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Same/different abstract-concept learning experiments were conducted with two primate species and three avian species by progressively increasing the size of the training stimulus set of distinctly different pictures from eight to 1,024 pictures. These same/different learning experiments were trained with two pictures presented simultaneously. Transfer tests of same and different learning employed interspersed trials of novel pictures to assess the level of correct performance on the very first time of subjects had seen those pictures. All of the species eventually performed these tests with high accuracy, contradicting the long-accepted notion that nonhuman animals are unable to learn the concept of same/different. Capuchin and rhesus monkeys learned the concept more readily than did pigeons. Clark's nutcrackers and black-billed magpies learned as readily as monkeys, and even showed a slight advantage with the smallest training stimulus sets. Those tests of same/different learning were followed by delay procedures, such that a delay was introduced after the subjects responded to the sample picture and before the test picture. In the sequential same/different task, accuracy was shown to diminish when the stimulus on a previous trial matched the test picture previously shown on a different trial. This effect is known as proactive interference. The pigeons' proactive interference was greater at 10-s delays than 1-s delays, revealing time-based interference. By contrast, time delays had little or no effect on rhesus monkeys' proactive interference, suggesting that rhesus monkeys have better explicit memory of where and when they saw the potential interfering picture, revealing better event-based memory.

Judgements of items viewed less than 100 ms prior are predominantly supported by a sensory, or iconic, memory system. Iconic memory is of high-capacity, but is also volatile and limited in duration. Judgements after longer delays increasingly rely on a working memory system, which is lower in capacity and volatility than sensory memory, but is longer in duration. In four experiments, several factors (e.g., length of delay, number of items, time to view items, presence of a visual mask) were manipulated during a spatial change-detection task conducted with humans and pigeons. Both species were exposed to trials with an array of colored circles (2, 3, and 4 circles in Experiment 1 and 2a; 4, 6, and 8 circles in Experiment 2b) followed by a brief delay (0, 50, and 100 ms in Experiment 1a; 0, 100, and 1000 ms in Experiments 1b and 2), and then were presented with a test display in which the position of one of the items had changed. Pigeons, like humans, were less accurate in selecting the changed item with more items in the display and after longer delays. Pigeons were equally accurate on trials with 0 and 100-ms delays, but worse on trials with a 1000-ms delay; whereas, humans were equally accurate on 100-ms and 1000-ms delays, but better on 0-ms delay trials. Accurate change detection was disrupted in both species when a visual mask was inserted between the sample and test display after a short (100 ms), but not a long (1000 ms) delay. The results support similarity between species in the functional relationships between delay and memory systems, despite time course differences related to sensory memory., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Rats exposed to variable-length, unique-odor lists were tested in distinctive contexts for odors second or forth from list-end. Accurate ability to recall odors backwards from the end of lists points to their ability to manipulate and replay odor-list episodic memories., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

This article describes an approach for training a variety of species to learn the abstract concept of same/different, which in turn forms the basis for testing proactive interference and list memory. The stimulus set for concept-learning training was progressively doubled from 8, 16, 32, 64, 128 . . . to 1,024 different pictures with novel-stimulus transfer following learning. All species fully learned the same/different abstract concept: capuchin and rhesus monkeys learned more readily than pigeons; nutcrackers and magpies were at least equivalent to monkeys and transferred somewhat better following initial training sets. A similar task using the 1,024-picture set plus delays was used to test proactive interference on occasional trials. Pigeons revealed greater interference with 10-s than with 1-s delays, whereas delay time had no effect on rhesus monkeys, suggesting that the monkeys' interference was event based. This same single-item same/different task was expanded to a 4-item list memory task to test animal list memory. Humans were tested similarly with lists of kaleidoscope pictures. Delays between the list and test were manipulated, resulting in strong initial recency effects (i.e., strong 4th-item memory) at short delays and changing to a strong primacy effect (i.e., strong 1st-item memory) at long delays (pigeons 0-s to 10-s delays; monkeys 0-s to 30-s delays; humans 0-s to 100-s delays). Results and findings are discussed in terms of these species' cognition and memory comparisons, evolutionary implications, and future directions for testing other species in these synergistically related tasks.

This article describes an approach for assessing and comparing complex cognition in rhesus monkeys and pigeons by training them in a sequence of synergistic tasks, each yielding a whole function for enhanced comparisons. These species were trained in similar same/different tasks with expanding training sets (8, 16, 32, 64, 128 … 1024 pictures) followed by novel-stimulus transfer eventually resulting in full abstract-concept learning. Concept-learning functions revealed better rhesus transfer throughout and full concept learning at the 128 set, versus pigeons at the 256 set. They were then tested in delayed same/different tasks for proactive interference by inserting occasional tests within trial-unique sessions where the test stimulus matched a previous sample stimulus (1, 2, 4, 8, 16 trials prior). Proactive-interference functions revealed time-based interference for pigeons (1, 10 s delays), but event-based interference for rhesus (no effect of 1, 10, 20 s delays). They were then tested in list-memory tasks by expanding the sample to four samples in trial-unique sessions (minimizing proactive interference). The four-item, list-memory functions revealed strong recency memory at short delays, gradually changing to strong primacy memory at long delays over 30 s for rhesus, and 10 s for pigeons. Other species comparisons and future directions are discussed., (© 2018 Society for the Experimental Analysis of Behavior.)

Since sensory measurements are noisy, an observer is rarely certain about the identity of a stimulus. In visual perception tasks, observers generally take their uncertainty about a stimulus into account when doing so helps task performance. Whether the same holds in visual working memory tasks is largely unknown. Ten human and two monkey subjects localized a single change in orientation between a sample display containing three ellipses and a test display containing two ellipses. To manipulate uncertainty, we varied the reliability of orientation information by making each ellipse more or less elongated (two levels); reliability was independent across the stimuli. In both species, a variable-precision encoding model equipped with an "uncertainty-indifferent" decision rule, which uses only the noisy memories, fitted the data poorly. In both species, a much better fit was provided by a model in which the observer also takes the levels of reliability-driven uncertainty associated with the memories into account. In particular, a measured change in a low-reliability stimulus was given lower weight than the same change in a high-reliability stimulus. We did not find strong evidence that observers took reliability-independent variations in uncertainty into account. Our results illustrate the importance of studying the decision stage in comparison tasks and provide further evidence for evolutionary continuity of working memory systems between monkeys and humans.

Corvids (birds of the family Corvidae) display intelligent behavior previously ascribed only to primates, but such feats are not directly comparable across species. To make direct species comparisons, we used a same/different task in the laboratory to assess abstract-concept learning in black-billed magpies ( Pica hudsonia). Concept learning was tested with novel pictures after training. Concept learning improved with training-set size, and test accuracy eventually matched training accuracy-full concept learning-with a 128-picture set; this magpie performance was equivalent to that of Clark's nutcrackers (a species of corvid) and monkeys (rhesus, capuchin) and better than that of pigeons. Even with an initial 8-item picture set, both corvid species showed partial concept learning, outperforming both monkeys and pigeons. Similar corvid performance refutes the hypothesis that nutcrackers' prolific cache-location memory accounts for their superior concept learning, because magpies rely less on caching. That corvids with "primitive" neural architectures evolved to equal primates in full concept learning and even to outperform them on the initial 8-item picture test is a testament to the shared (convergent) survival importance of abstract-concept learning.

relational concepts depend upon relationships between stimuli (e.g., same vs. different) and transcend features of the training stimuli. Recent evidence shows that learning abstract concepts is shared across a variety species including birds. Our recent work with a highly-skilled food-storing bird, Clark's nutcracker, revealed superior same/different abstract-concept learning compared to rhesus monkeys, capuchin monkeys, and pigeons. Here we test a more social, but less reliant on food-storing, corvid species, the Black-billed magpie (Pica hudsonia). We used the same procedures and training exemplars (eight pairs of the same rule, and 56 pairs of the different rule) as were used to test the other species. Magpies (n = 10) showed a level of abstract-concept learning that was equivalent to nutcrackers and greater than the primates and pigeons tested with these same exemplars. These findings suggest that superior initial abstract-concept learning abilities may be shared across corvids generally, rather than confined to those strongly reliant on spatial memory.

Previous work in discrimination learning has shown that nonmatching (oddity) tasks are learned faster and more accurately than comparable matching tasks. This learning advantage has been coined the oddity preference effect (Wright & Delius in Journal of Experimental Psychology: Animal Behavior Processes, 31, 425-432. doi: 10.1037/0097-7403.31.4.425 , 2005). Pigeons trained in a nonmatching task, following training in a same/different (S/D) task, learned the abstract concept of difference (Daniel et al., in Animal Cognition, 18(4), 831-837, 2015), but they did not show the expected faster acquisition or high levels of transfer from the oddity preference effect. In the present study, experimentally naïve pigeons were trained in an identical nonmatching task to examine whether they would show the oddity preference effect on abstract-concept learning. These experimentally naïve pigeons did show an oddity preference effect; their transfer to novel configurations was above chance with the initial (smallest) set size (3-item set) and was substantially more accurate than novel transfer in similar match-to-sample (MTS) or S/D tasks (Bodily et al., in Journal of Experimental Psychology: Animal Behavior Processes, 34, 178-184. doi: 10.1037/0097-7403.34.1.178 , 2008; Katz & Wright in Journal of Experimental Psychology: Animal Behavior Processes, 32, 80-86. doi: 10.1037/0097-7403.32.1.80 , 2006). As the number exemplars in the training set increased, transfer to novel configurations increased and reached equivalence to trained-stimulus performance with a 24-item set. Despite this transfer being equal to baseline performance with a 24-item set, subsequent transfers following training with larger set sizes declined before eventually rising again to baseline performance. This unusual set-size function (with inflection points at the 24- and 96-set sizes) suggests that these pigeons may have combined item-specific and relational learning strategies with differing emphasis as they acquired the abstract concept.

Three rhesus monkeys (Macaca mulatta) were tested in a same/different memory task for proactive interference (PI) from prior trials. PI occurs when a previous sample stimulus appears as a test stimulus on a later trial, does not match the current sample stimulus, and the wrong response "same" is made. Trial-unique pictures (scenes, objects, animals, etc.) were used on most trials, except on trials where the test stimulus matched potentially interfering sample stimulus from a prior trial (1, 2, 4, 8, or 16 trials prior). Greater interference occurred when fewer trials separated interference and test. PI functions showed a continuum of interference. Delays between sample and test stimuli and intertrial intervals were manipulated to test how PI might vary as a function of elapsed time. Contrary to a similar study with pigeons, these time manipulations had no discernable effect on the monkey's PI, as shown by compete overlap of PI functions with no statistical differences or interactions. These results suggested that interference was strictly based upon the number of intervening events (trials with other pictures) without regard to elapsed time. The monkeys' apparent event-based interference was further supported by retesting with a novel set of 1,024 pictures. PI from novel pictures 1 or 2 trials prior was greater than from familiar pictures, a familiar set of 1,024 pictures. Moreover, when potentially interfering novel stimuli were 16 trials prior, performance accuracy was actually greater than accuracy on baseline trials (no interference), suggesting that remembering stimuli from 16 trials prior was a cue that this stimulus was not the sample stimulus on the current trial-a somewhat surprising conclusion particularly given monkeys.

Three pigeons were trained to remember arrays of 2-6 colored squares and detect which of two squares had changed color to test their visual short-term memory. Procedures (e.g., stimuli, displays, viewing times, delays) were similar to those used to test monkeys and humans. Following extensive training, pigeons performed slightly better than similarly trained monkeys, but both animal species were considerably less accurate than humans with the same array sizes (2, 4 and 6 items). Pigeons and monkeys showed calculated memory capacities of one item or less, whereas humans showed a memory capacity of 2.5 items. Despite the differences in calculated memory capacities, the pigeons' memory results, like those from monkeys and humans, were all well characterized by an inverse power-law function fit to d' values for the five display sizes. This characterization provides a simple, straightforward summary of the fundamental processing of visual short-term memory (how visual short-term memory declines with memory load) that emphasizes species similarities based upon similar functional relationships. By closely matching pigeon testing parameters to those of monkeys and humans, these similar functional relationships suggest similar underlying processes of visual short-term memory in pigeons, monkeys and humans., (Copyright © 2015 Elsevier B.V. All rights reserved.)

Same/Different abstract-concept learning by Clark's nutcrackers (Nucifraga columbiana) was tested with novel stimuli following learning of training set expansion (8, 16, 32, 64, 128, 256, 512, and 1024 picture items). The resulting set-size function was compared to those from rhesus monkeys (Macaca mulatta), capuchin monkeys (Cebus apella), and pigeons (Columba livia). Nutcrackers showed partial concept learning following initial eight-item set learning, unlike the other species (Magnotti, Katz, Wright, & Kelly, 2015). The mean function for the nutcrackers' novel-stimulus transfer increased linearly as a function of the logarithm of training set size, which intersected its baseline function at the 128-item set size. Thus, nutcrackers on average achieved full concept learning (i.e., transfer statistically equivalent to baseline performance) somewhere between set sizes of 64 to 128 items, similar to full concept learning by monkeys. Pigeons required a somewhat larger training set (256 items) for full concept learning, but results from other experiments (initial training and transfer with 32- and 64-item set sizes) suggested carryover effects with smaller set sizes may have artificially prolonged the pigeon's full concept learning. We find it remarkable that these diverse species with very different neural architectures can fully learn this same/different abstract concept, and (at least under some conditions) do so with roughly similar sets sizes (64-128 items) and numbers of training exemplars, despite initial concept learning advantages (nutcrackers), learning disadvantages (pigeons), or increasing baselines (monkeys)., (© 2015 Society for the Experimental Analysis of Behavior.)

Many species have demonstrated the capacity to learn abstract concepts. Recent studies have shown that the quantity of stimuli used during training plays a critical role in how subjects learn abstract concepts. As the number of stimuli available in the training set increases, so too does performance on novel combinations. The role of set size has been explored with learning the concept of matching and same/different but not with learning the concept of difference. In the present study, pigeons were trained in a non-matching-to-sample task with an initial training set of three stimuli followed by transfer tests to novel stimuli. The training set was progressively doubled eight times with learning and transfer following each expansion. Transfer performance increased from chance level (50 %) at the smallest set size to a level equivalent to asymptotic training performance at the two largest training set sizes (384, 768). This progressive novel-stimulus transfer function of a non-matching (difference) rule is discussed in comparison with results from a similar experiment where pigeons were trained on a matching rule.

The ability to learn abstract relational concepts is fundamental to higher level cognition. In contrast to item-specific concepts (e.g. pictures containing trees versus pictures containing cars), abstract relational concepts are not bound to particular stimulus features, but instead involve the relationship between stimuli and therefore may be extrapolated to novel stimuli. Previous research investigating the same/different abstract concept has suggested that primates might be specially adapted to extract relations among items and would require fewer exemplars of a rule to learn an abstract concept than non-primate species. We assessed abstract-concept learning in an avian species, Clark's nutcracker (Nucifraga columbiana), using a small number of exemplars (eight pairs of the same rule, and 56 pairs of the different rule) identical to that previously used to compare rhesus monkeys, capuchin monkeys and pigeons. Nutcrackers as a group (N = 9) showed more novel stimulus transfer than any previous species tested with this small number of exemplars. Two nutcrackers showed full concept learning and four more showed transfer considerably above chance performance, indicating partial concept learning. These results show that the Clark's nutcracker, a corvid species well known for its amazing feats of spatial memory, learns the same/different abstract concept better than any non-human species (including non-human primates) yet tested on this same task., (© 2015 The Author(s) Published by the Royal Society. All rights reserved.)

Rhesus monkeys are widely used as an animal model for human memory, including visual working memory (VWM). It is, however, unknown whether the same principles govern VWM in humans and rhesus monkeys. Here, we tested both species in nearly identical change-localization paradigms and formally compared the same set of models of VWM limitations. These models include the classic item-limit model and recent noise-based (resource) models, as well as hybrid models that combine a noise-based representation with an item limit. By varying the magnitude of the change in addition to the typical set size manipulation, we were able to show large differences in goodness of fit among the five models tested. In spite of quantitative performance differences between the species, we find that the variable-precision model--a noise-based model--best describes the behavior of both species. Adding an item limit to this model does not help to account for the data. Our results suggest evolutionary continuity of VWM across primates and help establish the rhesus monkey as a model system for studying the neural substrates of multiple-item VWM.

Two adult rhesus monkeys were trained to detect which item in an array of memory items had changed using the same stimuli, viewing times, and delays as used with humans. Although the monkeys were extensively trained, they were less accurate than humans with the same array sizes (2, 4, & 6 items), with both stimulus types (colored squares, clip art), and showed calculated memory capacities of about 1 item (or less). Nevertheless, the memory results from both monkeys and humans for both stimulus types were well characterized by the inverse power-law of display size. This characterization provides a simple and straightforward summary of a fundamental process of visual short-term memory (STM; how VSTM declines with memory load) that emphasizes species similarities based upon similar functional relationships. By more closely matching monkey testing parameters to those of humans, the similar functional relationships strengthen the evidence suggesting similar processes underlying monkey and human VSTM., (PsycINFO Database Record (c) 2015 APA, all rights reserved.)

Change detection is commonly used to assess capacity (number of objects) of human visual short-term memory (VSTM). Comparisons with the performance of non-human animals completing similar tasks have shown similarities and differences in object-based VSTM, which is only one aspect ("what") of memory. Another important aspect of memory, which has received less attention, is spatial short-term memory for "where" an object is in space. In this article, we show for the first time that a monkey and pigeons can be accurately trained to identify location changes, much as humans do, in change detection tasks similar to those used to test object capacity of VSTM. The subject's task was to identify (touch/peck) an item that changed location across a brief delay. Both the monkey and pigeons showed transfer to delays longer than the training delay, to greater and smaller distance changes than in training, and to novel colors. These results are the first to demonstrate location-change detection in any non-human species and encourage comparative investigations into the nature of spatial and visual short-term memory.

Content-specific sub-systems of visual working memory (VWM) have been explored in many neuroimaging studies with inconsistent findings and procedures across experiments. The present study employed functional magnetic resonance imaging (fMRI) and a change detection task using a high number of trials and matched stimulus displays across object and location change (what vs. where) conditions. Furthermore, individual task periods were studied independently across conditions to identify differences corresponding to each task period. Importantly, this combination of task controls has not previously been described in the fMRI literature. Composite results revealed differential frontoparietal activation during each task period. A separation of object and location conditions yielded a distributed system of dorsal and ventral streams during the encoding of information corresponding to bilateral inferior parietal lobule (IPL) and lingual gyrus activation, respectively. Differential activity was also shown during the maintenance of information in middle frontal structures bilaterally for objects and the right IPL and left insula for locations. Together, these results reflect a domain-specific dissociation spanning several cortices and task periods. Furthermore, differential activations suggest a general caudal-rostral separation corresponding to object and location memory, respectively.

A recent study using a novel procedure to test the memory of rats for a preferred (chocolate) reinforcement shows many key characteristics that define source memory and episodic memory in humans., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Identifying critical features that control categorization of complex polymorphous pictures by animals remains a challenging and important problem. Toward this goal, experiments were conducted to isolate the properties controlling the categorization of two pictorial categories by pigeons. Pigeons were trained in a go/no-go task to categorize black and white line drawings of birds and mammals. They were then tested with a variety of familiar and novel exemplars of these categories to examine the features controlling this categorization. These tests suggested the pigeons were segregating and using the principal axis of orientation of the animal figures as the primary means of discriminating each category, although other categorical and item-specific cues were likely involved. This perceptual/cognitive reduction of the categorical stimulus space to a few visual features or dimensions is likely a characteristic of this species' processing of complex pictorial discrimination problems and is a critical property for theoretical accounts of this behavior., (Copyright © 2012. Published by Elsevier B.V.)

Functional relationships (from systematic manipulation of critical variables) are advocated for revealing fundamental processes of (comparative) cognition-through examples from my work in psychophysics, learning, and memory. Functional relationships for pigeon wavelength (hue) discrimination revealed best discrimination at the spectral points of hue transition for pigeons-a correspondence (i.e., functional relationship) similar to that for humans. Functional relationships for learning revealed: Item-specific or relational learning in matching to sample as a function of the pigeons' sample-response requirement, and same/different abstract-concept learning as a function of the training set size for rhesus monkeys, capuchin monkeys, and pigeons. Functional relationships for visual memory revealed serial position functions (a 1st order functional relationship) that changed systematically with retention delay (a 2nd order relationship) for pigeons, capuchin monkeys, rhesus monkeys, and humans. Functional relationships for rhesus-monkey auditory memory also revealed systematic changes in serial position functions with delay, but these changes were opposite to those for visual memory. Functional relationships for proactive interference revealed interference that varied as a function of a ratio of delay times. Functional relationships for change detection memory revealed (qualitative) similarities and (quantitative) differences in human and monkey visual short-term memory as a function of the number of memory items. It is concluded that these findings were made possible by varying critical variables over a substantial portion of the manipulable range to generate functions and derive relationships., (Copyright © 2012 Elsevier B.V. All rights reserved.)

We examined the role of the hippocampus in list-memory processing. Three rhesus monkeys that had extensive experience in this task and had demonstrated full abstract-concept learning and excellent list memory performance (Katz et al., 2002; Wright et al., 2003) received bilateral neurotoxic hippocampal lesions and were re-tested in the serial list memory task. Effects of delays on memory performance were measured in all monkeys, whereas the effects of proactive interference were assessed in only one. Despite a slight change in performance of one of the three animals during re-learning of the same/different task, selective hippocampal damage had little or no effects on list memory accuracy. In addition, the hippocampal damage did not impact serial list position functions (SPFs) but slightly altered the dynamic of the SPF curves. Finally, even more remarkable was that accurate memory performance of one animal remained intact despite the use of small set size of 8 items that created high proactive interference across lists thereby eliminating any use of familiarity judgments to support performance. Together the findings indicate that, with short list items and extensive training in the task (i.e., reference memory), monkeys with selective hippocampal lesions may be able to use alternative memory processes (i.e., working memory) that are mediated by structures other than the hippocampus., (Copyright © 2012 Elsevier B.V. All rights reserved.)

Seven adult human participants were tested in change detection tasks for object and location memory with large and small sets of four different stimulus types. Blocked tests demonstrated that participants performed similarly in separate object and location tests with matched parameters and displays. In mixed tests, participants were informed that they would be tested with either object changes or location changes; surprisingly, they were nearly as accurate remembering both objects and locations as when either was tested alone. By contrast, in the large-set condition, performance was lower than baseline on surprise probe test trials in which participants were tested (on 13% of trials) with the change type opposite to the present block (e.g., location probe trials during the object change block). These probe-test results were further supported by the reduction in probe-baseline differences when tested with small sets (6) of these item types. Small sets required remembering locations and objects to resolve object-location confounds. Together these results show that humans can remember both objects and locations with little loss of accuracy when instructed to do so, but do not learn these contextual associations without instruction., (Copyright © 2012 Elsevier B.V. All rights reserved.)

The study of visual memory has repeatedly shown qualitatively similar visual short-term memory (VSTM) systems between human and many nonhuman species. In studies of human VSTM using change detection, increasing visual object complexity has an inverse effect on accuracy. In the current study, we assessed the functional relationship between visual object complexity and memory performance in visual change detection in pigeons and humans. Visual object complexity was quantified for each object type within each species using visual target search. Change detection performance was inversely related to object complexity in both species, suggesting that pigeon VSTM, like human VSTM, is limited by visual object complexity. Human participants were able to use a verbal-labeling strategy to mitigate some of the effect of visual object complexity, suggesting a qualitative difference in how the two species may solve certain visual discriminations. Considering the visual complexity of novel objects may also help explain previous failures to transfer relational rules to novel visual objects., (Copyright © 2012 Elsevier B.V. All rights reserved.)

Two monkeys (Macaca mulatta) learned a color change-detection task where two colored circles (selected from a 4-color set) were presented on a 4 × 4 invisible matrix. Following a delay, the correct response was to touch the changed colored circle. The monkeys' learning, color transfer, and delay transfer were compared to a similar experiment with pigeons. Monkeys, like pigeons (Columba livia), showed full transfer to four novel colors, and to delays as long as 6.4 s, suggesting they remembered the colors as opposed to perceptual based attentional capture process that may work at very short delays. The monkeys and pigeons were further tested to compare transfer with other dimensions. Monkeys transferred to shape and location changes, unlike the pigeons, but neither species transferred to size changes. Thus, monkeys were less restricted in their domain to detect change than pigeons, but both species learned the basic task and appear suitable for comparative studies of visual short-term memory., (2012 APA, all rights reserved)

Processes of proactive interference were explored using the pigeon as a model system of memory. This study shows that proactive interference extends back in time at least 16 trials (and as many minutes), revealing a continuum of interference and providing a framework for studying memory. Pigeons were tested in a delayed same/different task containing trial-unique pictures. On interference trials, sample pictures from previous trials reappeared as test pictures on different trials. Proactive-interference functions showed greatest interference from the most recent trial and with the longer of two delays (10 s vs. 1 s). These interference functions are accounted for by a time-estimation model based on signal detection theory. The model predicts that accuracy at test is determined solely by the ratio of the elapsed time since the offset of the current-trial sample to the elapsed time since the offset of the interfering sample. Implications for comparing memory of different species and different types of memory (e.g., familiarity vs. recollection) are discussed.

This article has been withdrawn at the request of the authors. The Publisher apologizes for any inconvenience this may cause. The full Elsevier Policy on Article Withdrawal can be found at http://www.elsevier.com/locate/withdrawalpolicy. This article has been withdrawn at the request of the editor. The Publisher apologizes for any inconvenience this may cause. The full Elsevier Policy on Article Withdrawal can be found at http://www.elsevier.com/locate/withdrawalpolicy., (Copyright © 2011. Published by Elsevier Inc. All rights reserved.)

Change detection is a popular task to study visual short-term memory (STM) in humans [1-4]. Much of this work suggests that STM has a fixed capacity of 4 ± 1 items [1-6]. Here we report the first comparison of change-detection memory between humans and a species closely related to humans, the rhesus monkey. Monkeys and humans were tested in nearly identical procedures with overlapping display sizes. Although the monkeys' STM was well fit by a one-item fixed-capacity memory model, other monkey memory tests with four-item lists have shown performance impossible to obtain with a one-item capacity [7]. We suggest that this contradiction can be resolved using a continuous-resource approach more closely tied to the neural basis of memory [8, 9]. In this view, items have a noisy memory representation whose noise level depends on display size as a result of the distributed allocation of a continuous resource. In accord with this theory, we show that performance depends on the perceptual distance between items before and after the change, and d' depends on display size in an approximately power-law fashion. Our results open the door to combining the power of psychophysics, computation, and physiology to better understand the neural basis of STM., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Pigeons, capuchin monkeys and rhesus monkeys were trained in nearly identical same/different tasks with an expanding 8-item training set and showed qualitatively similar functional relationships: increasing novel-stimulus transfer (i.e., concept learning) as a function of the training-set size and the level of transfer eventually becoming equivalent to baseline training performance. There were also some quantitative functional differences: pigeon transfer increases were more gradual and baseline-equivalent transfer occurred at a larger set size (256 items) than for monkeys (128 items). Other pigeon groups trained at 32 and 64-item initial set sizes showed improved transfer (relative to expanding the 8-item training set), equivalent to the monkey species' transfer at these same training set sizes. This finding of equivalent concept learning over a portion of the functional range (8, 32, and 64 items or 64-4096 training pairs) is discussed in terms of species differences: carryover effects from smaller-set training, evolved neural systems, cognitive and cortical modules, and general distributed learning systems for "higher-order" cognitive abilities., (Copyright © 2010 Elsevier B.V. All rights reserved.)

Commonly recognized, the training procedure one employs often affects the results they obtain. Here, we demonstrate for the first time that abstract-concept learning is affected by employing a differential-outcomes procedure. The differential-outcome effect has been shown to occur for item-specific strategies but has not been established for relational strategies. To test whether different-outcome expectancies can facilitate a relational strategy, eight pigeons were trained and tested in a two-item same/different task with pictures. After pecking an upper picture, they pecked a lower picture if the pictures were the same or a white rectangle if the pictures were different. Two groups of pigeons were rewarded with either different outcomes (sounds and food amounts) or same outcomes. Both groups were trained to criterion with successively larger picture sets (8-1,024 items) and were transfer tested with novel pictures following each acquisition. With the smallest training sets, neither group showed any novel-stimulus transfer. But after acquiring the task with 32 pictures, the different-outcomes group responded more accurately to novel pictures than the same-outcome group. As the training set-size increased, both groups' transfer performance converged and became equivalent to training performance. These results show for the first time that training with different outcomes facilitates abstract-concept learning.

Six pigeons were trained in a change detection task with four colors. They were shown two colored circles on a sample array, followed by a test array with the color of one circle changed. The pigeons learned to choose the changed color and transferred their performance to four unfamiliar colors, suggesting that they had learned a generalized concept of color change. They also transferred performance to test delays several times their 50-msec training delay without prior delay training. The accurate delay performance of several seconds suggests that their change detection was memory based, as opposed to a perceptual attentional capture process. These experiments are the first to show that an animal species (pigeons, in this case) can learn a change detection task identical to ones used to test human memory, thereby providing the possibility of directly comparing short-term memory processing across species.

The translational-symmetry hypothesis of abstract-concept learning was tested in a same/different (S/D) task with pairs of pictures. The translational-symmetry hypothesis proposes that subjects discriminate same trials by the simultaneous repetition of features in the two pictures (and different trials by the lack of feature repetition). Pigeons that had learned a simultaneous S/D task were tested with delays between the two pictures to remove emergent perceptual cues. In Experiment 1, we tested delays of 0 and 1 sec. The results did not show the accuracy decrease expected according to the translational-symmetry hypothesis. In Experiment 2, we expanded the delays to 2 and 6 sec. Even at the longest delay, there was no evidence of the precipitous performance decline or default strategy that would be predicted by translational symmetry. The results provide evidence against translational symmetry (or other perceptually emergent features) that might control these pigeons' performance in our two-item S/D task.

The domain for relational learning was manipulated by varying the training set size for pigeons that had learned the same/different (S/D) concept. Six pigeons that had learned a S/D task with pairs of pictures with a set size of 1024 picture items had their training set size reduced to 8 items. Training on the reduced 8-item set was followed by transfer testing that was repeated four times. Transfer performance following reduction of the training set to 8 items was 9.2% less than it had been when the pigeons were trained with the 1024-item set, but 25.8% above chance. This partial abstract-concept learning remained constant over the four tests with novel stimuli. The results show that a broad domain established by a large expanding training set can once again become restricted by further training with a small training set., (Copyright 2009 Elsevier B.V. All rights reserved.)

Monkeys and pigeons learned a same/different task with pairs that were selected from a training set of eight picture stimuli. They showed no novel-stimulus transfer and hence no abstract-concept learning. They were also tested with novel pairs of the eight training pictures (i.e., combinations that had not been used in training) and with inverted pictures of the training pairs. If the subjects had learned the task item-specifically (e.g., if-then or configural learning), they should have failed these tests, but they performed well with novel combinations of training pictures and inverted pictures, suggesting that they learned the task relationally (i.e., on the basis of the relationship between the two pictures that were presented in each trial). This somewhat paradoxical conclusion of relational learning in the absence of abstract-concept learning is contrary to most theories of abstract-concept learning. The implications of this conclusion are discussed in the context of restricted-domain relational learning.

Three pigeons were trained in a three-item simultaneous same/different task. Three of six stimulus combinations were not trained (untrained set) and were tested later. Following acquisition, the subjects were tested with novel stimuli, the untrained set, training-stimulus inversions, and object shape and color manipulations. There was no novel-stimulus transfer--that is, no abstract-concept learning. Two pigeons showed partial transfer to untrained pairs and good transfer to stimulus inversions, suggesting that they had learned the relationship between the stimuli. Lack of transfer by the third pigeon suggests item-specific learning. The somewhat surprising finding of relational learning by 2 pigeons with only six training pairs suggests restricted-domain relational learning that was controlled more by color than by shape features. Individual differences of item-specific learning by 1 pigeon and relational learning by 2 others demonstrate that this task can be learned in different ways and that relational learning can occur in the absence of novel-stimulus transfer.

Three groups of pigeons were trained in a same/different task with 32, 64, or 1,024 color-picture stimuli. They were tested with novel transfer pictures. The training-testing cycle was repeated with training-set doublings. The 32-item group learned the same/different task as rapidly as a previous 8-item group and transferred better than the 8-item group at the 32-item training set. The 64- and 1,024-item groups learned the task only somewhat slower than other groups, but their transfer was better and equivalent to baseline performances. These results show that pigeons trained with small sets (e.g., 8 items) have carryover effects that hamper transfer when the training set is expanded. Without carryover effects (i.e., initial transfer from the 32- and 64-item groups), pigeons show the same degree of transfer as rhesus and capuchin monkeys at these same set sizes. This finding has implications for the general ability of abstract-concept learning across species with different neural architectures., (Copyright 2009 APA, all rights reserved.)

Drawings depicting familiar objects and unreal structures were presented twice, and participants (N=16) determined whether line drawings were real (familiar) or unreal (unfamiliar). The second presentation (repetition) of a drawing was typically responded to faster and more accurately than the first presentation and was accompanied by reduced activation in occipitotemporal (fusiform) and lateral precuneus regions, and increased activation in medial precuneus regions. The behavioral effects and reduced activations (e.g., lateral precuneus) on the second presentation were less pronounced for unreal objects than for real objects. Activation changes in the medial precuneus - increased activation on repetition and reduced activation for novel unreal objects - was further supported by the increased activation in this area during rest and reduced activation when workload was increased (i.e., processing novel unreal objects). The results from the present study in conjunction with those from several previous studies converge on the conclusion that the occipitotemporal and lateral regions of the precuneus are primarily involved in object priming, whereas the medial portion of precuneus primarily activates and deactivates as a function of workload.

Pigeons learned a matching-to-sample task with a split training-set design in which half of the stimulus displays were untrained and tested following acquisition. Transfer to the untrained displays along with no novel-stimulus transfer indicated that these pigeons learned the task (partially) via if-then rules. Comparisons to other performance measures indicated that they also partially learned the task via configural learning (learning the gestalt of the whole stimulus display). Differences in the FR-sample requirement (1 vs. 20) had no systematic effect on the type of learning or level of learning obtained. Differences from a previous study [Wright, A.A., 1997. Concept learning and learning strategies. Psychol. Sci. 8, 119-123] are discussed, including the effect of displaying the stimuli vertically (traditional display orientation) or horizontally from the floor.

Abstract concepts--rules that transcend training stimuli--have been argued to be unique to some species. Pigeons, a focus of much concept-learning research, were tested for learning a matching-to-sample abstract concept. Five pigeons were trained with three cartoon stimuli. Pigeons pecked a sample 10 times and then chose which of two simultaneously presented comparison stimuli matched the sample. After acquisition, abstract-concept learning was tested by presenting novel cartoons on 12 out of 96 trials for 4 consecutive sessions. A cycle of doubling the training set followed by retraining and novel-testing was repeated eight times, increasing the set size from 3 to 768 items. Transfer performance improved from chance (i.e., no abstract-concept learning) to a level equivalent to baseline performance (>80%) and was similar to an equivalent function for same/different abstract-concept learning. Analyses assessed the possibility that item-specific choice strategies accounted for acquisition and transfer performance. These analyses converged to rule out item-specific strategies at all but the smallest set-sizes (3-24 items). Ruling out these possibilities adds to the evidence that pigeons learned the relational abstract concept of matching-to-sample., (Copyright (c) 2008 APA, all rights reserved.)

The generalization hypothesis of abstract-concept learning was tested with a meta-analysis of rhesus monkeys (Macaca mulatta), capuchin monkeys (Cebus apella), and pigeons (Columba livia) learning a same/different (S/D) task with expanding training sets. The generalization hypothesis states that as the number of training items increases, generalization from the training pairs will increase and could explain the subjects' accurate novel-stimulus transfer. By contrast, concept learning is learning the relationship between each pair of items; with more training items subjects learn more exemplars of the rule and transfer better. Having to learn the stimulus pairs (the generalization hypothesis) would require more training as the set size increases, whereas learning the concept might require less training because subjects would be learning an abstract rule. The results strongly support concept or rule learning despite severely relaxing the generalization-hypothesis parameters. Thus, generalization was not a factor in the transfer from these experiments, adding to the evidence that these subjects were learning the S/D abstract concept., (Copyright 2007 APA.)

Rhesus monkeys were trained and tested in visual and auditory list-memory tasks with sequences of four travel pictures or four natural/environmental sounds followed by single test items. Acquisitions of the visual list-memory task are presented. Visual recency (last item) memory diminished with retention delay, and primacy (first item) memory strengthened. Capuchin monkeys, pigeons, and humans showed similar visual-memory changes. Rhesus learned an auditory memory task and showed octave generalization for some lists of notes--tonal, but not atonal, musical passages. In contrast with visual list memory, auditory primacy memory diminished with delay and auditory recency memory strengthened. Manipulations of interitem intervals, list length, and item presentation frequency revealed proactive and retroactive inhibition among items of individual auditory lists. Repeating visual items from prior lists produced interference (on nonmatching tests) revealing how far back memory extended. The possibility of using the interference function to separate familiarity vs. recollective memory processing is discussed.

-concept learning, including same/different and matching-to-sample concept learning, provides the basis for many other forms of "higher" cognition. The issue of which species can learn abstract concepts and the extent to which abstract-concept learning is expressed across species is discussed. Definitive answers to this issue are argued to depend on the subjects' learning strategy (e.g., a relational-learning strategy) and the particular procedures used to test for abstract-concept learning. Some critical procedures that we have identified are: How to present the items to-be-compared (e.g., in pairs), a high criterion for claiming abstract-concept learning (e.g., transfer performance equivalent to baseline performance), and systematic manipulation of the training set (e.g., increases in the number of rule exemplars when transfer is less than baseline performance). The research covered in this article on the recent advancements in abstract-concept learning show this basic ability in higher-order cognitive processing is common to many animal species and that "uniqueness" may be limited more to how quickly new abstract concepts are learned rather than to the ability itself.

Mechanisms of same/different concept learning by rhesus monkeys, capuchin monkeys, and pigeons were studied in terms of how these species learned the task (e.g., item-specific learning versus relational learning) and how rapidly they learned the abstract concept, as the training set size was doubled. They had similar displays, training stimuli, test stimuli, and contingencies. The monkey species learned the abstract concept at similar rates and more rapidly than pigeons, thus showing a quantitative difference across species. All species eventually showed full concept learning (novel-stimulus transfer equivalent to baseline: 128-item set size for monkeys; 256-item set for pigeons), thus showing a qualitative similarity across species. Issues of stimulus regularity/symmetry, generalization from item pairs, and familiarity processing were not considered to be major factors in the final performances, converging on the conclusion that these species were increasingly controlled by the sample-test relationship (i.e., relational processing) leading to full abstract-concept learning.

Eight pigeons were trained and tested in a simultaneous same/different task. After pecking an upper picture, they pecked a lower picture to indicate same or a white rectangle to indicate different. Increases in the training set size from 8 to 1,024 items produced improved transfer from 51.3% to 84.6%. This is the first evidence that pigeons can perform a two-item same/different task as accurately with novel items as training items and both above 80% correct. Fixed-set control groups ruled out training time or transfer testing as producing the high level of abstract-concept learning. Comparisons with similar experiments with rhesus and capuchin monkeys showed that the ability to learn the same/different abstract concept was similar but that pigeons require more training exemplars., (((c) 2006 APA, all rights reserved).)

Eight pigeons learned either matching (to sample) or oddity (from sample) with or without reward for sample responding. The training stimuli were coarse-white, fine-black, or smooth-mauve gravels in pots with buried grain as the reinforcer. Oddity without sample reward was learned most rapidly, followed by matching with sample reward, oddity with sample reward, and matching without sample reward. Transfer was related to acquisition rate: The oddity group without sample reward showed full (equal to baseline) color and texture transfer; the matching group with sample reward showed partial texture transfer; other groups showed no transfer. Sample reward was shown to determine rate of acquisition of matching and oddity and the oddity preference effect. The results are discussed in terms of item-specific associations operating early in learning prior to any relational learning between sample and comparison stimuli.

A rhesus monkey's memory was tested for single items and four-item lists of natural and environmental sounds. Memory items were presented from a center speaker, followed by a retention delay and then a choice response to a test sound presented simultaneously from two side speakers. Recognition of the last item of four-item lists was much poorer than that of single items at 0-, 1-, and 2-sec delays, despite there being the same temporal relations between study and test. This result showed that the first three items proactively interfered with memory of the last list item. Proactive interference dissipated after 2 sec, revealing a recency effect that eventually equaled single-item performance. Recognition of the first item of four-item lists was much poorer than single items at 20- and 30-sec delays, showing that the last three items retroactively interfered with memory of the first list item. The results point to the critical nature of interference processes in the understanding of serial position functions.

Three capuchin monkeys (Cebus apella) touched the lower of 2 pictures (same) or a white rectangle (different), increased same/different abstract-concept learning (52% to 87%) with set-size increases (8 to 128 pictures), and were better than 3 rhesus monkeys (Macaca mulatta). Three other rhesus that touched the top picture before choices learned similar to capuchins but were better at list-memory learning. Both species' serial position functions were similar in shape and changes with retention delays. Other species showed qualitatively similar shape changes but quantitatively different time-course changes. In abstract-concept learning, qualitative similarity was shown by complete concept learning, whereas a quantitative difference would have been a set-size slope difference. Qualitative similarity is discussed in relation to general-process versus modular cognitive accounts.

Experiments with 9 rhesus monkeys (Macaca mulatta) showed, for the first time, that abstract-concept learning varied with the training stimulus set size. In a same/different task, monkeys required to touch a top picture before choosing a bottom picture (same) or white rectangle (different) learned rapidly. Monkeys not required to touch the top picture or presented with the top picture for a fixed time learned slowly or not at all. No abstract-concept learning occurred after 8-item training but progressively improved with larger set sizes and was complete following 128-item training. A control monkey with a constant 8-item set ruled out repeated training and testing. Contrary to the unique-species account, it is argued that different species have quantitative, not qualitative, differences in abstract-concept learning.

A rhesus monkey was tested in an auditory list memory task with blocked and mixed retention delays. Each list of four natural or environmental sounds (from a center speaker) was followed by a retention delay (0, 1, 2, 10, 20, or 30 sec) and then by a recognition test (from two side speakers). The monkey had been tested for 12 years in tasks with blocked delays. An earlier (4 years prior) blocked-delay test was repeated, with virtually identical results. The results from the mixed-delay test were likewise similar. Thus, the peculiarities of blocked-delay testing, such as delay predictability or differences in list spacing, apparently do not alter this monkey's memory for auditory lists. It is concluded from this and other evidence that the monkey's serial position functions reflect mnemonic processes that change with changes in retention delay and are not artifacts of the blocked-delay procedure. The nature of the monkey's auditory memory is discussed.

Recognition of words and kaleidoscope pictures showed a double dissociation of left and right hippocampal activity using magnetic source imaging (MSI). MSI has advantages over alternative imaging techniques that measure hemodynamic changes for identifying regional changes in brain activity in real time and on an individual subject basis without the need for image subtraction. In this study, lists of words or kaleidoscope pictures were presented for memorization followed by tests of list items and foils during which brain activity was recorded. There was greater activation in the left than the right hippocampus with abstract nouns (e.g., relief) and greater activation in the right than the left hippocampus with kaleidoscope pictures. This dissociation was evident on a case by case basis. This study demonstrates the specialization of the two medial temporal lobe (MTL) regions, including the hippocampi, for mnemonic processing of verbal and pictorial items that are difficult to encode verbally.