Leaving patches: Effects of travel requirements

Five pigeons were trained in an analogue foraging procedure in which, by completing a travel requirement, they entered a “patch” in which a reinforcer might be available after an unpredictable time. They also had the opportunity, by emitting a defined response, to exit the patch and travel to another patch. Prey availability in a patch was not signaled. Data were collected on the length of time that subjects stayed in patches before exiting (residence times) as a function of various travel requirements: travel for a fixed time in blackout, fixed-interval schedule traveling, fixed-time traveling with an added response required to terminate traveling, and fixed-ratio traveling. For each of these conditions, the required amount of travel (time or responses) was varied over a wide range. As previously reported, residence times increased with increases in fixed-time traveling, as they did with increasing fixed-interval or fixed-ratio traveling. There was no evidence that adding response or work requirements systematically affected residence time except via increased travel time, although 3 of the 5 birds stayed longer in a patch under higher fixed-ratio values. A “threshold-maximization” model described the data well with a single parameter that was consistent across subjects, procedures, and experiments.     

Choice, changeover, and travel

Since foraging in nature can be viewed as instrumental behavior, choice between sources of food, known as “patches,” can be viewed as choice between instrumental response alternatives. Whereas the travel required to change alternatives deters changeover in nature, the changeover delay (COD) usually deters changeover in the laboratory. In this experiment, pigeons were exposed to laboratory choice situations, concurrent variable-interval schedules, that were standard except for the introduction of a travel requirement for changeover. As the travel requirement increased, rate of changeover decreased and preference for a favored alternative strengthened. When the travel requirement was small, the relations between choice and relative reinforcement revealed the usual tendencies toward matching and undermatching. When the travel requirement was large, strong overmatching occurred. These results, together with those from experiments in which changeover was deterred by punishment or a fixed-ratio requirement, deviate from the matching law, even when a correction is made for cost of changeover. If one accepted an argument that the COD is analogous to travel, the results suggest that the norm in choice relations would be overmatching. This overmatching, however, might only be the sign of an underlying strategy approximating optimization.     

Leaving patches: effects of economy, deprivation, and session duration

Three pigeons pecked keys for food reinforcers in a laboratory analogue of foraging in patches. Half the patches contained food (were prey patches). In prey patches, pecks to one key occasionally produced a reinforcer, followed by a fixed travel time and then the start of a new patch. Pecks to another key were exit responses, and immediately produced travel time and then a new patch. Travel time was varied from 0.25 to 16 s at each of three session durations: 1, 4, and 23.5 hr. This part of the experiment arranged a closed economy, in that the only source of food was reinforcers obtained in prey patches. In another part, food deprivation was manipulated by varying postsession feeding so as to maintain the subjects' body weights at percentages ranging from 85% to 95% of their ad lib weights, in 1-hr sessions with a travel time of 12 s. This was an open economy. Patch residence time, defined as the time between the start of a patch and an exit response, increased with increasing travel time, and consistently exceeded times predicted by an optimal foraging model, supporting previously published results. However, residence times also increased with increasing session duration and, in longer sessions, consistently exceeded previously reported residence times in comparable open-economy conditions. Residence times were not systematically affected by deprivation levels. In sum, the results show that the long residence times obtained in long closed-economy sessions should probably be attributed to session duration rather than to economy or deprivation. This conclusion is hard to reconcile with previous interpretations of longer-than-optimal residence times but is consistent with, in economic terms, a predicted shift in consumption towards a preferred commodity when income is increased.     

Pigeons' choices in situations of diminishing returns: fixed- versus progressive-ratio schedules.

In two different discrete-trial procedures, pigeons were faced with choices between fixed-ratio and progressive-ratio schedules. The latter schedules entail diminishing returns, a feature analogous to foraging situations in the wild. In the first condition (no reset), subjects chose between a progressive-ratio schedule that increased in increments of 20 throughout a session and a fixed-ratio schedule that was constant across blocks of sessions. The size of the fixed ratio was varied parametrically through an ascending and then a descending series. In the reset condition, the same fixed-ratio values were used, but each selection (and completion) of the fixed ratio reset the progressive-ratio schedule back to its minimal value. In the no-reset procedure, the pigeons tended to cease selecting the progressive ratio when it equaled or slightly exceeded the fixed-ratio value, whereas in reset, they chose the fixed ratio well in advance of that equality point. These results indicate sensitivity to molar as well as to molecular reinforcement rates, and those molar relationships are similar to predictions based on the marginal value theorem of optimal foraging theory (e.g., Charnov, 1976). However, although previous results with monkeys (Hineline & Sodetz, 1987) appeared to minimize responses per reinforcement, the present results corresponded more closely to predictions based on sums-of-reciprocals of distance from point of choice to each of the next four reinforcers. Results obtained by Hodos and Trumbule (1967) with chimpanzees in a similar procedure were intermediate between these two relationships. Variability of choices, as well as median choice points, differed between the reset and no-reset conditions.(ABSTRACT TRUNCATED AT 250 WORDS)     

Reinforcement frequency and contingency as factors in fixed-ratio behavior

Two variables often confounded in fixed-ratio schedules are reinforcement frequency and response requirement. These variables were isolated by a technique that yoked the distributions of reinforcements in time for one group of pigeons to those of pigeons responding on various fixed-ratio schedules. The contingencies for the yoked birds were then manipulated by adding various tandem fixed-ratio requirements to their schedules. Post-reinforcement pause was approximately equal for the yoked and ratio pigeons, and was relatively insensitive to changes in the tandem requirement. Terminal response rate increased with increases in the tandem requirement, even though reinforcement rate was invariant. This increase was attributed to the progressive interference of the tandem requirement with the differential reinforcement of long interresponse times.     

Soft commitment: self-control achieved by response persistence.

With reinforcement contingent on a single peck on either of two available keys (concurrent continuous reinforcement schedules) 4 pigeons, at 80% of free-feeding weights, preferred a smaller-sooner reinforcer (2.5 s of mixed grain preceded by a 0.5-s delay) to a larger-later reinforcer (4.5 s of mixed grain preceded by a 3.5-s delay). However, when the smaller-sooner and larger-later reinforcers were contingent on a concurrent fixed-ratio 31 schedule (the first 30 pecks distributed in any way on the two keys), all pigeons obtained the larger-later reinforcer much more often than they did when only a single peck was required. This "self-control" was achieved by beginning to peck the key leading to the larger-later reinforcer and persisting on that key until reinforcement occurred. We call this persistence "soft commitment" to distinguish it from strict commitment, in which self-control is achieved by preventing changeovers. Soft commitment also effectively achieved self-control when a brief (1-s) signal was inserted between the 30th and 31st response of the ratio and with concurrent fixed-interval 30-s schedules (rather than ratio schedules) of reinforcement. In a second experiment with the same subjects, the fixed ratio was interrupted by darkening both keys and lighting a third (center) key on which pecking was required for various fractions of the fixed-ratio count. The interruption significantly reduced self-control. When interruption was complete (30 responses on the center key followed by a single choice response), pigeons chose the smaller-sooner reinforcer as frequently as they did when only a single choice response was required.     

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.     

Fixed-ratio discrimination: effects of intermittent reinforcement

Four pigeons had discrimination training that required the choice of a left side-key after completing a fixed-ratio 10 on the center key, and a right side-key choice after fixed-ratio 20. Correct choices were reinforced on various fixed-interval, fixed-ratio, random-interval, and random-ratio schedules. When performance was examined across successive 15-second intervals (fixed-interval and fixed-ratio schedules) accuracy was high in the first 15-second interval, decreased in one or several of the next 15-second intervals, and then increased again as reinforcement was approached. When performance was examined across correct trials on fixed-interval and fixed-ratio schedules, accuracy was lowest immediately after reinforcement, followed by a systematic increase in accuracy as the number of correct choices increased. These patterns were due primarily to errors on fixed-ratio 20 trials. Systematic accuracy patterns did not occur on random-interval or random-ratio schedules. The results indicate that when choice patterns differed on fixed-interval and fixed-ratio schedules, the differences were due to the method of data analysis.     

Second-order schedules with fixed-ratio components: variation of component size

Key pecking by pigeons was reinforced with food under second-order schedules with fixed-ratio units. A constant total number of key pecks was required for reinforcement under each condition, but the size and, inversely, number of fixed-ratio components were varied. The total response requirement of 256 pecks was divided into fixed-ratio units of 128, 64, 32, 8, and 2 responses. A brief stimulus, which always preceded food reinforcement, was presented upon completion of each fixed-ratio unit. Under most conditions, the pattern of within-unit responding was typical of that under simple fixed-ratio schedules. Overall response rate was an inverted U-shaped function of component size. That is, response rates were highest under moderate sized units (fixed ratio 128 and 64). This relationship is consistent with previous determinations of rate as a function of fixed-ratio value for simple fixed-ratio schedules.     

Handling time and choice in pigeons

According to optimal foraging theory, animals should prefer food items with the highest ratios of energy intake to handling time. When single items have negligible handling times, one large item should be preferred to a collection of small ones of equivalent total weight. However, when pigeons were offered such a choice on equal concurrent variable-interval schedules in a shuttlebox, they preferred the side offering many small items per reinforcement to that offering one or a few relatively large items. This preference was still evident on concurrent fixed-cumulative-duration schedules in which choosing the alternative with longer handling time substantially lowered the rate of food intake.     

Aversive aspects of a schedule of positive reinforcement

Six male White Carneaux pigeons were trained to peck at one of two keys to obtain food on several fixed-ratio schedules of reinforcement. Concurrently, the first response on a second key could, I—change the conditions of visual stimulation and remove the food reinforcement contingency, II—change the conditions of stimulation and have no effect upon the reinforcement contingency, or III—do nothing. The second response on the stimulus change key always restored baseline conditions. When second-key responses produced a stimulus change, the number of such responses was a function of the ratio value on the first key. Typically, second-key responses occurred before the start of fixed-ratio runs. The duration of stimulus change periods was an exponential function of the number of responses required for reinforcement when the possibility for reinforcement was not disturbed by periods of stimulus change (Condition II).     

Septal area lesions impair spatial working memory in homing pigeons (Columba livia)

The septo-hippocampal system in birds resembles that of mammals, motivating research into the function of the avian hippocampus while surprisingly little attention has been given to the septum. To investigate a possible role of the avian septum in memory, the effects of septal area lesions on a spatial working memory (SpWM) task was tested in homing pigeons. After preoperative training on an analogue eight-arm (feeders) radial maze, now sham-operated control and septal lesioned pigeons were then trained again on the same task of locating the four feeders on the test phase of a trial that were not baited during the sample phase of a trial. During the test phase of a working memory trial, septal lesioned pigeons, compared to both their own preoperative performance and the performance of the controls, required significantly more choices to locate the four baited feeders not baited during the sample phase of a trial, and they made significantly fewer correct responses to the now baited feeders on their first four choices. The results demonstrate that, like its mammalian counterpart, the avian septum plays an important role in SpWM, suggesting that at least some functional properties of the septum are evolutionarily conserved in birds and mammals.     

Drug discrimination under a concurrent fixed-ratio fixed-ratio schedule.

Pigeons were trained to discriminate 5.0 mg/kg pentobarbital from saline under a two-key concurrent fixed-ratio 10 fixed-ratio 40 schedule of food presentation, in which the fixed-ratio component with the lower response requirement was programmed to reinforce responding on one key after drug administration (pentobarbital-biased key) and on the other key after saline administration (saline-biased key). After responding stabilized, pigeons averaged 98% of their responses on the pentobarbital-biased key during training sessions preceded by pentobarbital, and they averaged 90% of their responses on the saline-biased key during training sessions preceded by saline. In test sessions preceded by doses of pentobarbital, chlordiazepoxide, or ethanol, pigeons switched from responding on the saline-biased key at low doses to responding on the pentobarbital-biased key at higher doses (the dose-response curve was quantal). High doses of phencyclidine produced responding on both keys, whereas pigeons responded almost exclusively on the saline-biased key after all doses of methamphetamine. These and previous experiments using concurrent reinforcement schedules to study drug discrimination illustrate that the schedule of reinforcement is an important determinant of the shape of dose-effect curves in drug-discrimination experiments.     

The effect of punishment shock intensity upon responding under multiple schedules

In the first of two experiments, responses of two pigeons were maintained by multiple variable-interval, variable-ratio schedules of food reinforcement. Concurrent punishment was introduced, which consisted of a brief electric shock after each tenth response. The initial punishment intensities had no lasting effect upon responding. Then, as shock intensity increased, variable-ratio response rates were suppressed more quickly than variable-interval response rates. When shock intensity decreased, variable-interval responding recovered more quickly, but the rates under both schedules eventually returned to their pre-punishment levels. In the second experiment, the following conditions were studied in three additional pigeons: (1) With each shock intensity in effect for a number of sessions, punishment shock intensity was gradually increased and decreased and responding was maintained by multiple variable-ratio, fixed-ratio schedules of food reinforcement; (2) Changes in punishment shock intensity as described above with responding maintained by either a variable-ratio or a fixed-ratio schedule, which were presented on alternate days; (3) Session-to-session changes in shock intensity with responding maintained by multiple variable-ratio, fixed-ratio schedules. Responding under the two schedules was suppressed to approximately the same extent by a particular shock intensity. Also, post-reinforcement pauses under the fixed-ratio schedule increased as response suppression increased.     

A comparison of ratio and interval reinforcement schedules with comparable interreinforcement times

Pigeons were trained to peck keys on fixed-ratio and fixed-interval schedules of food reinforcement. Both schedules produced a pattern of behavior characterized as pause and run, but the relation of pausing to time between reinforcers differed for the two schedules even when mean time between reinforcers was the same. Pausing in the fixed ratio occupied less of the time between reinforcers for shorter interreinforcer times. For two of three birds, the relation was reversed at longer interreinforcer times. As an interreinforcer time elapsed, there was an increasing tendency to return to responding for the fixed interval, but a roughly constant tendency to return to responding for the fixed-ratio schedule. In Experiment 1 these observations were made for both single-reinforcement schedules and multiple schedules of fixed-ratio and fixed-interval reinforcement. In Experiment 2 the observations were extended to a comparison of fixed-ratio versus variable-interval reinforcement schedules, where the distribution of interreinforcement times in the variable interval approximated that for the fixed ratio.     

Temporal patterns of responding in small fixed-ratio schedules

Pigeons were exposed to an ascending series of small fixed-ratio schedules from fixed-ratio 1 to 7. Two of those pigeons were later placed on a fixed-ratio 30 schedule. The two primary dependent variables were the postreinforcement pause and the interresponse time. Changes in these variables under small fixed ratios were sometimes opposite to changes reported with large fixed ratios. For example, postreinforcement pauses decreased in length as the fixed-ratio requirement increased from fixed-ratio 1 to fixed-ratio 3. Also, the interresponse times early in the small fixed-ratio schedule were shorter than those immediately preceding reinforcement. These findings question the role of interresponse-time reinforcement in determining temporal patterns of responding under small fixed-ratio schedules. They also suggest that there may be a limited region in which the independent variable, fixed-ratio size, does not operate as previously described.     

Conjunctive schedules of reinforcement II: response requirements and stimulus effects

Responding of three pigeons was maintained under conjunctive fixed-ratio, fixed-interval schedules where a key peck produced food after both schedule requirements were completed. The individual schedule requirements were then successively removed and reinstated with responding maintained under the following conditions: conjunctive fixed-ratio, fixed-time; fixed-time; and fixed-interval schedules. Patterns of responding changed in accord with the successive removal of the schedule requirements. Compared to the conjunctive fixed-ratio, fixed-interval schedule, pause duration increased and response rate decreased under conjunctive fixed-ratio, fixed-time schedules and under fixed-time schedules alone. Overall mean rates of responding were highest and pause duration lowest under fixed-interval schedules. When changes in the keylight colors were correlated with completion of the fixed-ratio, the end of the fixed-interval, or both of these conditions, the pattern of responding was modified and indicated a greater degree of control by the individual schedules. Although two birds showed large increases in interreinforcement time when they were initially exposed to the conjunctive schedule, when responding stabilized this measure was largely invariant for all birds across most schedule conditions.     

Food-deprivation level alters the effects of morphine on pigeons'' key pecking.

Four pigeons pecked response keys under a multiple fixed-ratio 30 fixed-interval 5-min schedule of food presentation. Components alternated separated by 15-s timeouts; each was presented six times. Pigeons were maintained at 70%, 85%, and greater than 90% of their free-feeding weights across experimental conditions. When response rates were stable, the effects of morphine (0.56 to 10.0 mg/kg) and saline were investigated. Morphine reduced response rates in a dose-dependent manner under the fixed-ratio schedule and at high doses under the fixed-interval schedule. In some cases, low doses of morphine increased rates under the fixed-interval schedule. When pigeons were less food deprived, reductions in pecking rates occurred at lower doses under both schedules for 3 of 4 birds compared to when they were more food deprived. When pigeons were more food deprived, low doses of morphine increased rates of pecking in the initial portions of fixed intervals by a greater magnitude. Thus, food-deprivation levels altered both the rate-decreasing and rate-increasing effects of morphine. These effects may share a common mechanism with increased locomotor activity produced by drugs and with increased drug self-administration under conditions of more severe food deprivation.     

Pigeons discriminate between human feeders

Considered as plague in many cities, pigeons in urban areas live close to human activities and exploit this proximity to find food which is often directly delivered by people. In this study, we explored the capacity of feral pigeons to take advantage of this human-based food resource and discriminate between friendly and hostile people. Our study was conducted in an urban park. Pigeons were fed by two experimenters of approximately the same age and skin colour but wearing coats of different colours. During the training sessions, the two human feeders displayed different attitudes: one of the feeders was neutral and the second was hostile and chased away the pigeons. During the two test phases subsequent to the training phase, both feeders became neutral. Two experiments were conducted, one with one male and one female feeder and the second with two female feeders. In both experiments, the pigeons learned to quickly (six to nine sessions) discriminate between the feeders and maintained this discrimination during the test phases. The pigeons avoided the hostile feeder even when the two feeders exchanged their coats, suggesting that they used stable individual characteristics to differentiate between the experimenter feeders. Thus, pigeons are able to learn quickly from their interactions with human feeders and use this knowledge to maximize the profitability of the urban environment. This study provides the first experimental evidence in feral pigeons for this level of human discrimination.     

Behavioral contrast and relative reinforcement frequency in two multiple schedules

After preliminary variable-interval training, one group of pigeons was trained on a series of multiple variable-interval low-rate reinforcement schedules, while another group was trained on a series of multiple variable-interval fixed-ratio reinforcement schedules. Contrast effects were observed as variable-interval baseline rate changed in a direction away from the change in reinforcement frequency in the other component. The effects of the variable-interval component on performance in the low-rate and fixed-ratio reinforcement components in the multiple schedules were assessed by comparing the birds' performances on each of these schedules alone. Fixed-ratio reinforcement schedules showed a susceptibility to contrast effects, low-rate reinforcement schedules did not. The rate of reinforcement in fixed-ratio schedules at which no interaction occurred in the multiple schedules was higher than that in variable-interval 1-min schedules, suggesting that pigeons may prefer time-based, rather than response-based, reinforcement.