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.     

Figure-ground assignment in pigeons: evidence for a figural benefit

Four pigeons discriminated whether a target spot appeared on a colored figural shape or on a differently colored background by first pecking the target and then reporting its location: on the figure or the background. We recorded three dependent variables: target detection time, choice response time, and choice accuracy. The birds were faster to detect the target, to report its location, and to learn the correct response on figure trials than on background trials. Later tests suggested that the pigeons might have attended to the figural region as a whole rather than using local properties in performing the figure-background discrimination. The location of the figural region did not affect figure-ground assignment. Finally, when 4 other pigeons had to detect and peck the target without making a choice report, no figural advantage emerged in target detection time, suggesting that the birds' attention may not have been automatically summoned to the figural region.     

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.     

Prior experience affects amodal completion in pigeons

In a three-alternative forced-choice task, 4 pigeons were trained to discriminate a target stimulus consisting of two colored shapes, one of which partially occluded the other, from two foil stimuli that portrayed either a complete or an incomplete version of the occluded shape. The dependent measure was the percentage of total errors that the birds committed to the complete foil. At the outset of training, the pigeons committed approximately 50% of total errors to the complete foil, but as training progressed, the percentage of errors to the complete foil rose. When the pigeons were given a second exposure to the initial set of stimuli, they committed 70% of total errors to the complete foil, suggesting that they now saw the complete foil as more similar to the occluded target than the incomplete foil. These results suggest that experience with 2-D images may facilitate amodal completion in pigeons, perhaps via perceptual learning.     

PIGEONS ARE SENSITIVE TO THE SPATIAL ORGANIZATION OF COMPLEX VISUAL STIMULI

Abstract— Two experiments investigated the role of spatial organisation in the discrimination and generalization of complex visual stimuli by pigeons. In Experiment 1, after pigeons had been trained to discriminate line drawings of four objects, they were tested with novel pictures in which the same component parts of the objects were spatially rearranged. The spatially scrambled pictures led to a dramatic drop in recognition accuracy, hut responding remained above chance. In Experiment 2, pigeons reached a high level of discriminative performance when required to choose among four different spatial arrangements of the same object parts. These results confirm Cerella's (1980) conclusion that pigeons discriminate the component parts of complex visual stimuli, but. unless it is assumed that the scrambling deleted or created emergent features, the results disconfirm his conclusion that spatial organization plays no role in pigeons' picture perception.     

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.     

MATCHING-TO-SAMPLE PERFORMANCE IN RATS: A CASE OF MISTAKEN IDENTITY?

Three rats had previously acquired a simultaneous matching-to-sample performance with steady and blinking lights. In training, the sample stimulus had always appeared on the middle of three horizontally arranged keys with the comparison stimuli on the side keys. In Experiment 1, the sample stimulus appeared on any of the three keys with the comparison stimuli on the remaining two. The matching-to-sample performance broke down with variable sample and comparison locations; the sample stimulus did not control responding to the comparison stimuli when it appeared on a side key, but it retained control when it appeared on the middle key (as in training). In Experiment 2, the rats were trained with the sample always on the left key. When the sample appeared on either of the trained locations (left or middle key), it retained control for both locations. When the sample then appeared on any of the three keys, as in Experiment 1, sample control did not transfer to the untrained location (right key). The experiments demonstrate that training with fixed sample and comparison locations may establish spatial location as an additional controlling aspect of the stimuli displayed on the keys; stimulus location had become part of the definition of the controlling stimuli. The rats' performance seemed best described as specific discriminations involving the visual stimuli and their spatial locations rather than as identity matching.     

Peck tracking: a method for localizing critical features within complex pictures for pigeons

The pigeon is a standard animal in comparative psychology and is frequently used to investigate visuocognitive functions. Nonetheless, the strategies that pigeons use to discriminate complex visual stimuli remain a difficult area of study. In search of a reliable method to identify features that control the discrimination behaviour, pecking location was tracked using touch screen technology in a people-absent/people-present discrimination task. The correct stimuli contained human figures anywhere on the picture, but the birds were not required to peck on that part. However, the stimuli were designed in a way that only the human figures contained distinguishing information. All pigeons focused their pecks on a subarea of the distinctive human figures, namely the heads. Removal of the heads significantly impaired performance, while removal of other distinctive parts did not. Thus, peck tracking reveals the location within a complex visual stimulus that controls discrimination behaviour, and might be a valuable tool to reveal the strategies pigeons apply in visual discrimination tasks.     

Required pecking alters judgments of the passage of time by pigeons

There is evidence that how humans perceive time is affected by the activity in which they are engaged when they are judging time. In humans, typically, the more demanding the task, the faster time seems to pass. We asked whether a similar effect could be found in pigeons. Pigeons were trained to discriminate between a short-(2-sec) and a long-(10-sec) duration stimulus. Depending on the color of the stimulus (white or blue), the pigeons were required to peck (at least once per second or the trial was aborted) or to refrain from pecking (pecks aborted the trial). Once these tasks had been acquired to a high degree, probe trials involving white and blue stimuli were presented at durations between 2 and 10 sec. On trials in which the pigeons were required to peck, the point of subjective equality (i.e., the point at which pigeons are equally likely to choose the stimulus associated with long stimuli as the stimulus associated with short stimuli) was almost 1 sec longer than on trials in which the pigeons were required to refrain from pecking. In other words, on trials that required pecking, more time passed before the pigeons indicated that the probe duration was at the subjective midpoint between 2 and 10 sec than on trials that did not require pecking. This result suggests that like humans, the pigeons underestimated the passage of time when they were active or when attention to time-related cues had to be shared with attention to satisfying the response rate requirement.     

Discrimination of artificial polymorphous categories by rhesus monkeys (Macaca mulatta)

Monkeys (Macaca mulatta) were trained to discriminate between sets of artificial stimuli such as those used by Jitsumori (1993) for pigeons and humans. The stimuli were arrays of symbols differing along three two-valued (positive or negative) dimensions. The discrimination required was between polymorphous categories in which a positive stimulus was defined by possession of any 2 out of 3 positive features. Of the 5 monkeys, 3 learned the discrimination much faster than did pigeons, but transfer to novel stimuli was less impressive than had been shown in pigeons. The 3 monkeys showed high levels of transfer to the stimuli that contained either all 3 positive or all 3 negative features, but 2 of the 3 monkeys failed to show transfer to stimuli that had 1 of the 3 features replaced with a novel one. Analysis of the monkeys' performance raised doubts on the additive integration of features but supported learning of feature combinations as a basis for the discrimination of polymorphous categories by this species.     

Peak Shift on an Artificial Dimension

Blough (1975) proposed an elemental model of generalization and discrimination phenomena in which stimuli from physical dimensions, such as wavelength, are conceptualized as overlapping sets of elements. In order to test predictions generated by this model, we constructed an artificial 'dimension' of stimuli as a series of overlapping sets of arbitrary 'icons' (small, unrelated shapes). In Experiment 1, we trained pigeons to discriminate two neighbouring stimuli from this artificial dimension and then assessed their responses to other stimuli from the dimension. The results of these tests typified those obtained with stimuli from genuine physical dimensions-that is, we obtained positive and negative peak shifts. In Experiment 2, human subjects were given a similar discrimination task and produced an analogous set of results to those of the pigeons. These results support Blough's elemental theory and are not readily accommodated by Pearce's (1987) configural theory. In Experiment 3, both humans and pigeons were trained and tested with stimuli from a real physical dimension (luminance). The pigeons' results were again consistent with Blough's analysis, but those of the humans suggested the use of more sophisticated strategies that are unavailable to pigeons and that lie outside the scope of elemental models of discrimination learning.     

Sunk cost: pigeons (Columba livia), too, show bias to complete a task rather than shift to another

The sunk cost effect involves the bias to stay with an alternative because one has already invested resources, even when there is a better alternative available. In a series of experiments, at various points during a 30-peck requirement, pigeons (Columba livia) could choose between completing the response requirement (at a different location in Experiment 1 or the same location in Experiments 3 and 4) and switching to a constant number of pecks. In three experiments, the pigeons showed a bias to complete the pecks already started, even when that required more pecking. We also demonstrated that the bias depended on the initial investment and was not produced merely because the pigeons preferred a variable alternative over a fixed alternative. The deviation from optimal choice suggests that pigeons show a bias similar to the sunk cost effect in humans.     

Discrimination of five-dimensional stimuli by pigeons: Limitations of feature analysis

Pigeons were trained to discriminate between stimuli constructed using five orthogonal two-valued features. The stimuli consisted of stylized monochrome drawings of seeds. Two different training procedures (conditional and simultaneous discrimination) were used. In the first two experiments, the discrimination required was between polymorphous categories, in which a positive stimulus was defined as one in which three or more of the five features took their positive values. Discrimination in both experiments was imperfect; the pigeons' behaviour only came under the control of a subset of the available features (one to three in Experiment 1, three or four in Experiment 2). In Experiment 3, single features had to be discriminated, while the remaining features varied. It was found that all five features of the “seed” stimuli could be discriminated, but one of them was exceptionally difficult. The results show that pigeons do not reliably use all the features available to them when making category discriminations. This casts doubts on feature analysis as a basis for the excellent performance pigeons show when required to discriminate between categories of natural objects.     

Interference from irrelevant features on visual discrimination by macaques (Macaca fuscata): a behavioral analogue of the human Stroop effect

To study the operation of selective attention in a conflict situation with automatic processes, we trained 4 Japanese macaques (Macaca fuscata) extensively on a manual go/no-go task. The monkey had to discriminate either the color, shape, motion direction, or location of a visual stimulus. In each trial, the behavioral meaning of the relevant feature (go or no-go) could either be congruent or incongruent with irrelevant features of the same stimulus. Reaction times were slowed, and error rates increased when irrelevant stimulus features were incongruent with the required response. The effects were obtained when the monkey attended to the color, shape, or motion direction, but not when it attended to the location of the stimulus. The effects were cumulative so that the interference from 1 incongruent feature was smaller than that from 2 incongruent features. We propose that the present paradigm provides a behavioral analogue of the human Stroop effect.     

Subliminal stimuli in the near absence of attention influence top-down cognitive control

Recent research has shown that visual stimuli can influence cognitive control functions, even if subjects are unaware of the identity of the stimuli. However, in those previous studies, subjects actively attended to the location of the subliminal stimuli. Here we assessed the role of endogenous spatial attention in such paradigms. We required subjects to quickly prepare for one of two numerical judgment tasks on the basis of the direction of motion in patches of moving dots presented in cued spatial locations. We found that irrelevant motion patches presented in the uncued spatial locations also influenced task performance. Motion in the uncued patches was weak and did not affect the perception of the cued patches. Further analyses suggested that the effect of priming by the uncued stimuli was present even for subjects who could only discriminate such stimuli at chance level. Three additional experiments confirmed that subjects paid minimal attention to the uncued locations, in that the subjects could not perform simple discriminations of conjunctions of features in those locations.     

Pigeons learn stimulus identity and stimulus relations when both serve as redundant,relevant cues during same-different discrimination training

The authors taught pigeons to discriminate displays of 16 identical items from displays of 16 nonidentical items. Unlike most same-different discrimination studies--where only stimulus relations could serve a discriminative function--both the identity of the items and the relations among the items were discriminative features of the displays. The pigeons learned about both stimulus identity and stimulus relations when these 2 sources of information served as redundant, relevant cues. In tests of associative competition, identity cues exerted greater stimulus control than relational cues. These results suggest that the pigeon can respond to both specific stimuli and general relations in the environment.     

Discrimination of shape reflections and shape orientations by Columba livia

By using a free-operant instrumental discrimination procedure, it was demonstrated that pigeons find two-dimensional mirror-image visual forms more difficult to distinguish than otherwise similar forms. Variations in orientation of the discriminanda exacerbated the relative confusability of mirror images. No significant difference was found in the pigeons' performance whether the birds were discriminating vertically or horizontally reflected mirror-image pairs. Mirror images of shapes were also shown to be less discriminable than upside-down versions of shapes. The similarity of mirror-image patterns is discussed in relation to the generalized recognition of bilaterally symmetrical forms by pigeons. Pigeons found an orientation discrimination task involving a 45 degree tilt comparatively hard. A second experiment with a discrete-trial conditional paradigm confirmed that discriminations of shape orientations can be difficult for these birds. The addition of shape cues improved the performance on the orientation discrimination task, more so when arbitrary shapes were employed than when mirror images were used, which indicates again that the latter were more difficult to discriminate than the former. The relative insensitivity to shape orientations is ascribed to normal ecological demands on pigeons.     

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.     

The pigeon's discrimination of movement patterns (Lissajous figures) and contour-dependent rotational invariance

The ability of pigeons to discriminate complex motion patterns was investigated with the aid of moving Lissajous figures. The pigeons successfully learned to differentiate two successively presented cyclic trajectories of a single moving dot. This suggests that they can recognize a movement Gestalt when information about shape is minimal. They also quickly learned a new discrimination between moving-outline stimuli with repetitively changing contour patterns. Contrasting results were obtained when the dot or outline stimuli were axis-rotated through 90 degrees. Rotational invariance of pattern discrimination was clearly demonstrated only when moving contours were visible. Nevertheless, pigeons could discriminate the axis-orientation of a moving-dot or moving-outline pattern when trained to do so. Discrimination did not seem to depend on single parameters of motion but rather on the recognition of a temporally integrated movement Gestalt. The visual system of pigeons, as well as that of humans, may be well adapted to recognize the types of oscillatory movements that represent components of the motor behaviour shown by many living organisms.