Muhammad A. J. Qadri

Shape from shading in pigeons

Light is the origin of vision. The pattern of shading reflected from object surfaces is one of several optical features that provide fundamental information about shape and surface orientation. To understand how surface and object shading is processed by birds, six pigeons were tested with differentially illuminated convex and concave curved surfaces in five experiments using a go/no-go procedure. We found that pigeons rapidly learned this type of visual discrimination independent of lighting direction, surface coloration and camera perspective. Subsequent experiments varying the pattern of the lighting on these surfaces through changes in camera perspective, surface height, contrast, material specularity, surface shape, light motion, and perspective movement were consistent with the hypothesis that the pigeons were perceiving these illuminated surfaces as three-dimensional surfaces containing curved shapes. The results suggest that the use of relative shading for objects in a visual scene creates highly salient features for shape processing in birds.

Visual control of an action discrimination in pigeons

Recognizing and categorizing behavior is essential for all animals. The visual and cognitive mechanisms underlying such action discriminations are not well understood, especially in nonhuman animals. To identify the visual bases of action discriminations, four pigeons were tested in a go/no-go procedure to examine the contribution of different visual features in a discrimination of walking and running actions by different digital animal models. Two different tests with point-light displays derived from studies of human biological motion failed to support transfer of the learned action discrimination from fully figured models. Tests with silhouettes, contours, and the selective deletion or occlusion of different parts of the models indicated that information about the global motions of the entire model was critical to the discrimination. This outcome, along with earlier results, suggests that the pigeons’ discrimination of these locomotive actions involved a generalized categorization of the sequence of configural poses. Because the motor systems for locomotion and flying in pigeons share little in common with quadruped motions, the pigeons’ discrimination of these behaviors creates problems for motor theories of action recognition based on mirror neurons or related notions of embodied cognition. It suggests instead that more general motion and shape mechanisms are sufficient for making such discriminations, at least in birds.

Shape from shading in starlings (Sturnus vulgaris).

Birds behave as if they quickly and accurately perceive an object-filled visual world. Beyond the extensive research with pigeons, however, there is a large and important gap in our knowledge about the mechanisms of object perception and recognition in other avian visual systems. The pattern of shading reflected from the surfaces of objects is one important optical feature that provides fundamental information about shape. To better understand how surface and object shading is processed by a passerine species, 5 starlings were tested with differentially illuminated convex and concave curved surfaces in 3 experiments using a simultaneous visual discrimination procedure. Starlings rapidly learned this shape-from-shading discrimination independent of varied lighting direction, surface color, and camera perspective. Variations in the pattern of lighting through experimental manipulations of camera perspective, surface height, contrast, material specularity, and surface shape were consistent with the hypothesis that the starlings perceived these illuminated surfaces as having 3-dimensional shape, similar to results previously collected with pigeons. These similarities across different orders of birds indicate that the relative shading of objects in a visual scene is a highly salient feature for shape processing in birds and is likely a highly conserved visual process that is widely distributed within this class of animal.

Discrimination of Complex Human Behavior by Pigeons (Columba livia) and Humans

The cognitive and neural mechanisms for recognizing and categorizing behavior are not well understood in non-human animals. In the current experiments, pigeons and humans learned to categorize two non-repeating, complex human behaviors (“martial arts” vs. “Indian dance”). Using multiple video exemplars of a digital human model, pigeons discriminated these behaviors in a go/no-go task and humans in a choice task. Experiment 1 found that pigeons already experienced with discriminating the locomotive actions of digital animals acquired the discrimination more rapidly when action information was available than when only pose information was available. Experiments 2 and 3 found this same dynamic superiority effect with naïve pigeons and human participants. Both species used the same combination of immediately available static pose information and more slowly perceived dynamic action cues to discriminate the behavioral categories. Theories based on generalized visual mechanisms, as opposed to embodied, species-specific action networks, offer a parsimonious account of how these different animals recognize behavior across and within species.

The Perception of Glass Patterns by Starlings (Sturnus vulgaris)

Glass patterns are structured dot stimuli used to investigate the visual perception of global form. Studies have demonstrated that humans and pigeons differ in their processing of circular versus linearly organized Glass patterns. To test whether this comparative difference is characteristic of birds as a phylogenetic class, we investigated for the first time how a passerine (starlings, Sturnus vulgaris) discriminated multiple Glass patterns from random-dot stimuli in a simultaneous discrimination. By examining acquisition, steady-state performance, and the effects of diminishing global coherence, it was found that the perception of Glass patterns by 5 starlings differed from human perception and corresponded to that established with pigeons. This suggests an important difference in how birds and primates are specialized in their processing of circular visual patterns, perhaps related to face perception, or in how these highly visual animals direct attention to the global and local components of spatially separated form stimuli.

Pigeons and humans use action and pose information to categorize complex human behaviors

Abstract The biological mechanisms used to categorize and recognize behaviors are poorly understood in both human and non‐human animals. Using animated digital models, we have recently shown that pigeons can categorize different locomotive animal gaits and types of complex human behaviors. In the current experiments, pigeons (go/no‐go task) and humans (choice task) both learned to conditionally categorize two categories of human behaviors that did not repeat and were comprised of the coordinated motions of multiple limbs. These “martial arts” and “Indian dance” action sequences were depicted by a digital human model. Depending upon whether the model was in motion or not, each species was required to engage in different and opposing responses to the two behavioral categories. Both species learned to conditionally and correctly act on this dynamic and static behavioral information, indicating that both species use a combination of static pose cues that are available from stimulus onset in addition to less rapidly available action information in order to successfully discriminate between the behaviors. Human participants additionally demonstrated a bias towards the dynamic information in the display when re‐learning the task. Theories that rely on generalized, non‐specific visual mechanisms involving channels for motion and static cues offer a parsimonious account of how humans and pigeons recognize and categorize behaviors within and across species.

Visualizing search behavior with adaptive discriminations

We examined different aspects of the visual search behavior of a pigeon using an open-ended, adaptive testing procedure controlled by a genetic algorithm. The animal had to accurately search for and peck a gray target element randomly located from among a variable number of surrounding darker and lighter distractor elements. Display composition was controlled by a genetic algorithm involving the multivariate configuration of different parameters or genes (number of distractors, element size, shape, spacing, target brightness, and distractor brightness). Sessions were composed of random displays, testing randomized combinations of these genes, and selected displays, representing the varied descendants of displays correctly identified by the pigeon. Testing a larger number of random displays than done previously, it was found that the birds solution to the search task was highly stable and did not change with extensive experience in the task. The location and shape of this attractor was visualized using multivariate behavioral surfaces in which element size and the number of distractors were the most important factors controlling search accuracy and search time. The resulting visualizations of the birds search behavior are discussed with reference to the potential of using adaptive, open-ended experimental techniques for investigating animal cognition and their implications for Bond and Kamils innovative development of virtual ecologies using an analogous methodology. This article is part of a Special Issue entitled: CO3 2013.

Dynamic cue use in pigeon mid-session reversal

The systematic anticipation and preservation errors produced by pigeons around the reversal point in midsession reversal (MSR) learning experiments suggest that an internal time estimation cue, instead of a more efficient external cue provided by reinforcement, controls behavior over the course of a session. The current experiments examined the role and effectiveness of other external cues in the MSR task. In Experiment 1, providing differential outcomes based on response key location produced fewer errors prior to, but not after, the reversal as compared with a non-differential outcomes condition. Experiment 2a used alternating differentially colored ITIs (cued sessions) or dark ITIs (un-cued sessions) during each half of the session. The ITI cues improved switch efficiency both prior to and after the reversal. Experiment 2b introduced probe trials around the reversal, testing ITI color cues added to un-cued sessions or removed from cued sessions. Results showed control by the ITI cues when they were available and control by the time-based cue when they were unavailable. This suggests both cues were being simultaneously processed when available and that the cues could also independently provide sufficient information about future reinforcement. In Experiment 2c, ITI cues were inserted as probe trials in the opposite half of the session (miscues). The closer such miscue trials were to the reversal, the more the ITI cues exerted control over behavior. Together, these results indicate that as the utility of internal temporal cues is reduced, the use of external visual cues increases. These results have implications for the way in which cues dynamically shift in controlling behavior over time based on their relative rates of utility, and are discussed in light of an occasion setting perspective.

Examination of long-term visual memorization capacity in the Clark’s nutcracker (Nucifraga columbiana)

Clark’s nutcrackers exhibit remarkable cache recovery behavior, remembering thousands of seed locations over the winter. No direct laboratory test of their visual memory capacity, however, has yet been performed. Here, two nutcrackers were tested in an operant procedure used to measure different species’ visual memory capacities. The nutcrackers were incrementally tested with an ever-expanding pool of pictorial stimuli in a two-alternative discrimination task. Each picture was randomly assigned to either a right or a left choice response, forcing the nutcrackers to memorize each picture–response association. The nutcrackers’ visual memorization capacity was estimated at a little over 500 pictures, and the testing suggested effects of primacy, recency, and memory decay over time. The size of this long-term visual memory was less than the approximately 800-picture capacity established for pigeons. These results support the hypothesis that nutcrackers’ spatial memory is a specialized adaptation tied to their natural history of food-caching and recovery, and not to a larger long-term, general memory capacity. Furthermore, despite millennia of separate and divergent evolution, the mechanisms of visual information retention seem to reflect common memory systems of differing capacities across the different species tested in this design.

Detection and discrimination of complex sounds by pigeons (Columba livia)

Auditory scene analysis is the process by which sounds are separated and identified from each other and from the background to make functional auditory objects. One challenge in making these psychological units is that complex sounds often continuously differ in composition over their duration. Here we examined the acoustic basis of complex sound processing in four pigeons by evaluating their performance in an ongoing same/different (S/D) task. This provided an opportunity to investigate avian auditory processing in a non-vocal learning, non-songbird. These pigeons were already successfully discriminating 18.5 s sequences of all different 1.5 s sounds (ABCD…) from sequences of one sound repeating (AAAA…, BBBB…, etc.) in a go/no-go procedure. The stimuli for these same/different sequences consisted of 504 tonal sounds (36 chromatic notes×14 different instruments), 36 pure tones, and 72 complex sounds. Not all of these sounds were equally effective in supporting S/D discrimination. As identified by a stepwise regression modeling of ten acoustic properties, tonal and complex sounds with intermediate levels of acoustic content tended to support better discrimination. The results suggest that pigeons have the auditory and cognitive capabilities to recognize and group continuously changing sound elements into larger functional units that can serve to differentiate long sequences of same and different sounds.