Olga F. Lazareva

Nonaccidental properties underlie shape recognition in mammalian and nonmammalian vision

An infinite number of 2D patterns on the retina can correspond to a single 3D object. How do visual systems resolve this ill-posed problem and recognize objects from only a few 2D retinal projections in varied exposure conditions? Theories of object recognition rely on the nonaccidental statistics of edge properties, mainly symmetry, collinearity, curvilinearity, and cotermination. These statistics are determined by the image-formation process (i.e., the 2D retinal projection of a 3D object ); their existence under a range of viewpoints enables viewpoint-invariant recognition. An important question in behavioral biology is whether the visual systems of nonmammalian animals have also evolved biases to utilize nonaccidental statistics . Here, we trained humans and pigeons to recognize four shapes. With the Bubbles technique, we determined which stimulus properties both species used to recognize the shapes. Both humans and pigeons used cotermination, the most diagnostic nonaccidental property of real-world objects, despite evidence from a model computer observer that cotermination was not the most diagnostic pictorial information in this particular task. This result reveals that a nonmammalian visual system that is different anatomically from the human visual system is also biased to recognize objects from nonaccidental statistics.

Pigeons concurrently categorize photographs at both basic and superordinate levels

We studied categorization in pigeons, using carefully controlled photographs. Within daily sessions, 4 pigeons had to classify each of 32 photographs into either its proper basic-level category (cars, chairs, flowers, or people; four-key forced choice procedure) or its proper superordinate-level category (natural or artificial; two-key forced choice procedure). The pigeons successfully classified the same stimuli at both levels. Overall, the pigeons learned the basic discrimination more quickly than the superordinate discrimination, but this difference was reliable only for artificial stimuli (cars and chairs), not for natural stimuli (flowers and people). The pigeons also exhibited reliable discrimination transfer to novel photographs, attesting to the open-endedness of these basic and superordinate categories.

Pigeons and humans are more sensitive to nonaccidental than to metric changes in visual objects.

Humans and macaques are more sensitive to differences in nonaccidental image properties, such as straight vs. curved contours, than to differences in metric properties, such as degree of curvature [Biederman, I., Bar, M., 1999. One-shot viewpoint invariance in matching novel objects. Vis. Res. 39, 2885-2899; Kayaert, G., Biederman, I., Vogels, R., 2003. Shape tuning in macaque inferior temporal cortex. J. Neurosci. 23, 3016-3027; Kayaert, G., Biederman, I., Vogels, R., 2005. Representation of regular and irregular shapes in macaque inferotemporal cortex. Cereb. Cortex 15, 1308-1321]. This differential sensitivity allows facile recognition when the object is viewed at an orientation in depth not previously experienced. In Experiment 1, we trained pigeons to discriminate grayscale, shaded images of four shapes. Pigeons made more confusion errors to shapes that shared more nonaccidental properties. Although the images in that experiment were not well controlled for incidental changes in metric properties, the same results were apparent with better controlled stimuli in Experiment 2: pigeons trained to discriminate a target shape from a metrically changed shape and a nonaccidentally changed shape committed more confusion errors to the metrically changed shape, suggesting that they perceived it to be more similar to the target shape. Humans trained with similar stimuli and procedure exhibited the same tendency to make more errors to the metrically changed shape. These results document the greater saliency of nonaccidental differences for shape recognition and discrimination in a non-primate species and suggest that nonaccidental sensitivity may be characteristic of all shape-discriminating species.

Multiple-pair training enhances transposition in pigeons.

We studied transposition in pigeons by training them to select the smaller (or the larger) of a pair of circles. In training, different groups of pigeons were given one pair, two pairs, or three pairs of circles along the size dimension. Testing included two stimulus pairs for which, according to theoretical postdiscrimination generalization gradients, transposition should decrease from one-pair to two-pair to three-pair training. On the basis of the results of our earlier study (Lazareva, Wasserman, & Young, 2005) and contrary to these predictions, we expected that transposition should increase from one-pair to two-pair to three-pair training. We found that multiple-pair discrimination training enhanced transposition, which, on average, rose from 47% (one-pair training) to 52% (two-pair training) to 64% (three-pair training). In addition, we found that the overall similarity of the testing pair to the training pair(s) modulated the strength of relational responding. These results demonstrate that encountering multiple instances of a rule leads to stronger relational learning, even when reinforcement history predicts the opposite trend. These results also provide strong evidence against stimulus generalization as the sole determinant of relational responding in transposition.

Transposition in Pigeons: Reassessing spence (1937) with multiple discrimination training

We studied transposition in pigeons, using multiple-pair discrimination training. Four birds discriminated two pairs of circles: 1+ 2− and 5+ 6− or 1− 2+ and 5− 6+ (digits denote circle diameters and plus and minus signs denote reward and nonreward, respectively). Four other birds discriminated four pairs of circles: 1+2−, 1+3−, 4+6−, and 5+6−or 1−2+, 1−3+, 4−6+, and 5−6+. Finally, 4 birds discriminated only one pair of circles: 1+ 2−, 1+ 2−, 5+ 6−, or 5− 6+. Testing included five new pairs—1/5, 2/3, 2/6, 3/4, and 4/5—that distinguished absolute from relational accounts of transposition. The pigeons’ relational responding rose from one- to two- to four-pair training. The similarity of the testing stimuli to one another also affected relational responding: Transposition increased with highly dissimilar stimuli. Neither Spence’s (1937) theory nor existing relational accounts could predict the obtained pattern of relational responding.

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.

Nonverbal transitive inference: Effects of task and awareness on human performance

We studied human nonverbal transitive inference in two paradigms: with choice stimuli orderable along a physical dimension and with non-orderable choice stimuli. We taught 96 participants to discriminate four overlapping pairs of choice stimuli: A+ B-, B+ C-, C+ D-, and D+ E-. Half of the participants were provided with post-choice visual feedback stimuli which were orderable by size; the other half were not provided with orderable feedback stimuli. In later testing, we presented novel choice pairs: BD, AC, AD, AE, BE, and CE. We found that transitive responding depended on task awareness for all participants. Additionally, participants given ordered feedback showed clearer task awareness and stronger transitive responding than did participants not given ordered feedback. Associative models (Wynne, 1995; Siemann and Delius, 1998) failed to predict the increase in transitive responding with increasing awareness. These results suggest that ordered and non-ordered transitive inference tasks support different patterns of performance.

Effect of between-category similarity on basic-level superiority in pigeons

Children categorize stimuli at the basic level faster than at the superordinate level. We hypothesized that between-category similarity may affect this basic level superiority effect. Dissimilar categories may be easy to distinguish at the basic level but be difficult to group at the superordinate level, whereas similar categories may be easy to group at the superordinate level but be difficult to distinguish at the basic level. Consequently, similar basic level categories may produce a superordinate-before-basic learning trend, whereas dissimilar basic level categories may result in a basic-before-superordinate learning trend. We tested this hypothesis in pigeons by constructing superordinate level categories out of basic level categories with known similarity. In Experiment 1, we experimentally evaluated the between-category similarity of four basic level photographic categories using multiple fixed interval-extinction training (Astley and Wasserman, 1992). We used the resultant similarity matrices in Experiment 2 to construct two superordinate level categories from basic level categories with high between-category similarity (cars and persons; chairs and flowers). We then trained pigeons to concurrently classify those photographs into either the proper basic level category or the proper superordinate level category. Under these conditions, the pigeons learned the superordinate level discrimination faster than the basic level discrimination, confirming our hypothesis that basic level superiority is affected by between-category similarity.

Object discrimination by pigeons: effects of object color and shape

Can nonhuman animals attend to visual stimuli as whole, coherent objects? We investigated this question by adapting for use with pigeons a task in which human participants must report whether two visual attributes belong to the same object (one-object trial) or to different objects (two-object trial). We trained pigeons to discriminate a pair of differently colored shapes that had two targets either on a single object or on two different objects. Each target equally often appeared on the one-object and two-object stimuli; therefore, a specific target location could not serve as a discriminative cue. The pigeons learned to report whether the two target dots were located on a single object or on two different objects; follow-up tests demonstrated that this ability was not entirely based on memorization of the dot patterns and locations. Additional tests disclosed predominate stimulus control by the color, but not by the shape of the two objects. These findings suggest that human psychophysical methods are readily applicable to the study of object discrimination by nonhuman animals.

Hippocampal lesion and transitive inference: Dissociation of inference-based and reinforcement-based strategies in pigeons

A typical nonverbal transitive inference task (TI) consists of several overlapping discriminations (A+ B−, B+ C−, C+ D−, D+ E−, where letters indicate stimuli and pluses and minuses denote reinforcement and nonreinforcement). A choice of stimulus B in a novel pair BD is interpreted as indicative of a TI (if B > C and C > D, then B > D). Although hippocampus has been implicated in nonverbal TI, it is not clear whether it simply maintains memory of associative values or stores an ordered representation of stimuli. We investigated the effect of hippocampal lesion on TI in pigeons while controlling reinforcement history so that reliance on associative values would lead to a choice of a stimulus D in the pair BD instead of a choice of a stimulus B expected by inferential mechanisms. Prior to the lesion, some of the pigeons (relational group; n = 4) have selected B over D indicating TI, while other birds (associative group; n = 6) chose D over B suggesting reliance on associative values. Hippocampal lesion had no effect on postlesion performance in associative group. In contrast, the relational group that preferred stimulus B in a pair BD before lesion showed a near‐chance performance after the lesion. Our results demonstrate that hippocampus may be involved in creating a representation of an ordered series of the stimuli instead of maintaining reinforcement history of each stimulus. In addition, we provide a behavioral procedure suitable for dissociating different behavioral strategies used to solving TI task. Finally, we show for the first time the involvement of avian hippocampus in the task that is not explicitly spatial in nature. These results further confirm the notion that avian hippocampus is functionally analogous to mammalian hippocampus despite the significant differences in their anatomy.