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Behavioral flexibility, the ability to adapt behavior to new circumstances, is thought to play an important role in a species’ ability to successfully adapt to new environments and expand its geographic range. However, flexibility is rarely directly tested in a way that would allow us to determine how flexibility works to predict a species’ ability to adapt their behavior to new environments. We use great-tailed grackles (Quiscalus mexicanus; a bird species) as a model to investigate this question because they have recently rapidly expanded their range into North America. We attempted to manipulate grackle flexibility using shaded (light and dark gray) tube reversal learning to determine whether flexibility is generalizable across contexts (multi-access box), and what learning strategies grackles employ. We found that flexibility was manipulable: birds in the manipulated group took fewer trials to pass criterion with increasing reversal number, and they reversed a shade preference in fewer trials by the end of their serial reversals compared to control birds who had only one reversal. Birds that passed their last reversal faster were also more flexible (faster to switch between loci) and innovative (solved more loci) on a multi-access box. All grackles in the manipulated reversal learning group used one learning strategy (epsilon-decreasing) in all reversals, and none used a particular exploration or exploitation strategy earlier or later in their serial reversals. Understanding how flexibility causally relates to other traits will allow researchers to develop robust theory about what flexibility is and when to invoke it as a primary driver in a given context, such as a rapid geographic range expansion.

Behavioral flexibility, the ability to change behavior when circumstances change based on learning from previous experience, is thought to play an important role in a species ability to successfully adapt to new environments and expand its geographic range. It is alternatively or additionally possible that causal cognition, the ability to understand relationships beyond their statistical covariations, could play a significant role in rapid range expansions via the ability to learn faster: causal cognition could lead to making better predictions about outcomes through exerting more control over events. We aim to determine whether great-tailed grackles (Quiscalus mexicanus), a species that is rapidly expanding its geographic range, use causal inference and whether this ability relates to their behavioral flexibility (flexibility measured in these individuals by Logan et al. (2019): reversal learning of a color discrimination and solution switching on a puzzle box). Causal cognition was measured using a touchscreen where individuals learned about the relationships between a star, a tone, a clicking noise, and food. They were then tested on their expectations about which of these causes the food to become available. We found that eight grackles showed no evidence of making causal inferences when given the opportunity to intervene on observed events using a touchscreen apparatus, and that performance on the causal cognition task did not correlate with behavioral flexibility measures. This could indicate that our test was inadequate to assess causal cognition. Because of this, we are unable to speculate about the potential role of causal cognition in a species that is rapidly expanding its geographic range. We suggest further exploration of this hypothesis using larger sample sizes and multiple test paradigms.

The Partial Reinforcement Extinction Effect (PREE) is the finding of more rapid extinction to a continuously reinforced cue than to a partially reinforced cue. Despite its well-documented existence, it has challenged many theoretical accounts of learning that assume the likelihood of responding is correlated with the strength of the learned Stimulus (S) – Response (R) association. While a number of explanations (notably by Amsel and Capaldi) of the PREE have been proposed, Rescorla (1999) reported a PREE using an entirely within-subjects design—a finding that runs counter to these proposed explanations. We describe how the PREE might be explained by the formation of inhibitory S-R associations that develop during the acquisition of partial reinforcement, but not continuous reinforcement. To test this proposal, we trained pigeons to peck a response key in the presence of one partially reinforced stimulus but also omitted the response key during the training of an alternative partially reinforced stimulus—thus preventing the development of inhibitory S-Rs to that latter stimulus. We found evidence for the PREE only for the partially reinforced stimulus where inhibitory S-Rs were capable of being formed, thus supporting our proposal. Alternative explanations of our procedure and its relevance to broader computational rules of learning is discussed.

Subjects are often willing to pay a cost for information. In a procedure that promotes paradoxical choices, animals choose between a richer option followed by a cue that is rewarded 50% of the time (No Info) vs. a leaner option followed by one of two cues that signal certain outcomes: one always rewarded (100%) and the other never rewarded, 0% (Info). Since decisions involve comparing the subjective value of options after integrating all their features, preference for information may rely on cortico-amygdalar circuitry. To test this, male and female rats were prepared with bilateral inhibitory Designer Receptors Exclusively Activated by Designer Drugs (DREADDs) in the anterior cingulate cortex, orbitofrontal cortex, basolateral amygdala, or null virus (control). We inhibited these regions after stable preference was acquired. We found that inhibition of the anterior cingulate cortex destabilized choice preference in female rats without affecting latency to choose or response rate to cues. A logistic regression fit revealed that previous choice predicted current choice in all conditions, however previously rewarded Info trials strongly predicted preference in all conditions except in female rats following anterior cingulate cortex inhibition. The results reveal a causal, sex-dependent role for the anterior cingulate cortex in decisions involving information.

When information about an event is perceptually occluded, individuals might recognize that the event might be present but that they cannot detect it because of the occluder. In such situations, an individual should continue to believe that the prevailing contingencies have not changed. This is in stark contrast to conditions where an expected event is explicitly absent, which should lead the individual to update their contingency knowledge. In an autoshaping procedure, we tested whether pigeons can discriminate conditions of perceptual ambiguity from perceptual certainty. Pigeons first learned to peck at two Pavlovian visual cues, followed by extinction of one of the cues. During extinction, the feeder was occluded by a metal shield for pigeons in Group Occluded, while the metal shield was placed next to but not covering the feeder for pigeons in Group Un-Occluded. On a final test with the metal shield removed, pigeons in Group Un-occluded pecked less at the extinguished cue than at the un-extinguished cue; while pigeons in Group Occluded pecked at an equally high rate to both cues. These results replicate in pigeons similar results reported in rats by Waldmann et al. (2012), and show that pigeons are able to discriminate conditions of certainty from conditions of ambiguity.

Research into animal cognitive abilities is increasing quickly and often uses methods where behavioral performance on a task is assumed to represent variation in the underlying cognitive trait. However, because these methods rely on behavioral responses as a proxy for cognitive ability, it is important to validate that the task structure does, in fact, target the cognitive trait of interest rather than non-target cognitive, personality, or motivational traits (construct validity). Although it can be difficult, or impossible, to definitively assign performance to one cognitive trait, one way to validate that task structure is more likely to elicit performance based on the target cognitive trait is to assess the temporal and contextual repeatability of performance. In other words, individual performance is likely to represent an inherent trait when it is consistent across time and across similar or different tasks that theoretically test the same trait. Here, we assessed the temporal and contextual repeatability of performance on tasks intended to test the cognitive trait behavioral flexibility in great-tailed grackles (Quiscalus mexicanus). For temporal repeatability, we quantified the number of trials to form a color preference after each of multiple color reversals on a serial reversal learning task. For contextual repeatability, we then compared performance on the serial color reversal task to the latency to switch among solutions on each of two different multi-access boxes. We found that the number of trials to form a preference in reversal learning was repeatable across serial color reversals and the latency to switch a preference was repeatable across color reversal learning and the multi-access box contexts. This supports the idea that the reversal learning task structure elicits performance reflective of an inherent trait, and that reversal learning and solution switching on multi-access boxes similarly reflect the inherent trait of behavioral flexibility.

We studied object-location binding in pigeons using a sequence learning procedure. A sequence of four objects was presented, one at a time at one of four locations on a touchscreen. A single peck at the object ended the trial, and food reinforcement was delivered intermittently. In Experiment 1, a between-subjects design was used to present objects, locations, or both in a regular sequence or randomly. Response time costs on nonreinforced probe tests on which object order, location order, or both were disrupted revealed sequence learning effects. Pigeons encoded location order when it was consistent, but not object order when it alone was consistent. When both were consistent, pigeons encoded both, and showed evidence of object-location binding. In Experiment 2, two groups of pigeons received training on sequences where the same object always appeared at the same location. For some pigeons a consistent sequence was used while for others sequence order was randomized. Only when sequence order was consistent was object-location binding found. These experiments are the first demonstrations of strong and lasting feature binding in pigeons and are consistent with a functional account of learning.

To examine the cognitive and physical changes associated with consuming an energy drink concurrent to video gaming, we examined a convenience sample of nine elite League of Legends (LoL) e-sport players (21 ± 2 y, BMI 25.6 ± 3.4 kg/m2) consuming an energy drink (ReloadTM) or placebo (Placebo) in a randomized, double-blind, placebo-controlled cross-over trial. Participants completed the same test battery prior to treatment consumption and after playing each of three competitive LoL games. Primary outcomes included measures of attention (Erikson Flanker Test), reaction time (Go/No-Go test) and working memory (n-back test). Secondary outcomes examined fatigue (hand grip strength and finger tap speed). Statistical analysis was performed by repeated-measures analysis of variance (RM-ANOVA) and reported as the mean (standard deviation [SD]) or mean change (95% confidence interval [CI]). Participants reported sleeping 8.1 (1.2) h/night, playing LoL 10.3 (2.1) h/d, playing other video games 1.8 (2.8) h/d, and exercising 4.2 (1.7) times per week. Overall, we observed no significant time, group, or group-by-time interactions for any measured performance index with the exception of a significant improvement for the n-back test, where the Reload group demonstrated a significant within-group improvement: Reload [-171 ms (95% CI, -327.91, -14.09), p < 0.004], Placebo [-92 ms (95% CI, -213.63, 29.63)]. However, no between-group differences were noted (38.50 ms, 95% CI, -141.89, 64.89, p = 0.803). Our findings suggest that elite eSport athletes do not demonstrate a mental or physical improvement in performance relative to the treatment supplement or indices measured in this study.

Deeply rooted within the history of experimental psychology is the search for general laws of learning that hold across tasks and species. Central to this enterprise has been the notion of equipotentiality; that any two events have the same likelihood of being associated with one another as any other pair of events. Much work, generally summarized as ‘biological constraints on learning,’ has challenged this view, and demonstrates pre-existing relations between cues and outcomes, based on genes and prior experience, that influence potential associability. Learning theorists and comparative psychologists have thus recognized the need to consider how the evolutionary history as well as prior experience of the organism being studied influences its ability to learn about and navigate its environment. We suggest that current models of human memory, and human memory research in general, lack sufficient consideration of how human evolution has shaped human memory systems. We review several findings that suggest the human memory system preferentially processes information relevant to biological fitness, and highlight potential theoretical and applied benefits afforded by adopting this functionalist perspective.

We've shown that pigeons can integrate separately acquired spatial maps into a cognitive map. Integration requires an element shared between maps. In two experiments using a spatial-search task in pigeons, we test spatial combination rules when no shared element was present during training. In all three experiments, pigeons first learned individual landmark-target maps. In subsequent tests involving combinations of landmarks, we found evidence that landmarks collaborate in guiding spatial choice at test (Experiment 1). In Experiment 2, pigeons were trained on two landmarks with different proximities to the target. On tests on a compound of both landmarks, pigeons showed stronger spatial control by the more proximal landmark, a performance overshadowing effect. Extinction of the proximal landmark shifted spatial control to the non-extinguished distal landmark. This reveals that the performance overshadowing effect was associative in nature, and not due to perceptual or spatial biases. This emphasis on spatial control during performance reflects the emphasis on performance processes that were a major focus in Ralph Miller's lab.

Age-related decrements in cognitive ability have been proposed to stem from deteriorating function of the hippocampus. Many birds are long lived, especially for their relatively small body mass and elevated metabolism, making them a unique model of resilience to aging. Nevertheless, little is known about avian age-related changes in cognition and hippocampal physiology. We studied spatial cognition and hippocampal expression of the age-related gene, Apolipoprotein D (ApoD), and the immediate early gene Egr-1 in zebra finches at various developmental time points. In a first experiment, middle-aged adult males outperformed middle-aged females in learning correct food locations in a four-arm maze, but all birds remembered the task equally well after a 5- or 10-day delay. In a second experiment comparing young and old birds, aged birds showed minimal evidence for deterioration in spatial cognition or motivation relative to young birds, except that aged females showed less rapid gains in accuracy during spatial learning than young females. These findings indicate that sex differences in hippocampus-dependent spatial learning and decline with age are phylogenetically conserved. With respect to hippocampal gene expression, adult females expressed Egr-1 at significantly greater levels than males after memory retrieval, perhaps reflecting a neurobiological compensation. Contrary to mammals, ApoD expression was elevated in young zebra finches compared with aged birds. This may explain the near absence of decrements in spatial memory due to age, possibly indicating an alternative mechanism of managing oxidative stress in aged birds. (PsycINFO Database Record

Organisms are believed to attempt to maximize their net energy intake while foraging. The paradoxical choice task shows that they may instead prefer to obtain information rather than primary reward when the outcome is uncertain. That is, they prefer stimuli that consistently predict food or no food (informative option), to stimuli that inconsistently predict both food and no food in larger amounts (noninformative option). This task also seems to indicate that some species (like pigeons, Columba livia, and starlings, Sturnus vulgaris) are more prone to choose the informative option, while other species (like rats, Rattus norvegicus, and humans, Homo sapiens) tend to favor reward procurement through the noninformative option. There is empirical evidence for and against this view. However, an analysis of the literature suggests that species differences in paradoxical choice might be less pronounced than often believed. We argue that pigeons and rats are usually not tested under conditions that are motivationally equivalent for both species—in particular, the opportunities to track consistent stimulus–food pairings are less often met in the rat studies than in the pigeon studies. [ABSTRACT FROM AUTHOR]

Prior work in our lab has shown that an expanding image on a computer screen elicits a hiding response in the Caribbean terrestrial hermit crab (Coenobita clypeatus). We conducted two experiments to identify what properties of the expanding stimulus contribute to its effectiveness as a visual threat. First we found that an expanding geometric star evoked a strong hiding response while a contracting or full-sized stationary star did not. A second experiment revealed that the more quickly the stimulus expanded the shorter the latency to hide. These findings suggest that the anti-predator response to looming stimulus relies heavily on visual cues relating to the manner of approach. The simulated visual threat on a computer screen captures key features of a real looming object that elicits hiding behavior in crabs in the wild.

Identifying statistical patterns between environmental stimuli enables organisms to respond adaptively when cues are later observed. However, stimuli are often obscured from detection, necessitating behavior under conditions of ambiguity. Considerable evidence indicates decisions under ambiguity rely on inference processes that draw on past experiences to generate predictions under novel conditions. Despite the high demand for this process and the observation that it deteriorates disproportionately with age, the underlying mechanisms remain unknown. We developed a rodent model of decision-making during ambiguity to examine features of experience that contribute to inference. Rats learned either a simple (positive patterning) or complex (negative patterning) instrumental discrimination between the illumination of one or two lights. During test, only one light was lit while the other relevant light was blocked from physical detection (covered by an opaque shield, rendering its status ambiguous). We found experience with the complex negative patterning discrimination was necessary for rats to behave sensitively to the ambiguous test situation. These rats behaved as if they inferred the presence of the hidden light, responding differently than when the light was explicitly absent (uncovered and unlit). Differential expression profiles of the immediate early gene cFos indicated hippocampal involvement in the inference process while localized microinfusions of the muscarinic antagonist, scopolamine, into the dorsal hippocampus caused rats to behave as if only one light was present. That is, blocking cholinergic modulation prevented the rat from inferring the presence of the hidden light. Collectively, these results suggest cholinergic modulation mediates recruitment of hippocampal processes related to past experiences and transfer of these processes to make decisions during ambiguous situations. Our results correspond with correlations observed between human brain function and inference abilities, suggesting our experiments may inform interventions to alleviate or prevent cognitive dysfunction. © 2015 Wiley Periodicals, Inc.

We investigated theoretical accounts of spatial overshadowing using a landmark-based spatial-search task in a touchscreen preparation with pigeons. Pigeons first learned to find a hidden target on a screen using a compound of two visual cues as landmarks. Landmark A was proximal to the target while landmark X was distal to the target. Experiment 1 replicated our prior spatial overshadowing effect whereby landmark A overshadowed the development of spatial control by X. Spatial control by X was also poorer than by landmark Y which had been paired with the target alone but with the same absolute distance to the target as X had. Thus, the poor spatial control by X was not merely due to the greater X-target distance (relative to the A-target distance). Experiments 2a and 2b failed to find recovery from spatial overshadowing of X through either post-training extinction or counterconditioning of overshadowing landmark A, respectively. We interpret our results as being consistent with acquisition-focused models of elementary associative learning, but not with performance-focused models.

Hermit crabs hide into shells when confronted with potential dangers, including images presented on a monitor. We do not know, however, what hermit crabs can see and how they perceive different objects. We examined the hiding response of the Caribbean hermit crab (Coenobita clypeatus) to various stimuli presented on a monitor in seven experiments to explore whether crabs could discriminate different properties of a threatening digital image, including color, brightness, contrast, shape and orientation. We found crabs responded differently to expanding circles presented in wavelengths of light corresponding to what humans see as red, blue, and green. "Blue" stimuli elicited the strongest hiding response (Experiments 1, 2, & 7). "Blue" was also more effective than a gray stimulus of similar brightness (Experiment 3). Hermit crabs were sensitive to the amount of contrast between a stimulus and its background rather than absolute brightness of the stimulus (Experiment 4). Moreover, we did not find evidence that crabs could discriminate orientation (Experiment 6), and mixed evidence that they could discriminate stimulus shape (Experiments 5 & 7). These results suggest that the Caribbean hermit crab is sensitive to color features, but not spatial features, of a threatening object presented on a computer screen. This is the first study to use the hiding response of the hermit crab to examine its visual ability, and demonstrates that the hiding response provides a useful behavioral approach with which to study learning and discrimination in the hermit crab.

Many studies investigating cue competition have focused on the blocking effect. We investigated the blocking effect with pigeons using a landmark-based spatial search task in both a touchscreen preparation (Exp. 1a) and an automated remote environmental navigation apparatus (Exp. 1b). In Phase 1, two landmarks (LMs: A and Z) appeared on separate trials as colored circles among a row of eight (touchscreen) or six (ARENA) identical response units. Subjects were rewarded for pecking at a target response unit to the right of LM A and to the left of an extraneous LM, Z. During the blocking trials in Phase 2, LM X was presented in compound with a second LM (A) that had been previously trained. On control trials, LM Y was presented in compound with LM B and a target in the same manner as in the trials of AX, except that neither landmark had previously been trained with the target. All subjects were then tested with separate trials of A, X, B, and Y. Testing revealed poor spatial control by X relative to A and Y. We report the first evidence for a spatial-blocking effect in pigeons and additional support for associative effects (e.g., blocking) occurring under similar conditions (e.g., training sessions, spatial relationships, etc.) in 3-D and 2-D search tasks.

IntroductionPurified high-fat diet (HFD) feeding causes deleterious metabolic and cognitive effects when compared with unrefined low-fat diets in rodent models. These effects are often attributed to the diet's high content of fat, while less attention has been paid to other mechanisms associated with the diet's highly refined state. Although the effects of HFD feeding on cognition have been explored, little is known about the impact of refined vs. unrefined food on cognition. We tested the hypothesis that a refined low-fat diet (LFD) increases body weight and adversely affects cognition relative to an unrefined diet.Materials and methodsRats were allowed ad libitum access to unrefined rodent chow (CON, Lab Diets 5001) or a purified low-fat diet (REF, Research Diets D12450B) for 6 months, and body weight and performance on an instrumental lever pressing task were recorded.ResultsAfter six months on their respective diets, group REF gained significantly more weight than group CON. REF rats made significantly fewer lever presses and exhibited dramatically lower breaking points than CON rats for sucrose and water reinforcement, indicating a chronic reduction of motivation for instrumental performance. Switching the rats' diet for 9 days had no effect on these measures.ConclusionsDiet-induced obesity produces a substantial deficit in motivated behavior in rats, independent of dietary fat content. This holds implications for an association between obesity and motivation. Specifically, behavioral traits comorbid with obesity, such as depression and fatigue, may be effects of obesity rather than contributing causes. To the degree that refined foods contribute to obesity, as demonstrated in our study, they may play a significant contributing role to other behavioral and cognitive disorders.

Prior research, using two- and three-dimensional environments, has found that when both human and nonhuman animals independently acquire two associations between landmarks with a common landmark (e.g., LM1-LM2 and LM2-LM3), each with its own spatial relationship, they behave as if the two unique LMs have a known spatial relationship despite their never having been paired. Seemingly, they have integrated the two associations to create a third association with its own spatial relationship (LM1-LM3). Using sensory preconditioning (Experiment 1) and second-order conditioning (Experiment 2) procedures, we found that human participants integrated information about the boundaries of pathways to locate a goal within a three-dimensional virtual environment in the absence of any relevant landmarks. Spatial integration depended on the participant experiencing a common boundary feature with which to link the pathways. These results suggest that the principles of associative learning also apply to the boundaries of an environment.

The human body—an amazing biological system that scales up fractally from its cellular building blocks—exhibits an incredible ability to self heal. Why then, are chronic diseases and degeneration on the rise in the population? Why are we sicker, more obese, and more depressed and stressed than ever before in human history? Why can’t we heal? The answers to these questions may lie in our ancestry, and modern departure from the human ecological niche. The ability to heal requires proper spatio-temporal inputs—nutrition, sleep, stress, activity, and socialization—in order for cellular signaling to occur properly across semi-permeable cell membranes. We first review key steps in the evolutionary history of multicellular life, focusing on the fundamental role of cell-cell interactions. Next, we present this as an important framework by which to understand how the entrainment of physiological signals in homeostatic mechanisms reveals new insights into the processes of disease. Examples are drawn from the evolution of metabolism, nutrition, and respiration in multicellular life. We argue that disease processes result from a mismatch between the physiological inputs an individual receives and their optimal amount and fractal distribution as determined by an individual’s ancestry. A comparative analysis is a useful tool by which to illuminate deep homologies that reveal a mechanistic account for disease processes. This cell-molecular approach provides a useful contrast to the traditional reductionist approach to disease exemplified by the human genome project. As an example, we describe how cell-cell communication drives the ontogeny and phylogeny of physiology, producing the tissues, organs, and organ systems that hierarchically serve human physiology on various levels. Modern society, with its disconnected and stress-riddled lifestyle, is increasingly failing to provide the proper inputs for healthy gene expression and physiological function. Thus, the answers to our modern health woes—physical, mental, and social—may lie in acknowledging the powerful roles that our past has played in shaping our bodies. Finding ways to provide the proper inputs of the human ecological niche in the modern day may lead to significant, perhaps staggering improvements in our health and wellness. The fractal mathematics underpinning these dynamics also serves as a metaphor for the Ancestral Health Movement, which is currently arising as a multi-cultural, multi-national grass-roots pluralistic phenomenon.

Children love to play. Why do they find such a frivolous activity so pleasurable and desirable? Perhaps it is not frivolous, but instead is an adaptation designed to guide proper cognitive development in human children. To understand why, I marshal evidence from different fields to build a case for play as a central behavioral mechanism of human brain and cognitive development. I start with a discussion of human evolution, focusing on the evolution of human physiology, tool-use, the human brain, and life-history strategy, and development, and how these are all connected as an adaptive suite. The anthropological and developmental evidence suggests the existence of an extended childhood adapted to establish the skills, knowledge, and understanding necessary to become a successful hunter-gatherer. I also compare human and chimpanzee brain development, and how brain-specific genes evolved uniquely in humans to foster human brain development. I conclude with the evidence from developmental psychology that even contemporary, first-world children are born with the drive to learn and develop intellectually through play. In this framework, human play can be viewed as an adaptation that guides human brain development to produce curious, intelligent and well-adjusted adults. I close by speculating on the possibility that barriers to or constraints on play may hamper intellectual and cognitive development. I focus on the important concept of developmental decanlization as a mechanism of evolutionary mismatch. I argue that more empirical study is needed to better understand the importance of play compared to other forms of education for optimal intellectual and cognitive development.

Welcome to the first issue of the Journal of Evolution and Health! The Journal of Evolution and Health is the peer-reviewed, open-access journal of the Ancestral Health Society, a community of scientists, healthcare professionals, and laypersons who collaborate to understand health challenges from an evolutionary perspective.

The goal of three experiments was to study whether rats are aware of the difference between absence of events and lack of evidence. We used a Pavlovian extinction paradigm in which lights consistently signaling sucrose were suddenly paired with the absence of sucrose. The crucial manipulation involved the absent outcomes in the extinction phase. Whereas in the Cover conditions, access to the drinking receptacle was blocked by a metal plate, in the No Cover conditions, the drinking receptacle was accessible. The Test phase showed that in the Cover conditions, the measured expectancies of sucrose were clearly at a higher level than in the No Cover conditions. We compare two competing theories potentially explaining the findings. A cognitive theory interprets the observed effect as evidence that the rats were able to understand that the cover blocked informational access to the outcome information, and therefore the changed learning input did not necessarily signify a change of the underlying contingency in the world. An alternative associationist account, renewal theory, might instead explain the relative sparing of extinction in the Cover condition as a consequence of context change. We discuss the merits of both theories as accounts of our data and conclude that the cognitive explanation is in this case preferred.

Six pigeons were trained using a matching-to-sample procedure where sample and rewarded comparisons matched on both attributional (color) and relational (horizontal or vertical orientation) dimensions. Probes then evaluated the pigeons' preference to comparisons that varied in these dimensions. A strong preference was found for the attribute of color. The discrimination was not found to transfer to novel colors, however, suggesting that a general color rule had not been learned. Further, when color could not be used to guide responding, some influence of other attributional cues such as shape, but not relational cues, was found. We conclude that pigeons based their performance on attributional properties of but not on relational properties between elements in our matching-to-sample procedure. Future studies should look at examining other attributes to compare attributional versus relational processing.

The study of human and animal behavior in psychology is almost always framed at either the associative or the cognitive level of explanation. Despite continued debate between proponents of each approach, we appear to be no closer to a consensus view than we were when the debate began in earnest in the 1960 s. Could it be that the two levels of explanation are irreconcilable? Or is it possible that both frameworks are useful, though incompatible? Perhaps these frameworks merely account for the same behaviors but at different levels of explanation, as characterized by hardware-software or genotype-phenotype analogies. This special issue provides a venue for contemporary scientists involved in this debate to express their views, and follows from a Focus Session of the same title held at the 2010 meeting of the Winter Conference in Animal Learning & Behavior.

The connection between reduced reinforcement probability and increased behavioral variability hasbeen well established in recent years (Gharib, Gade, & Roberts, 2004; Stahlman, Roberts, & Blaisdell, 2010). Researchers have hypothesized that this relationship is an adaptive one - it is beneficial for animals to increase behavioral variability in response to low likelihood of success,because this increase in variability potentially allows them to discover new behavioral options that are more highly rewarded. We conducted a study to investigate the relationship between behavioral variability and reward probability in an ecologically valid experimental task. We trained rats tosearch for hidden food in the presence of either of two landmarks, each that signaled a different likelihood of reward (i.e., HI and LO). Variation in locations searched was higher in the presence ofthe LO probability landmark. These results build on prior findings that reward expectation drives behavioral variability.

Artificial sensations can be produced by direct brain stimulation of sensory areas through implanted microelectrodes, but the perceptual psychophysics of such artificial sensations are not well understood. Based on prior work in cortical stimulation, we hypothesized that perceived intensity of electrical stimulation may be explained by the population response of the neurons affected by the stimulus train. To explore this hypothesis, we modeled perceived intensity of a stimulation pulse train with a leaky neural integrator. We then conducted a series of two-alternative forced choice behavioral experiments in which we systematically tested the ability of rats to discriminate frequency, amplitude, and duration of electrical pulse trains delivered to the whisker barrel somatosensory cortex. We found that the model was able to predict the performance of the animals, supporting the notion that perceived intensity can be largely accounted for by spatiotemporal integration of the action potentials evoked by the stimulus train.

In two experiments, rats received sensory preconditioning treatment in which an auditory conditioned stimulus (CS) X was followed by visual CS A in Phase 1, and CS A was followed by an appetitive US (sucrose) in Phase 2. Rats also received presentations of auditory CS Y unpaired with other events. At test, rats looked for sucrose more following CS X than following CS Y on non-reinforced probe test trials only if the light bulb on which CS A had been presented during training was removed from the chamber at the time of testing. With the light bulb present (but unlit), rats showed nodifference in amount of nose poking between CS X and CS Y. These results suggest that rats distinguish between the explicit absence of events and lack of information

Grau and Joynes (2005) assess the current state of the field of animal learning and behavior, with particular emphasis on pedagogical and curricular issues. They suggest that the conventional framework which organizes lecture material around methodology is flawed and that an organization around mechanism should be used instead. They also advocate a shift from a purely behavioral approach to research on learning and behavior to a neural-functionalist approach more akin to contemporary behavioral neuroscience. While I support many of the suggestions for improving instruction, I disagree with their proposed shift away from purely behavioral investigations of animal behavior. Behavioral research continues to be a thriving and productive source of empirical and theoretical discoveries. The diverse array of specialized methodologies that have been developed to pursue this work are still paying dividends by illuminating the nature of behavioral mechanisms. Banishing purely behavioral approaches to learning and behavior, such as those used to study associative learning, animal cognition, and comparative psychology, would severely hamper our knowledge of behavioral mechanism.

We report the first successful demonstration of a simultaneous, two-item same-different (S/D) discrimination by 6 pigeons, in which nonpictorial color and shape stimuli were used. This study was conducted because the majority of recently successful demonstrations of S/D discrimination in pigeons have employed displays with more than two items. Two pairs of stimulus items were simultaneously presented on a touch screen equipped computer monitor. Pigeons were reinforced for consistently pecking at either the same (i.e., identical) or the different (i.e., nonidentical) pair of items. These pairs were created from combinations of simple colored shapes drawn from a pool of six colors and six shapes. After acquiring the discrimination with item pairs that differed redundantly in both the shape and the color dimensions, the pigeons were tested for transfer to items that varied in only one of these dimensions. Although both dimensions contributed to the discrimination, greater control was exhibited by the color dimension. Most important, the discrimination transferred in tests with novel colored, shaped, and sized items, suggesting that the mechanisms involved were not stimulus specific but were more generalized in nature. These results suggest that the capacity to judge S/D relations is present in pigeons even when only two stimuli are used to implement this contrast.

The integration of spatial maps in pigeons was investigated using a spatial analog to sensory preconditioning. The pigeons were tested in an open-field arena in which they had to locate hidden food among a 4x4 grid of gravel-filled cups. In phase 1, the pigeons were exposed to a consistent spatial relationship (vector) between landmark L (a red L-shaped block of wood), landmark T (a blue T-shaped block of wood) and the hidden food goal. In phase 2, the pigeons were then exposed to landmark T with a different spatial vector to the hidden food goal. Following phase 2, pigeons were tested with trials on which they were presented with only landmark L to examine the potential integration of the phase 1 and 2 vectors via their shared common elements. When these test trials were preceded by phase I and phase 2 reminder trials, pigeons searched for the goal most often at a location consistent with their integration of the L --> T phase 1 and T --> phase 2 goal vectors. This result indicates that integration of spatial vectors acquired during phases I and 2 allowed the pigeons to compute a novel L --> goal vector. This suggests that spatial maps may be enlarged by successively integrating additional spatial information through the linkage of common elements.