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.     

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.     

Recognizing rotated views of objects: Interpolation versus generalization by humans and pigeons

Pigeons and humans were trained to discriminate between pictures of three-dimensional objects that differed in global shape. Each pair of objects was shown at two orientations that differed by a depth rotation of 90° during training. Pictures of the objects at novel depth rotations were then tested for recognition. The novel test rotations were 30°, 45°, and 90° from the nearest trained orientation and were either interpolated between the trained orientations or extrapolated outside of the training range. For both pigeons and humans, recognition accuracy and/or speed decreased as a function of distance from the nearest trained orientation. However, humans, but not pigeons, were more accurate in recognizing novel interpolated views than novel extrapolated views. The results suggest that pigeons’ recognition was based on independent generalization from each training view, whereas humans showed view-combination processes that resulted in a benefit for novel views interpolated between the training views.     

Symmetry brings an impression of familiarity but does not improve recognition memory

Studies have shown that symmetric stimuli are recognized better than asymmetric stimuli but evidence suggests that this advantage may result from a familiarity bias induced by symmetry. We used a classic episodic memory paradigm to test this bias and see if it truly accounts for the symmetry advantage. Subjects first encoded symmetric and asymmetric figures. During a subsequent recognition phase, they discriminated the encoded (old) figures from new intermixed figures. The recognition rate of old figures was higher with symmetric figures than asymmetric figures. However, the tendency to falsely recognize new figures was also higher when they were symmetric, meaning that the higher recognition rate for symmetric figures was artificially inflated by a response bias. Three other experiments further tested this finding and examined the influence of some variables (rotation in virtual 3D space, stimulus meaningfulness, and redundancy of information) on the bias. A fifth experiment with photo stimuli confirmed that the response bias also applies to objects that we regularly encounter in everyday life. In conclusion, our results show that symmetry does not enhance mnemonic processes but instead induces a response bias leading individuals to judge such stimuli as having been seen.     

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 effect of distinctive parts on recognition of depth-rotated objects by pigeons (Columba livia) and humans.

To explore whether effects observed in human object recognition represent fundamental properties of visual perception that are general across species, the authors trained pigeons (Columba livia) and humans to discriminate between pictures of 3-dimensional objects that differed in shape. Novel pictures of the depth-rotated objects were then tested for recognition. Across conditions, the object pairs contained either 0, 1, 3, or 5 distinctive parts. Pigeons showed viewpoint dependence in all object-part conditions, and their performance declined systematically with degree of rotation from the nearest training view. Humans showed viewpoint invariance for novel rotations between the training views but viewpoint dependence for novel rotations outside the training views. For humans, but not pigeons, viewpoint dependence was weakest in the 1-part condition. The authors discuss the results in terms of structural and multiple-view models of object recognition.     

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.     

Visual and haptic discrimination of symmetry in unfamiliar displays extended in the z-axis

We investigated, in two experiments, the discrimination of bilateral symmetry in vision and touch using four sets of unfamiliar displays. They varied in complexity from 3 to 30 turns. Two sets were 2-D flat forms (raised-line shapes and raised surfaces) while the other two were 3-D objects constructed by extending the 2-D shapes in height (short and tall objects). Experiment 1 showed that visual accuracy was excellent but latencies increased for raised-line shapes compared with 3-D objects. Experiment 2 showed that unimanual exploration was more accurate for asymmetric than for symmetric judgments, but only for 2-D shapes and short objects. Bimanual exploration at the body midline facilitated the discrimination of symmetric shapes without changing performance with asymmetric ones. Accuracy for haptically explored symmetric stimuli improved as the stimuli were extended in the third dimension, while no such a trend appeared for asymmetric stimuli. Unlike vision, haptic response latency decreased for 2-D shapes compared with 3-D objects. The present results are relevant to the understanding of symmetry discrimination in vision and touch.     

The effect of vertical and horizontal symmetry on memory for tactile patterns in late blind individuals

Visual stimuli that exhibit vertical symmetry are easier to remember than stimuli symmetric along other axes, an advantage that extends to the haptic modality as well. Critically, the vertical symmetry memory advantage has not been found in early blind individuals, despite their overall superior memory, as compared with sighted individuals, and the presence of an overall advantage for identifying symmetric over asymmetric patterns. The absence of the vertical axis memory advantage in the early blind may depend on their total lack of visual experience or on the effect of prolonged visual deprivation. To disentangle this issue, in this study, we measured the ability of late blind individuals to remember tactile spatial patterns that were either vertically or horizontally symmetric or asymmetric. Late blind participants showed better memory performance for symmetric patterns. An additional advantage for the vertical axis of symmetry over the horizontal one was reported, but only for patterns presented in the frontal plane. In the horizontal plane, no difference was observed between vertical and horizontal symmetric patterns, due to the latter being recalled particularly well. These results are discussed in terms of the influence of the spatial reference frame adopted during exploration. Overall, our data suggest that prior visual experience is sufficient to drive the vertical symmetry memory advantage, at least when an external reference frame based on geocentric cues (i.e., gravity) is adopted.     

Associative symmetry by pigeons after few-exemplar training

The present experiment investigated whether pigeons can show associative symmetry on a two-alternative matching-to-sample procedure. The procedure consisted of a within-subject sequence of training and testing with reinforcement, and it provided (a) exemplars of symmetrical responding, and (b) all prerequisite discriminations among test samples and comparisons. After pigeons had learned two arbitrary-matching tasks (A-B and C-D), they were given a reinforced symmetry test for half of the baseline relations (B1-A1 and D1-C1). To control for the effects of reinforcement during testing, two novel, nonsymmetrical responses were concurrently reinforced using the other baseline stimuli (D2-A2 and B2-C2). Pigeons matched at chance on both types of relations, thus indicating no evidence for symmetry. These symmetrical and nonsymmetrical relations were then directly trained in order to provide exemplars of symmetry and all prerequisite discriminations for a second test. The symmetrical test relations were now B2-A2 and D2-C2 and the nonsymmetrical relations were D1-A1 and B1-C1. On this test, 1 pigeon showed clear evidence of symmetry, 2 pigeons showed weak evidence, and 1 pigeon showed no evidence. The previous training of all prerequisite discriminations among stimuli, and the within-subject control for testing with reinforcement seem to have set favorable conditions for the emergence of symmetry in nonhumans. However, the variability across subjects shows that methodological variables still remain to be controlled.     

Categorical Discrimination of Human Facial Expressions by Pigeons: A Test of the Linear Feature Model

Pigeons were trained to discriminate human facial expressions, happiness and anger, in a go/ no-go discrimination procedure. Five pigeons learned to discriminate photographs of the happy and angry faces of 25 different people and showed high levels of transfer to novel faces expressing the training emotions. The pigeons directed their pecks predominantly to the mouth, eyes, or the area between these features. The pigeons were then tested with familiar stimuli in which the upper and lower parts of the face were manipulated separately by substitution or removal of facial features ('eyes-and-eyebrows' and 'mouth'). It was shown that the salience of particular features differed considerably among the birds, but that a linear feature model adequately accounted for discriminative performance of the birds with these stimuli. Furthermore, the discrimination was maintained when these features were inverted. Thus, the so-called Thatcher illusion did not occur. It is suggested that the discrimination is based not on a feature configuration or perceptual gestalt but on an additive integration of individual features.     

Discrimination of moving video images of self by pigeons (<Emphasis Type="Italic">Columba livia</Emphasis>)

The ability to recognize self has been known to be limited to some animal species, but previous research has focused almost exclusively on the animal's reaction to a mirror. Recent studies suggest that the temporal contingency between a subject's action and the corresponding visual scene reflected in a mirror plays an important role in self-recognition. To assess the roles of visual-proprioceptive contiguity in self-recognition, we explored whether pigeons are able to discriminate videos of themselves with various temporal properties. We trained five pigeons to respond to live video images of themselves (live self-movies) and not to video filmed during previous training sessions (pre-recorded self-movies). Pigeons learned to peck trial-unique live self-movies more frequently than pre-recorded self-movies. We conducted two generalization tests after pigeons learned to discriminate between the two conditions. First, discrimination acquired during training sessions was transferred to a test session involving live self-movies and new pre-recorded self-movies. Second, the same pigeons were tested in extinction procedure using delayed live self-movies and new pre-recorded self-movies. Although pigeons responded to delayed presentations of live self-movies more frequently than to new pre-recorded self-movies, the relative response rate to delayed presentation of live self-movies gradually decreased as the temporal discrepancy between pigeons' own behavior and the corresponding video increased. These results indicate that pigeons' discrimination of self-movies with various temporal properties was based on the temporal contiguity between their behavior and its visual feedback. The methodology used in the present experiment is an important step toward improving the experimental analysis of self-recognition in non-human animals.     

Independent feature processing in the visual system

Summary A recognition experiment with a set of simple visual patterns was performed. The patterns were composed of three binary features: Lines, angles and curve segments. The pattern sets differed with respect to the distance between components and the symmetry of the arrangement. The empirical confusion matrices were analysed assuming two different models: A model assuming independent feature analysis and a model that assumes that one of the features is analysed independently of the other two. The main result was that quite a large distance between features was necessary to achieve independent processing of the features. Moreover, in the asymmetric pattern set the predictions of the independence model were better than in the symmetric pattern set.     

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.     

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.     

Discrimination of intentional and random motion paths by pigeons

Twelve pigeons (Columba livia) were trained on a go/no-go schedule to discriminate between two kinds of movement patterns of dots, which to human observers appear to be "intentional" and "non-intentional" movements. In experiment 1, the intentional motion stimulus contained one dot (a "wolf") that moved systematically towards another dot as though stalking it, and three distractors ("sheep"). The non-intentional motion stimulus consisted of four distractors but no stalker. Birds showed some improvement of discrimination as the sessions progressed, but high levels of discrimination were not reached. In experiment 2, the same birds were tested with different stimuli. The same parameters were used but the number of intentionally moving dots in the intentional motion stimulus was altered, so that three wolves stalked one sheep. Despite the enhanced difference of movement patterns, the birds did not show any further improvement in discrimination. However, birds for which the non-intentional stimulus was associated with reward showed a decline in discrimination. These results indicated that pigeons can discriminate between stimuli that do and do not contain an element that human observer see as moving intentionally. However, as no feature-positive effect was found in experiment 1, it is assumed that pigeons did not perceive or discriminate these stimuli on the basis that the intentional stimuli contained a feature that the non-intentional stimuli lacked, though the convergence seen in experiment 2 may have been an effective feature for the pigeons. Pigeons seem to be able to recognise some form of multiple simultaneously goal-directed motions, compared to random motions, as a distinctive feature, but do not seem to use simple "intentional" motion paths of two geometrical figures, embedded in random motions, as a feature whose presence or absence differentiates motion displays. Electronic Publication     

Generalization of visual regularities in newly hatched chicks (<Emphasis Type="Italic">Gallus gallus</Emphasis>)

Evidence of learning and generalization of visual regularities in a newborn organism is provided in the present research. Domestic chicks have been trained to discriminate visual triplets of simultaneously presented shapes, implementing AAB versus ABA (Experiment 1), AAB versus ABB and AAB versus BAA (Experiment 2). Chicks distinguished pattern-following and pattern-violating novel test triplets in all comparisons, showing no preference for repetition-based patterns. The animals generalized to novel instances even when the patterns compared were not discriminable by the presence or absence of reduplicated elements or by symmetry (e.g., AAB vs. ABB). These findings represent the first evidence of learning and generalization of regularities at the onset of life in an animal model, revealing intriguing differences with respect to human newborns and infants. Extensive prior experience seems to be unnecessary to drive the process, suggesting that chicks are predisposed to detect patterns characterizing the visual world.     

Local rather than global processing of visual arrays in numerosity discrimination by pigeons (Columba livia)

A series of experiments investigated which stimulus properties pigeons use when they discriminate pairs of visual arrays that differ in numerosity. Transfer tests with novel stimuli confirmed that the birds’ choices were based on relative differences in numerosity. However, pigeons differed from other species in the non-numerical cues that affected their choices. In human and non-human primates, numerical discrimination is often influenced by continuous variables such as surface area or overall stimulus brightness. Pigeons showed little evidence of using those cues, even when summed area and brightness had been correlated with numerosity differences and reward outcome. But when array-element sizes were asymmetrically distributed across numerosities, the birds readily utilized information about item sizes as an additional discriminative cue. These novel results are discussed in relation to pigeons’ tendency to focus on local, rather than global dimensions when they process other non-numerical complex visual stimuli. The findings suggest there may be inter-specific differences in the type of perceptual information that provides the input stage for mechanisms underlying numerical processing. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

European starlings are capable of discriminating subtle size asymmetries in paired stimuli

Small deviations from bilateral symmetry (fluctuating asymmetries) are cues to fitness differences in some animals. Therefore, researchers have considered whether animals use these small asymmetries as visual cues to determine appropriate behavioral responses (e.g., mate preferences). However, there have been few systematic studies of animals' abilities to visually discriminate such minor asymmetries. If the asymmetries cannot be discriminated, fluctuating asymmetry can not be a visual cue. Here, we report an investigation of European starlings' (Sturnus vulgaris) abilities to discriminate small size asymmetries. We trained starlings, through operant conditioning in a free-flight aviary, to discriminate achromatic, symmetric paired stimuli from size-matched asymmetric stimuli. By starting the learning process with a large asymmetry and progressing through sequential trials of decreasing asymmetry, we elucidated a behavioral limit to asymmetry discrimination. We found that starlings are capable of discriminating a 10% size asymmetry. There was weaker evidence for discrimination of 5% asymmetry but no evidence for signal discrimination at 2.5% size asymmetry. This level of asymmetry discrimination suggests that many size asymmetry cues in nature can be discriminated by birds. At each level of asymmetry discrimination, we also tested whether starlings could generalize their learned symmetry preference to unreinforced novel images. Consistent with previous findings, we found that starlings could generalize their symmetry preferences.     

Motion as a natural category for pigeons: Generalization and a feature-positive effect

Three groups of pigeons were trained with a modified discriminative autoshaping procedure to discriminate video images of other pigeons on the basis of movement. Birds of all groups were shown the same video images of other pigeons, which were either moving or still. The group to whom food was presented only after moving images learned the discrimination very quickly. A second group, to whom food was given only after still images, and a pseudocategory group, to whom food was presented after arbitrarily chosen stimuli, showed no evidence of discrimination during acquisition training. Extinction conditions led to clear differences in peck rates to moving and still images in the second group but not in the pseudocategory group. The result is related to the feature-positive effect. Generalization tests showed that the discrimination performance was based on visual features of the stimuli but was invariant against changes of size, perspective, brightness, and color. Furthermore, discrimination was maintained when novel images of pigeons under different viewing angles and seven other types of motion categories were presented. It is argued that the discrimination is based not on a common motion feature but on motion concepts or high-order generalization across motion categories.