinued and gradients of stimulus generalization were assessed by presenting tones with frequencies above, below, and at the frequencies of those used to develop conditioned suppression. When the training tones had frequencies at 670 and 1500 cps, resulting gradients were bi-modal with peak suppression at 670 and 1500 cps. Of the other test tones, 1000 cps produced the most suppression. When the training tones had frequencies at 450 and 2250 cps, bi-modal gradients were again obtained with peak suppression to the 450 and 2250 cps tones. Of the other test tones, 1000 cps produced the least suppression. These results support the hypothesis that generalized response tendencies summate.     

Mechanisms underlying the effects of unsignaled delayed reinforcement on key pecking of pigeons under variable-interval schedules.

Three experiments were conducted to test an interpretation of the response-rate-reducing effects of unsignaled nonresetting delays to reinforcement in pigeons. According to this interpretation, rates of key pecking decrease under these conditions because key pecks alternate with hopper-observing behavior. In Experiment 1, 4 pigeons pecked a food key that raised the hopper provided that pecks on a different variable-interval-schedule key met the requirements of a variable-interval 60-s schedule. The stimuli associated with the availability of the hopper (i.e., houselight and keylight off, food key illuminated, feedback following food-key pecks) were gradually removed across phases while the dependent relation between hopper availability and variable-interval-schedule key pecks was maintained. Rates of pecking the variable-interval-schedule key decreased to low levels and rates of food-key pecks increased when variable-interval-schedule key pecks did not produce hopper-correlated stimuli. In Experiment 2, pigeons initially pecked a single key under a variable-interval 60-s schedule. Then the dependent relation between hopper presentation and key pecks was eliminated by arranging a variable-time 60-s schedule. When rates of pecking had decreased to low levels, conditions were changed so that pecks during the final 5 s of each interval changed the keylight color from green to amber. When pecking produced these hopper-correlated stimuli, pecking occurred at high rates, despite the absence of a peck-food dependency. When peck-produced changes in keylight color were uncorrelated with food, rates of pecking fell to low levels. In Experiment 3, details (obtained delays, interresponse-time distributions, eating times) of the transition from high to low response rates produced by the introduction of a 3-s unsignaled delay were tracked from session to session in 3 pigeons that had been initially trained to peck under a conventional variable-interval 60-s schedule. Decreases in response rates soon after the transition to delayed reinforcement were accompanied by decreases in eating times and alterations in interresponse-time distributions. As response rates decreased and became stable, eating times increased and their variability decreased. These findings support an interpretation of the effects of delayed reinforcement that emphasizes the importance of hopper-observing behavior.     

Stimulus aspects of aversive controls: the retention of conditioned suppression

Three years ago a tone ending in unavoidable electrical shock was periodically presented to pigeons while they pecked a key for food. When pecking was disrupted by tone, shock was disconnected and the training tone as well as tones of different frequencies were presented. At first, all tones caused a reduction in the rate of pecking, but as testing proceeded, suppression began to extinguish and the gradient narrowed. In the present work, testing was resumed after a 2½-yr interruption. Analysis of the gradients obtained just before and just after the interruption yielded no evidence of changes with the passage of time. As testing proceeded, however, extinction of suppression continued and the gradient all but disappeared. In subsequent experiments with these subjects (Ss) it was found that the presentation of free shocks caused a reappearance of the gradient and that this effect persisted in reduced amount for several sessions after the shock condition was terminated.     

Effects of search cost on foraging and feeding: a three-component chain analysis

An experiment determined whether pigeons minimize number of key pecks per food delivery and maintain their baseline intake of food while key pecking on a three-component chain schedule. Pigeons at either 80% or 100% body weight obtained all their food during baseline and contingency sessions. During baseline sessions, pecks on the left and center keys had no consequences; each peck on the right key activated the feeder. During contingency sessions, pigeons key pecked on a three-component chain schedule simulating components of a foraging chain. In the search component either 3, 9 or 15 key pecks (varied parametrically across blocks of sessions) on the left key produced a stimulus on the middle key, indicating an encounter with either the low-cost prey (3 key pecks) or an equally probable high-cost prey (21 key pecks). In the procurement component the pigeon pecked either: (a) the left key once, thus returning to the search component, or (b) the middle key either 3 or 21 times, which activated the right response key. In the handling component one peck on the right key operated the feeder. The pigeons always procured the low-cost prey and minimized the number of key pecks per hopper by procuring the high-cost prey when the search-cost ratio was high (15 key pecks) but not when it was low (3 key pecks). All pigeons maintained their baselines of eating during contingency sessions by key pecking more frequently and eating more efficiently. The 80% body-weight birds produced higher overall rates of key pecking and eating. These results have implications for ecological theories of optimal foraging and for psychological theories of learned performance.     

The relative law of effect: effects of shock intensity on response strength in multiple schedules

Key pecking of four birds was reinforced with food according to a two-component multiple variable-interval 1-minute variable-interval 4-minute schedule. In addition, key pecking was punished by a brief shock according to a variable-interval 30-second schedule during both components of the multiple schedule. The intensity of the shock was varied. For all birds, punishment had a stronger suppressive effect on the responding maintained by the leaner food schedule, and the ratio of responding during the two components of the multiple schedule became closer to the ratio of reinforcement as shock intensity was increased, as the relative law of effect predicts. At the higher shock intensity, there was some evidence that the ratio of responses overmatched the ratio of reinforcements.     

Enhancement of off-key pecking by on-key punishment

Under a variable-interval food reinforcement schedule, some of a pigeon's pecking responses land on the wall area adjacent to the response key. These off-key pecks increase in frequency when key responses produce shocks and decrease when shock is removed.     

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.     

Constraints on Pavlovian conditioning of the pigeon: Relative conditioned reinforcing effects of red-light and tone CSs paired with food

Recent Pavlovian conditioning experiments presented all possible CS-US combinations of red-light and tone CSs and food and shock USs to separate groups of pigeons. Pigeons receiving shock USs demonstrated conditioned head raising followed by prancing to both CSs, but CRs were acquired more rapidly to tone than to red light. Although pigeons receiving food USs rapidly acquired a conditioned response of pecking to the red-light CS, there was no evidence of conditioned responding in groups receiving tone-food pairings. This outcome left open the possibility that Pavlovian pairings of tone and food may have resulted in association formation that was not revealed in performance. The present series of experiments attempted to reveal that association, using an indirect method of assessment, conditioned reinforcement. Experiment 1 demonstrated that both red light and tone paired with food became positive conditioned reinforcers, suggesting that an association between tone and food was formed in the same number of Pavlovian conditioning trials that previously failed to yield any direct evidence of conditioning. Experiment 2, which presented fewer conditioning trials, revealed that the tone-food association was formed less rapidly than the red light-food association. Experiment 3 demonstrated that the observed outcomes were not attributable to unconditioned, rather than conditioned, reinforcing effects of the Css.     

On some determinants of choice in pigeons

A pigeon's pecking at each of two or three simultaneously available red keys was reinforced at different frequencies with a conditioned reinforcer, an orange key, on which 25 pecks resulted in a presentation of grain. Pecking was occasionally punished with a period of no reinforcement during which each key was dark. Both with two and with three keys, the relative frequency of pecking on a key was equal to the relative frequency of reinforcement obtained by pecks on that key. Also, the absolute frequency of pecking on each key was a linear function with zero intercept of the absolute frequency of reinforcement associated with that key. The slope of this function varied with the number of available keys; it was steeper with two than with three. The relative frequency of switching from any key (two successive pecks on different keys) approximated a linear function with zero intercept and slope slightly greater than 1.0 of the total relative frequency of reinforcement associated with the keys to which the bird could switch. However, the relative frequency of switching to a particular key often showed systematic irregularities. The invariance in these data is the equality between the relative frequency of pecks on one of two or three keys and the relative frequency of reinforcement associated with that key.     

Appetitive-aversive interactions: Superconditioning of fear by an appetitive cs

Four groups of rats received conditioned suppression training in which a tone and light compound was reinforced with shock. If the light had been previously paired with free food, enhanced fear conditioning accrued to the tone during compound training relative to control groups pre-exposed to the light alone, the light semi-randomly associated with food, or the light unpaired with food. The second experiment replicated the difference in aversive conditioning for the groups receiving the light either paired or unpaired with food. The results are discussed in terms of the functional similarity of a conditioned excitor for food and a conditioned inhibitor for shock.     

The effects upon food-reinforced pecking and treadle-pressing of auditory and visual signals for response-independent food

Seven pigeons whose key-pecking was maintained by food reinforcement on a differential-reinforcement-of-low-rates 12-sec limited-hold 4-sec schedule and 12 other pigeons whose treadle-pressing was maintained by the same schedule received appetitive Pavlovian conditioning trials superimposed upon the instrumental baseline. Half the birds in each group received a tone as the CS, and the other half received a stimulus change on the key. Each CS was 20 sec long, and was immediately followed by 10-sec access to grain. The visual CS markedly facilitated the rate of pecking on the key for the birds whose baseline response was pecking. The visual CS produced auto-shaping of the key-peck and tended to produce suppression of treadle-pressing for the birds whose baseline response was treadle-pressing. The auditory CS produced inconsistent effects across birds regardless of the baseline response. In all cases the conditioned effects extinguished when response-independent food was omitted.     

Negatively reinforced key pecking

A reinforcement-switching procedure was used to produce negatively reinforced key pecking in pigeons. First, key pecking on a chain schedule (fixed-interval 10-sec variable-interval 60-sec) was conditioned using grain reinforcement. Second, intermittent shock in the initial link was introduced at a low intensity and gradually increased. Third, food reinforcement in the terminal link was eliminated. With shock at 90 V occurring on the average every 3 sec, initial-link pecking was maintained with no terminal-link food. Three of four pigeons responded consistently at shock intensities of 90, 70, and 50 V but not at 30 V. A fourth pigeon responded at but not below 90 V. Rate of response was directly related to shock frequency. Eliminating food deprivation did not affect the negatively reinforced performance.     

Two different kinds of key peck in the pigeon: some properties of responses maintained by negative and positive response-reinforcer contingencies

Pigeons emitted almost exclusively short-duration key pecks (shorter than 20 msec) when on negative automaintenance procedures, in which pecks prevented reinforcement. Peck durations under fixed-interval and fixed-ratio reinforcement schedules were generally two to five times longer than pecks under a negative automaintenance schedule. However, initial key pecks were of short duration, independent of procedure. The frequency of short-duration pecks was insensitive to differential reinforcement, while the frequency of long-duration pecks was sensitive to differential reinforcement. It is proposed that short-duration pecks arise from the pigeon's normal feeding pattern and are directly enhanced by food presentation, while long-duration pecks are controlled by the contingent effects of food presentation. The implications of the existence of two classes of pecks for the functional definition of operants and the separation of phylogenetic and ontogenetic sources of control of key pecking are discussed.     

Time-dependent changes in conditioned suppression

Time-dependent changes in a response following aversive conditioning were investigated using a conditioned suppression procedure in a within-subjects design. Four groups of pigeons received Pavlovian conditioning “off the baseline”, immediately followed by an operant task. During the Pavlovian phase, two groups received a forward pairing of a tone with shock, one group received a backward pairing, and one group received a truly random pairing. One of the forward pairing groups also received a delay between the Pavlovian and operant phases. For all groups, key pecking was reinforced on a variable-interval schedule during the operant phase. Testing sessions were identical to training sessions, except that the tone used during Pavlovian conditioning was presented either 0, 15, 30, 45, of 60 minutes after the operant phase began. Testing sessions in which the Pavlovian phase was omitted were also included. The results showed suppression to change as a function of the retention interval, with maximum suppression occurring at intermediate intervals. This U-shaped function was obtained for 11 of the 12 pigeons in the forward-pairing groups and for three of the five in the truly random group. Pigeons in the background pairing group did not show changes in suppression as a function of the retention interval.