Learning and organization of within-session sequences by pigeons (Columba livia).

Most animals engage in complex activities that are the combination of simpler actions expressed over a period of time. The mechanisms organizing such sequential behavior have been of long-standing biological and psychological interest. Previously, we observed pigeons' anticipatory behavior with a within-session sequence involving four choice alternatives suggestive of a potential understanding of the overall order and sequence of the items within a session. In that task, each colored alternative was correct for 24 consecutive trials as presented in a predictable sequence (i.e., A first, then B, then C, then D). To test whether these four already-trained pigeons possessed a sequential and linked representation of the ABCD items, we added a second four-item sequence involving new and distinct colored choice alternatives (i.e., E first for 24 trials, then F, then G, then H) and then alternated these ABCD and EFGH sequences over successive sessions. Over three manipulations, we tested and trained trials composed of combinations of elements from both sequences. We determined that pigeons did not learn any within-sequence associations among the elements. Despite the availability and explicit utility of such sequence cues, the data suggest instead that pigeons learned the discrimination tasks as a series of temporal associations among independent elements. This absence of any sequential linkage is consistent with the hypothesis that such representations are difficult to form in pigeons. This pattern of data suggests that for repeated sequential activities in birds, and potentially other animals including humans, there are highly effective, but underappreciated, clock-like mechanisms that control the ordering of behaviors.

Mechanisms of within-session sequential behavior in pigeons.

Correctly and efficiently selecting among options is critical to the organization of behavior across different time scales (minutes, days, seasons). As a result, understanding the mechanisms underlying the sequential behavior of animals has been a long-standing aim. In three experiments, four pigeons were tested in a four-choice simultaneous color discrimination. Across a session, they had to sequentially select a colored stimulus, and the correct color changed over four 24-trial phases (A→B→C→D). After learning this ABCD within-session sequence, tests identified that both timing and outcome feedback mechanisms contributed to the organization of pigeons' behavior. Different representational mechanisms are considered as accounts for the pigeons' observed sequential behavior.

A putative social concept in dolphins.

In playback studies, wild dolphins respond distinctly to group members regardless of individual relationships, implying a mediating associative concept of the group. However, confirming associative concept formation in wild animals is out of our current technological reach, and theories of associative concepts do not account for reciprocal, cooperative relationships which may cause the individual to be a member of the associative concept.

Towards describing scenes by animals: Pigeons' ordinal discrimination of objects varying in depth.

The perception of a complex scene requires visual mechanisms that include identifying objects and their relative placement in depth. To examine apparent depth perception in birds, we tested four pigeons with a novel multiple-sequential-choice procedure. We created 3D-rendered scene stimuli containing three objects located at different apparent depths based on a variety of pictorial cues and placed small circular target response areas on them. The pigeons were trained to sequentially choose among the multiple response areas to report the object closest in apparent depth (ordinal position; front then middle object). After the pigeons learned this sequential depth discrimination, their use of three different monocular depth cues (occlusion, relative size, height in field) was tested, and their flexibility evaluated using three novel objects. In addition to the contribution to understanding apparent depth perception in birds, the use of more flexible open-ended choice discriminations, as employed here, has considerable promise for creating informative production-like tasks in nonverbal animals.

Pigeons process actor-action configurations more readily than bystander-action configurations.

Behavior requires an actor. Two experiments using complex conditional action discriminations examined whether pigeons privilege information related to the digital actor who is engaged in behavior. In Experiment 1, each of two video displays contained a digital model, one an actor engaged in one of two behaviors (Indian dance or martial arts) and one a neutrally posed bystander. To correctly classify the display, the pigeons needed to conditionally process the action in conjunction with distinctive physical features of the actor or the bystander. Four actor-conditional pigeons learned to correctly discriminate the actions based on the identity of the actors, whereas four bystander-conditional birds failed to learn. Experiment 2 established that this failure was not due to the latter group's inability to spatially integrate information across the distance between the two models. Potentially, the colocalization of the relevant model identity and the action was critical due to a fundamental configural or integral representation of these properties. These findings contribute to our understanding of the evolution of action recognition, the recognition of social behavior, and forms of observational learning by animals.

Perception of Ebbinghaus-Titchener stimuli in starlings (Sturnus vulgaris).

Whether animals experience visual illusions is a fertile area of study for examining the evolution and operation of visual cognition across different species. Here, five starlings were tested to examine whether they experienced the Ebbinghaus-Titchener illusion. Across two experiments using an absolute target circle size discrimination, the size, similarity, distance, and number of the surrounding flankers were manipulated. The results suggest that this passerine species exhibits behavior inconsistent with the perception of the illusion, neither in a human-like fashion nor, as suggested by the first experiment, a reversed illusion. Instead, the typical training used to investigate this illusion caused the starlings to learn to integrate the irrelevant flankers into their decision process in a manner that precludes the study of illusory perception. The resulting discriminative behavior might best be described using a template-matching account. While illusion perception by animals remains an important comparative question, it requires additional validation to confirm the exact mechanisms of any illusory reports.

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.

Pigeons discount continuously changing perspective during action recognition.

An important challenge for animal and artificial visual systems is separating the system's own motions from the movements of other animals or events. To examine this issue in birds, we conducted three experiments testing four pigeons in a go/no-go action discrimination. The pigeons discriminated whether a digital human model was exhibiting an extended series of articulated motions or one of a set of static poses from the same video. They were required to do so while the rendering camera's perspective changed continually during each trial's 20-s video presentation. Experiment 1 found that pigeons easily discount the camera's continuous motion. Experiments 2 and 3, by testing novel sequences of the behavior, novel behaviors, silhouettes, and a form of conditional discrimination, revealed this to be a general capacity. Overall, the discrimination was predominantly mediated by global action cues, although a small contribution of image-based statistical features was detected. Collectively, the experiments reveal pigeons can readily separate and discount constantly changing perspectives while processing others' actions. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Dynamically occluded action recognition by pigeons.

Identifying the behaviors of organisms is essential for an animal's survival. This ability is particularly challenged when the "actors" are dynamically occluded by other objects and become fragmented as they move through an environment. Even when fragmented in time and across space, humans readily recognize the behavior of these dynamically occluded objects and actors. How animals process such fragmented information, especially when involving motion, remains uncertain. In three experiments, we investigated the ability of six pigeons to discriminate between the running and walking actions of digital animal models when dynamically occluded. The pigeons were tested in a go/no-go procedure using three models that transited behind multiple occluders in a semirealistic scene. Without ever seeing the entirety of the animal model at one time, all the pigeons learned to discriminate among these two behaviors. This discrimination transferred to an unfamiliar model, transit direction, transiting rates, camera perspectives, and occluders. Tests with different static and dynamic features indicated that the pigeons relied on motion features for the discrimination, especially articulated motion. These experiments demonstrate that pigeons, like humans, can discriminate actions even when their view of the actor is fragmented in time and space.

Modeling within-session dynamics of categorical and item-memory mechanisms in pigeons.

Past studies have shown that pigeons can learn complex categories and can also remember large numbers of individual objects. In recent work, Cook et al. Psychonomic Bulletin & Review, 28, 548-555, (2021) provided evidence that pigeons may use a dynamic combination of both category-based information and item-specific memorization to solve a categorical variation of the mid-session reversal (MSR) task, which is an influential task for exploring the nature of temporally organized behaviors in animals. To provide greater insight into these pigeons' behaviors, in this article we developed and investigated different computational models and their variations to account for these data. Of these, two models emerged as good candidates. One was a multinomial-processing-tree categorization/memory model, formalizing the two-process mechanism initially proposed by Cook et al. Psychonomic Bulletin & Review, 28, 548-555, (2021). The second was a new object/time-coding model, which posits the storage of object-specific memories with an additional within-session time code and assumes that a basic stimulus generalization process underlies the pigeons' choice behavior. Both provided high-quality fits to the published sets of training and transfer data collected in the categorical MSR task. These computational efforts give deeper insights into the theoretical mechanisms underlying the temporal and sequential structure of behavior in animals and stimulate future empirical research further revealing the organization of the pigeons' cognitive processes.

Same/different discrimination of motion by pigeons.

Telling that one object or moment is different from another one is fundamental to cognition and intelligent behavior. Most investigations examining same/different (S/D) concepts in animals have relied on testing static visual stimuli. To move beyond this limitation, we investigated how five pigeons learned and performed a motion S/D discrimination. Using a go/no-go task, dynamic motion fields built from dot elements were presented in sequence to display repeating (same) or changing (different) motions. Each trial consisted of 10 motion segments presented in succession using the direction and rate of dot movement in the motion field to exemplify the S/D relations. The pigeons learned this motion S/D discrimination. We further tested their performance by varying the number and persistence of the dots in the motion fields. The results indicated the pigeons likely extracted globally integrated perceptual summaries of the motions for comparison across the segments. Testing differing organizations of the S/D relations across segments indicated that this discrimination could be determined from as few as two segments and involved an updating comparison of at least four or more segments of the sequence during their presentation. Collectively, the experiments establish for the first time that pigeons can use motion features to classify sequential same and different experiences. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Multiple expressions of "expert" abnormality gist in novices following perceptual learning.

With a brief half-second presentation, a medical expert can determine at above chance levels whether a medical scan she sees is abnormal based on a first impression arising from an initial global image process, termed "gist." The nature of gist processing is debated but this debate stems from results in medical experts who have years of perceptual experience. The aim of the present study was to determine if gist processing for medical images occurs in naïve (non-medically trained) participants who received a brief perceptual training and to tease apart the nature of that gist signal. We trained 20 naïve participants on a brief perceptual-adaptive training of histology images. After training, naïve observers were able to obtain abnormality detection and abnormality categorization above chance, from a brief 500 ms masked presentation of a histology image, hence showing "gist." The global signal demonstrated in perceptually trained naïve participants demonstrated multiple dissociable components, with some of these components relating to how rapidly naïve participants learned a normal template during perceptual learning. We suggest that multiple gist signals are present when experts view medical images derived from the tens of thousands of images that they are exposed to throughout their training and careers. We also suggest that a directed learning of a normal template may produce better abnormality detection and identification in radiologists and pathologists.

Use of different attentional strategies by pigeons and humans in multidimensional visual search.

To study comparative attentional allocation strategies, pigeons and humans were tested using simultaneously available discrimination tasks. Given visual search displays containing 32 items from two orthogonal dimensions, participants were reinforced for selecting the eight brightest (or darkest) of 16 brightness items and the eight most vertical (or horizontal) of 16 orientation items. Consistent with a sequential dimensional strategy, humans preferentially chose items from one dimension before switching to the other to complete the search. In contrast, the pigeons did not preferentially stay within one dimension over consecutive choices. Instead, they chose the items most likely to yield reward based on item discriminability. Computational models that incorporated a "dimensional staying" factor accounted best for the human data, while simulations using only discriminability reproduced the pigeons' data. These results suggest that humans are sensitive to the benefits of attentional staying and the costs of switching between dimensional tasks, while there was no evidence that these factors influenced the pigeons' choice behavior. These findings suggest fundamental differences in how pigeons and humans allocate attention in complex choice situations. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Adaptive testing of the critical features in 2D-shape discrimination by pigeons and starlings.

An innovative adaptive discrimination procedure examined how two bird species, pigeons and starlings, recognize and discriminate two-dimensional (2D) visual shapes. Prior results suggest a comparative divergence between mammals and birds in their relative reliance on vertices versus line segments to mediate discrimination. To address this potentially important difference, four pigeons and five starlings were tested with a square versus triangle discrimination in two experiments. An adaptive genetic algorithm guided the selection and organization of the training and test stimuli. Both species showed considerable flexibility in accurately selecting triangles despite wide variation in stimulus appearance and location. Most critically, Experiment 2 revealed that both bird species relied more on the figures' vertices during successful discrimination than their connecting line segments. This reliance was revealed by both traditional accuracy differences using contour-deleted displays and genetic algorithm-based shifts in "gene values" caused by the birds' selection. These results, in contrast to previous findings, indicate that mammals and birds likely converge in their reliance on vertices as a highly critical feature in visual shape discrimination. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

Within-session dynamics of categorical and memory mechanisms in pigeons.

The current experiments used categorical mid-session reversal (MSR) to examine how eight pigeons utilized categorical and item-specific mechanisms to learn and solve a novel variation of this task. Employing a fixed order of trial-unique pictorial items from two categories (flowers and cars) on each simultaneous discrimination trial, categorical and item-specific information was available during each session's 80 trials. Choices to one category were rewarded for the first 40 trials, after which the correct category was reversed (e.g., car correct early → flower correct late). This procedure selectively impacts the time-modulated utility of categorical identification, but leaves exclusively item-specific information intact. Results revealed that categorical control emerged rapidly and before item-specific memorization, which came after extended experience. Both types of control occurred within a session, with control modulated by their time-based relative utility. The implications for the timing, ordering, and attention by animals to categorical and item-specific information is considered.

Pigeons simultaneously attend to static and dynamic features of complex displays.

The simultaneous processing and attention to temporally dynamic and static features remains an open and important question in theories of avian visual cognition. Here, four pigeons (Columba livia) learned to discriminate complex displays involving concurrently available static and dynamic features. These displays consisted of 20 elements built from combinations of two, binary-valued, static visual dimensions: red vs. green element color, large vs. small element size; and two binary-valued dynamic dimensions; fast vs slow element motion, right vs up motion direction. One combination of these four features was reinforced on a VI schedule. The remaining 15 combinations of element color, size, speed and direction were never reinforced. During acquisition, all four dimensions were simultaneously discriminated. Varying the number of elements revealed that a single element was sufficient to support discrimination of all four dimensions. The pigeons agreed on the relative discriminability of stimuli within and across the different dimensions, with the difference in motion direction being the hardest for all birds. Redundant facilitation suggested rapid, perhaps parallel, processing of both dynamic and static features. No attentional trade-offs between dynamic or static dimensions were observed. These results agree with theories of avian vision employing the notion of multiple independent channels for different types of information.

Testing analogical rule transfer in pigeons (Columba livia).

Categorization is an essential cognitive process useful for transferring knowledge from previous experience to novel situations. The mechanisms by which trained categorization behavior extends to novel stimuli, especially in animals, are insufficiently understood. To understand how pigeons learn and transfer category membership, seven pigeons were trained to classify controlled, bi-dimensional stimuli in a two-alternative forced-choice task. Following either dimensional, rule-based (RB) or information integration (II) training, tests were conducted focusing on the "analogical" extension of the learned discrimination to novel regions of the stimulus space (Casale, Roeder, & Ashby, 2012). The pigeons' results mirrored those from human and non-human primates evaluated using the same analogical task structure, training and testing: the pigeons transferred their discriminative behavior to the new extended values following RB training, but not after II training. Further experiments evaluating rule-based models and association-based models suggested the pigeons use dimensions and associations to learn the task and mediate transfer to stimuli within the novel region of the parametric stimulus space.

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.

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

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.

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.