Amodal completion in bonobos

We trained two bonobos to discriminate among occluded, complete, and incomplete stimuli. The occluded stimulus comprised a pair of colored shapes, one of which appeared to occlude the other. The complete and incomplete stimuli involved the single shape that appeared to have been partially covered in the occluded stimulus; the complete stimulus showed the full shape, whereas the incomplete stimulus showed the same portion of the shape that was visible in the occluded stimulus. The correct response was to select the two-part occluded stimulus. Consistent with amodal completion, the bonobos committed a higher percentage of errors to the complete stimuli than to the incomplete stimuli. As well, the percentage of errors to the complete stimuli rose after repeated training with several different shapes. Extensive experience with two-dimensional images enhances amodal completion of partially occluded stimuli in bonobos.     

Amodal completion of moving objects by pigeons

In a series of four experiments, we explored whether pigeons complete partially occluded moving shapes. Four pigeons were trained to discriminate between a complete moving shape and an incomplete moving shape in a two-alternative forced-choice task. In testing, the birds were presented with a partially occluded moving shape. In experiment 1, none of the pigeons appeared to complete the testing stimulus; instead, they appeared to perceive the testing stimulus as incomplete fragments. However, in experiments 2, 3, and 4, three of the birds appeared to complete the partially occluded moving shapes. These rare positive results suggest that motion may facilitate amodal completion by pigeons, perhaps by enhancing the figure - ground segregation process.     

Does the use of natural stimuli facilitate amodal completion in pigeons?

Three experiments were carried out to investigate whether amodal completion in pigeons can be facilitated by the use of colour photographs instead of highly artificial stimuli such as geometrical shapes. Ten pigeons were trained in a go/no-go procedure to discriminate between photographs of complete and of incomplete pigeon figures. In the subsequent test, the birds classified pictures of partly occluded pigeons as though they were complete (experiment 1). However, we found evidence that classification was based on spurious stimulus features that paralleled the intended class rule of figural completeness versus incompleteness. In particular, classification was shown to be guided by white background gaps that separated the parts of the fragmented pigeon figures (experiment 2), as well as by cues related to overall Gestalt (experiment 3). In summary, the present results indicate that the use of more natural stimuli such as photographs instead of geometrical shapes is insufficient for providing amodal completion in pigeons. It is suggested that a combination of various cues, including, eg, 3-D information and common motion in addition to surface and contour properties, may be required to induce a perceptual bias favouring visual completion of occluded portions.     

Perceptual completion in rhesus monkeys (Macaca mulatta) and pigeons (Columba livia)

In a two-dimensional drawing, when the narrow edge of a bar appears to touch the edge of a large rectangle, humans overestimate the length of the bar (Kanizsa, 1979). Kanizsa has suggested that this illusion occurs because humans perceive the bar as continuing behind the rectangle and complete the “occluded” portion of the bar. Rhesus monkeys and pigeons were trained to classify black target bars with a variety of lengths as “long” or “short.” In training, the bar was always located at the same distance from a gray box. After learning this discrimination, the subjects were tested on novel stimuli, in which the bar was located at three new locations. Monkeys showed a consistent response bias for “long” when the bar touched the box, but pigeons did not. Monkeys appear to have completed the “occluded” part like humans, whereas pigeons failed to do so. Because this procedure does not require animals to complete the “occluded” part with any particular form, their failure suggests that pigeons do not even perceive the target bar as continuing behind the “occluding” figure. The failure of pigeons may be due to difficulty in perceiving depth from two-dimensional drawings.     

Perceptual completion in rhesus monkeys (Macaca mulatta) and pigeons (Columbia livia)

In a two-dimensional drawing, when the narrow edge of a bar appears to touch the edge of a large rectangle, humans overestimate the length of the bar (Kanizsa, 1979). Kanizsa has suggested that this illusion occurs because humans perceive the bar as continuing behind the rectangle and complete the "occluded" portion of the bar. Rhesus monkeys and pigeons were trained to classify black target bars with a variety of lengths as "long" or "short." In training, the bar was always located at the same distance from a gray box. After learning this discrimination, the subjects were tested on novel stimuli, in which the bar was located at three new locations. Monkeys showed a consistent response bias for "long" when the bar touched the box, but pigeons did not. Monkeys appear to have completed the "occluded" part like humans, whereas pigeons failed to do so. Because this procedure does not require animals to complete the "occluded" part with any particular form, their failure suggests that pigeons do not even perceive the target bar as continuing behind the "occluding" figure. The failure of pigeons may be due to difficulty in perceiving depth from two-dimensional drawings.     

Effects of occlusion on pigeons' visual object recognition

Casual observation suggests that pigeons and other animals can recognize occluded objects; yet laboratory research has thus far failed to show that pigeons can do so. In a series of experiments, we investigated pigeons' ability to 'name' shaded, textured stimuli by associating each with a different response. After first learning to recognize four unoccluded objects, pigeons had to recognize the objects when they were partially occluded by another surface or when they were placed on top of another surface; in each case, recognition was weak. Following training with the unoccluded stimuli and with the stimuli placed on top of the occluder, pigeons' recognition of occluded objects dramatically improved. Pigeons' improved recognition of occluded objects was not limited to the trained objects but transferred to novel objects as well. Evidently, the recognition of occluded objects requires pigeons to learn to discriminate the object from the occluder; once this discrimination is mastered, occluded objects can be better recognized.     

Pigeons' recognition of partially occluded objects depends on specific training experience

DiPietro et al (2002 Perception 31 1299-1312) reported a dramatic improvement in pigeons' recognition of partially occluded objects after the birds had been trained to recognize objects that were placed on top of another surface. Here, we investigated whether training with partially erased stimuli or with notched stimuli that had a thin gap between the object and another surface would similarly enhance pigeons' recognition of partially occluded objects. We found that erased training had no effect on the birds' recognition of partially occluded objects. Training pigeons to recognize notched objects improved their performance with the same objects when they were partially occluded; but this improvement did not transfer to novel objects, a result that DiPietro et al reported after on-top training. Together, the present results and those of DiPietro et al implicate prior experience as a key factor in pigeons' recognition of partially occluded objects. Training experiences which improve recognition of partially occluded objects may do so because they improve decomposition of complex two-dimensional scenes by pigeons into separate entities.     

Transfer from “edible” categorization training to feeding behavior in pigeons (Columba livia)1

Abstract: We investigated a transfer from an operant experimental situation to a feeding situation in pigeons using real objects as stimuli. Four pigeons were trained in an operant box to categorize familiar edible items as positives and inedible items as negatives with a go/no‐go procedure. Next, two pairs of unfamiliar edible items were added as stimuli. One of the paired stimuli was arbitrarily assigned as positive and the other as negative. We tested the subjects in their home cages to see whether they would feed on the items they were trained to categorize as positives. In three of the six cases in which categorization training was successful, they continued to peck the positive items. This result suggests that the pigeons transfer what they learned in the operant training situation to the feeding situation.     

Discrimination of polymorphous stimulus sets by pigeons

Pigeons were trained to perform a visual discrimination between stimulus sets in which the presence of any two of three positive features made a stimulus positive, while any two of three negative features made it negative (there were thus three different positive and three different negative stimuli). After training, the birds were exposed to test stimuli that contained either all three positive or all three negative features. In Experiment I three pigeons were successfully trained by a successive method, and subsequently responded to the test stimuli as though they were positive or negative respectively. In Experiment II four pigeons were trained by a simultaneous method. Three learned the discrimination and generalized appropriately to the test stimuli, but they showed no preference between positive test and positive training stimuli, nor any consistent difference in speed of response to them; and similar results were found for negative stimuli. It is argued from this that the pigeons learned to respond to the stimuli as patterns (configurations of features) rather than to the constituent features, but that they generalized to the test stimuli by using the common features. The experiments show that pigeons could in principle learn to discriminate natural polymorphous classes (such as “pigeon” or “person”) without using any single feature, but neither the present experiments nor earlier ones demonstrating discriminations of such natural classes establish that pigeons make use of polymorphous concepts in the same way as people.     

Do bantams (Gallus gallus domesticus) amodally complete partly occluded lines? An analysis of line classification performance

Humans perceive a line touching an edge of a large rectangle longer than the reality. Kanizsa (1979) has suggested that this illusion occurs because we perceive that the line is partly "hidden" behind the rectangle and automatically completes it. We tested whether bantams (Gallus gallus domesticus) would experience this perceptual phenomenon using a line classification task on the touch monitor, which was used in our previous study with rhesus monkeys and pigeons (Fujita, 2001). We trained three bantams to classify six lengths of black target lines into two categories, "short" or "long," ignoring a gray rectangle (Experiment 1) or a gray area (i.e., a left or a right half of the monitor was filled with gray; Experiment 2) located at the same distance (8 pixels) from the target line. In the test, the gap between the line and the gray rectangle (or area) sometimes changed (0, 4, or 8 pixels; we labeled these stimuli as G0, G4, and G8 respectively). Both of the two successfully trained bantams showed an illusion for G0, but the direction of illusion was reversed; that is, they judged the line in G0 to be "shorter" than that in G4 and G8. Further analyses proved that neither the gaps between the target line and the gray rectangle nor the total widths of the stimuli could account for the bantams' responses. These results suggest that bantams do not complete the "occluded" portion even when identification of its shape is not required.     

Color-naming functions for the pigeon

Six pigeons were trained to match wavelengths in a three-key matching-to-sample paradigm. Test trials were occasionally presented, where probe wavelengths appeared on the center key and choices were made to the training stimuli presented on the side keys. Color naming functions were obtained by plotting the percentage of test trials that each training stimulus wavelength was chosen for each center key probe wavelength. The wavelength where the functions intersected was interpreted as a transition point between pigeon hues. Three experiments employed different wavelengths as training stimuli. The first two experiments demonstrated that the intersection of the color-naming functions occurred in all cases at 540 nm and 595 nm. The third experiment employed 540 nm and 595 nm as two of the three training stimuli, and the relatively slow acquisition, together with the resulting color-naming functions, supported the proposition that 540 nm and 595 nm may be transition point wavelengths between pigeon hues.     

A new approach to the formation of equivalence classes in pigeons

Four pigeons were given simultaneous discrimination training with visual patterns arbitrarily divided into two sets, with the stimuli in one set designated A1, B1, C1, and D1 and those in the other set designated A2, B2, C2, and D2. In sequentially introduced training phases, the pigeons were exposed to a series of reversals to establish AB and then CD equivalences. In subsequent testing sessions, a subset of stimuli from one set served as positive stimuli and those from the other set as negative stimuli on training trials, and transfer of the reinforced relation to other members of the sets was tested with nonreinforced probe trials. The pigeons were trained further on AC and BD equivalences and then were tested for the emergence of untrained AD and BC equivalences. Two of the 4 pigeons exhibited the emergence of one of these untrained equivalences, evidence for the emergence of transitive relations. This finding suggests that the pigeons established three-member functional equivalence classes by incorporating separately trained multiple equivalence relations. Repeated reversal training and probe testing enabled us to explore the formation and expansion of functional equivalence classes in pigeons.     

Choice as a dependent measure in autoshaping: sensitivity to frequency and duration of food presentation.

Previous investigations have shown that rate, latency, and percentage of trials with at least one response are somewhat insensitive measures of the strength of autoshaped responding. In the present studies, these measures were contrasted with the allocation of responding during simultaneous choice tests, a measure of response strength frequently used in operant paradigms. In two experiments, nine pigeons were exposed to a forward pairing autoshaping procedure. Training sessions consisted of the successive presentation of three stimuli, each followed by food on either 100%, 50%, or 0% of the trials. Choice testing involved the simultaneous presentation of the three stimuli. In Experiment I, all pigeons consistently directed their initial choice responses and the majority of subsequent responses to the stimulus always followed by food, despite the fact that during training sessions the response rates of most birds were highest in the presence of the stimulus followed by food on 50% of the trials. In Experiment II, rate, latency, and percentage of trials with at least one response did not change appreciably as a function of duration of feeder presentations. However, choice responding was lawfully affected by duration of feeder presentations. These data suggest that choice is perhaps a more sensitive measure of the strength of autoshaped responding than other, more commonly employed, indices.     

Contrast and stimulus generalization following prolonged discrimination training

Different groups of pigeons received discrimination training in which the reinforcement-associated and extinction-associated stimuli were respectively either (a) a line tilt vs a blank key, (b) a blank key vs a line tilt, or (c) two different line tilts. The high response rates that developed to the positive stimulus in all groups during discrimination learning were maintained over 64 sessions of training. After these sessions, all subjects were tested for stimulus generalization along the line-tilt dimension. Gradients of relative (per cent) generalization around the stimulus associated with reinforcement (so-called excitatory gradients) and around the stimulus associated with extinction (so-called inhibitory gradients) were as steep as they typically are after much briefer training periods. These results do not support several of Terrace's predictions on the basis of the hypothesis that emotional responses develop to the stimulus associated with extinction during discrimination learning with errors, but eventually dissipate after extended training.     

Pigeons can discriminate “good” and “bad” paintings by children

Humans have the unique ability to create art, but non-human animals may be able to discriminate “good” art from “bad” art. In this study, I investigated whether pigeons could be trained to discriminate between paintings that had been judged by humans as either “bad” or “good”. To do this, adult human observers first classified several children’s paintings as either “good” (beautiful) or “bad” (ugly). Using operant conditioning procedures, pigeons were then reinforced for pecking at “good” paintings. After the pigeons learned the discrimination task, they were presented with novel pictures of both “good” and “bad” children’s paintings to test whether they had successfully learned to discriminate between these two stimulus categories. The results showed that pigeons could discriminate novel “good” and “bad” paintings. Then, to determine which cues the subjects used for the discrimination, I conducted tests of the stimuli when the paintings were of reduced size or grayscale. In addition, I tested their ability to discriminate when the painting stimuli were mosaic and partial occluded. The pigeons maintained discrimination performance when the paintings were reduced in size. However, discrimination performance decreased when stimuli were presented as grayscale images or when a mosaic effect was applied to the original stimuli in order to disrupt spatial frequency. Thus, the pigeons used both color and pattern cues for their discrimination. The partial occlusion did not disrupt the discriminative behavior suggesting that the pigeons did not attend to particular parts, namely upper, lower, left or right half, of the paintings. These results suggest that the pigeons are capable of learning the concept of a stimulus class that humans name “good” pictures. The second experiment showed that pigeons learned to discriminate watercolor paintings from pastel paintings. The subjects showed generalization to novel paintings. Then, as the first experiment, size reduction test, grayscale test, mosaic processing test and partial occlusion test were carried out. The results suggest that the pigeons used both color and pattern cues for the discrimination and show that non-human animals, such as pigeons, can be trained to discriminate abstract visual stimuli, such as pictures and may also have the ability to learn the concept of “beauty” as defined by humans.     

Repeated acquisition of conditional discriminations

A new technique was developed to study the repeated acquisition of conditional discriminations. Using a discrete trial procedure, pigeons were required to learn during each session a different two-member chain of conditional discriminations. Key color and geometric forms were used as stimuli. After the pigeons had reached a steady state of relearning (40 to 60 sessions), the technique was used to investigate variables that have previously been shown to affect the repeated acquisition of response sequences. Various (0 to 90 seconds) durations of timeout for errors were investigated in Experiment I. The stimulus change associated with a timeout, rather than its duration, was found to be the critical variable in acquisition of the discrimination. Extended training on a single chain was found to reduce total errors across sessions in Experiment II. Extended training (three sessions) did not, however, change the pattern of within-session error reduction. In some cases, extended training facilitated acquisition of a partially reversed discrimination. In Experiment III, color rather than chain position was found to control behavior, for three of the four birds, as the second stimulus dimension in the conditional situation. The results of these experiments replicate and extend previous findings concerning some of the variables that affect the repeated acquisition of response sequences.     

Symbolic Representation by Pigeons

The capacity for symbolic representation is a prerequisite for the development of human language because words, the basic units of language, are symbols that represent things. But symbolic representation may also serve a nonlinguistic role of organizing events into categories having the same meaning, and such a capacity could have considerable survival value for many species. In a number of experiments, my co-workers and I have found that pigeons that are trained to treat two different stimuli similarly also learn that those stimuli are commonly represented and, thus, that they have the same meaning. We have demonstrated evidence for such common representations in a number of ways, but perhaps the most convincing is when pigeons learn a new association involving one of the presumed commonly represented stimuli, and without further training demonstrate that they have learned a similar association involving the other stimulus. Furthermore, we have found that when pigeons are trained to treat two stimuli similarly, one of those stimuli is represented in terms of the other. These results have implications not only for the generality of cognitive processes across species, but also for the generality of symbolic representation beyond language use.     

Stimulus control of differential-reinforcement-of-low-rate responding

Five pigeons were given single-stimulus training on an 8-sec differential-reinforcement-of-low-rate schedule followed by steady-state generalization training using 12 wavelength stimuli. Three birds had a high percentage of reinforced responses on the training schedule and flat generalization gradients of total responses. The birds with fewer reinforced responses had much steeper generalization gradients. Generalization gradients plotted as a function of both stimulus wavelength and interresponse time showed that for most birds, stimulus control was restricted to responses with long interresponse times. Responses with very short interresponse times were not under stimulus control and there was some evidence of inhibitory control of short interresponse times. Interresponse-times-per-opportunity functions, plotted as a function of stimulus wavelength, showed that stimulus wavelength controlled the temporal distribution of responses, rather than the overall rate of response. The data indicate that the differential-reinforcement-of-low-rate schedule generates several response categories that are controlled in different ways by wavelength and time-correlated stimuli, and that averaging responses regardless of interresponse-time length obscures this control.     

CONCEPT LEARNING AND LEARNING STRATEGIES

Abstract —Concept teaming and learning strategies of pigeons were manipulated in a matching-Co-sample task. Groups of 4 pigeons responded either 0, J, 10, or 20 times to a sample stimulus, and then chose between a matching comparison stimulus and a nonmatching comparison stimulus. Tests with unfamiliar arrangements of the three training stimuli showed that learning was not by if-then rules. Tests with novel stimuli showed that as the number of sample responses increased, learning about the configural pattern of each display gave way to more teaming about the sample-comparison relationship and more concept learning. Pigeons making the most sample responses showed complete concept teaming.     

Secondary generalization and categorization in pigeons

In Experiment 1, one group of pigeons learned to classify a set of stimuli into the human language classes cat, flower, car, and chair (categorization); another group learned to classify the same set into arbitrary classes (pseudocategorization). Then, both groups were trained on a new categorization task and their performance compared to that of a control group that had no initial classification training. Hull's (1943) notion of secondary generalization (generalization that is not based on physical similarity but on mediating associations) predicts that categorization experience will facilitate the learning of a new categorization task, whereas pseudocategorization experience will impair it. However, in Experiment 1, performance on the new categorization task was not differently affected by prior experience. In Experiment 2, pigeons initially trained to classify a set of 48 stimuli (original training) were later trained to classify a subset of four of these stimuli using new responses (reassignment training). Then, they were tested on the 44 remaining stimuli. Performance better accorded with original than with reassignment training, indicating that categorization training did not lead to the formation of equivalence classes of stimuli, in which the equivalence relationship is mediated by secondary generalization. The lack of evidence of secondary generalization implies that our pigeons failed to meet Lea's (1984) criterion for conceptual behavior.