The behavioral origins of phylogenic responses and ontogenic habits.

An examination of innate behavior and its possible origins suggests parallels with the formation of habitual behavior. Inflexible but adaptive responses-innate reflexive behavior, Pavlovian conditioned responses, and operant habits-may have evolved from variable behavior in phylogeny and ontogeny. This form of "plasticity-first" scientific narrative was unpopular post-Darwin but has recently gained credibility in evolutionary biology. The present article seeks to identify originating events and contingencies contributing to such inflexible but adaptive behavior at both phylogenic and ontogenic levels of selection. In ontogeny, the development of inflexible performance (i.e., habit) from variable operant behavior is reminiscent of the genetic accommodation of initially variable phylogenic traits. The effects characteristic of habit (e.g., unresponsiveness to reinforcer devaluation) are explicable as the result of a conflict between behaviors at distinct levels of selection. The present interpretation validates the practice of seeking hard analogies between evolutionary biology and operant behavior. Finding such parallels implies the validity of a claim that organismal behavior, both innate and learned, is a product of selection by consequences. A complete and coherent account of organismal behavior may ultimately focus on functional selective histories in much the same way evolutionary biology does with its subject matter.

The effects of feature extinction in dual-response feature-positive discriminations.

In a typical feature-positive discrimination, responding is reinforced (+) during the target stimulus (A) on trials with the feature stimulus (X), but not during target-alone trials (A-). When X and A are presented simultaneously, direct control by X is typically observed; however, when the stimuli are presented serially, X sets the occasion for responding to A. In the current dual-response procedures, one response (e.g., left lever press) was reinforced during feature-target trials (XA+) and a different response (e.g., right lever press) was reinforced during target-alone trials (A+). In Experiment 1, rats received either serial (X → A+) or simultaneous (X:A+) presentations of the feature-target compound along with target-alone trials (A+). Contrary to our predictions, the serial group failed to learn the discrimination and the simultaneous group demonstrated occasion setting. In Experiment 2, the salience of the feature was increased, which resulted in direct control by the feature in both groups. In Experiment 3, an additional serial group was included with a longer interval between the feature (X) and target (A). Despite the reduced temporal proximity of X to reinforcement, direct control was again observed in all groups. The current pattern of results in the simultaneous and serial groups is interpreted in relation to the enhanced salience of A relative to X, due to separate pairings of A-alone with reinforcement in the dual-response procedure. Consistent with previous findings, occasion setting was observed when A was most salient relative to X (Experiment 1, simultaneous group), but direct control was found when the salience of X was increased (Experiments 2-3). (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Devaluation of a conditioned reinforcer requires its reexposure.

A change in motivational state does not guarantee a change in operant behaviour. Only after an organism has had contact with an outcome while in a relevant motivational state does behaviour change, a phenomenon called incentive learning. While ample evidence indicates that this is true for primary reinforcers, it has not been established for conditioned reinforcers. We performed an experiment with rats where lever-presses were reinforced by presentations of an audiovisual stimulus that had previously preceded food delivery; in the critical experimental groups, the audiovisual stimulus was then paired a single time with a strong electric shock. Some animals were reexposed to the audiovisual stimulus. Lever-presses yielding no outcomes were recorded in a subsequent test. Animals that had been reexposed to the audiovisual stimulus after the aversive training responded less than did those that had not received reexposure. Indeed, those animals that were not reexposed did not differ from a control group that received no aversive conditioning of the audiovisual stimulus. Moreover, these results were not mediated by a change in the food's reinforcement value, but instead reflect a change in behaviour with respect to the conditioned reinforcer itself. These are the first data to indicate that the affective value of conditioned stimuli, like that of unconditioned ones, is established when the organism comes into contact with them.

The coelacanth still lives: Bringing selection back to the fore in a science of behavior.

There is little scientific debate regarding the validity of Darwin's theory of evolution by natural selection, which effectively describes how relevant ancestral histories produce both an organism's genetic characteristics and innate behavioral repertoires. The combination of variation and selection in the production of novel forms can be extended beyond Darwinian theory to encompass facts of ontogeny. The present article sheds light on an underappreciated and critical insight, namely, that the consequences of behavior have a selective effect analogous to that observed in biological evolution. Three levels of environmental selection (phylogenic, ontogenic, and cultural) constitute a full account of the causes for action. This perspective identifies the relevant functional contingencies of which behavior is a product, it accurately and parsimoniously predicts a wide variety of disparate behavioral findings, it resolves old debates on nativism and empiricism, it unites psychological science under a central organizing principle, and it specifies psychology's position in relation to biology. Wholesale adoption of this perspective should be considered a positive advance for the field of psychology. (PsycINFO Database Record (c) 2018 APA, all rights reserved).

The influence of landmark stability on control by occasion setters.

In an operant serial feature-positive procedure, an occasion setter (OSX) signals that a response will be reinforced in the presence of a second stimulus (e.g., a discriminative stimulus, A). During a transfer test, the OS is paired with a different discriminative stimulus. Experiment 1a tested transfer effects in a touchscreen-based spatial occasion setting task with pigeons. During training, four OSs (OSW, OSX, OSY, and OSZ) were paired on separate trials with landmark A (LMA) or B (LMB) and the opportunity for a reinforced response at one location to the immediate left (R1) or right (R2) of the LM (OSW→LMA:R1, OSX→LMA:R2, OSY→LMB:R1, OSZ→LMB:R2). Training also included non-reinforced trials of LMA and LMB alone (LMA- and LMB-) and trials of a non-modulated LM with R1 and R2 reinforced across separate trials (LMC:R1 and LMC:R2). After training, the number and spatial location of responses during test trials of a LM paired with the same OS as in training did not differ reliably from transfer tests of an OS paired with a different, modulated LM (OSW→LMB and OSY→LMA), but did differ from transfer to the non-modulated LM (OSX→LMC). Experiment 1b utilized the same pigeons and training with LMB to test the degree to which the spatial stability of a LM influenced transfer. Retraining with LMA was intended to establish it as a non-modulated, stable LM (LMA:R2). Subsequent tests with LMA revealed reduced modulation by the formerly trained OS (OSW), and complete disruption of modulation of spatial location during transfer with a different OS (OSY). These findings further our understanding of the conditions under which OSs may develop and transfer modulation.

Midsession shifts in reward probability and the control of behavioral variability.

Extensive research has shown that the variability of organismal behavior is great when contingent reinforcement is delayed, small, or improbable. This research has generally employed stable response-reinforcer relationships, and therefore is limited in its explanatory scope with respect to a dynamic environment. We conducted two experiments to investigate whether pigeons' conditioned pecking behavior shows anticipatory or perseverative patterns of behavioral variability when the reinforcement probability reliably changes within experimental sessions. In Experiment 1, three pigeons received alternating sessions in which the reinforcement probability (35% or 4.2%) was shifted at the midpoint of each session in the presence of the same discrete cue. Experiment 2 featured a similar design, but with the inclusion of a discrete visual cue that changed at the session midpoint, and thus unambiguously indicated reinforcement probability. Local reinforcement rates only reliably controlled response variability when a discrete visual cue was available. Without this, pigeons did not discriminate between trial types in the first halves of sessions, and showed evidence of perseveration of response variability following a within-session shift. Critically, this is the first experimental demonstration that the relationship between reinforcement probability and behavioral variability is moderated by another factor (i.e., trial position within a session). This study thus expands our understanding of the control of behavioral variability as a function of experiential factors.

Extinction and spontaneous recovery of spatial behavior in pigeons.

We investigated extinction and spontaneous recovery of spatial associations using a landmark-based appetitive search task in a touchscreen preparation with pigeons. Four visual landmarks (A, B, C, and D) were separately established as signals of a hidden reinforced target among an 8 × 7 array of potential target locations. The target was located above landmarks (LM) A and C and below B and D. After conditioning, A and B were extinguished. Responding to A and C was assessed on probe tests 2 days following extinction, whereas, B and D were tested 14 days after extinction. We observed spontaneous recovery from spatial extinction following a 14-day, but not a 2-day, postextinction retention interval. Furthermore, by plotting the spatial distribution of responding across the X and Y axes during testing, we found that spontaneous recovery of responding to the target in our task was due to enhanced spatial control (i.e., a change in the overall distribution of responses) following the long delay to testing. These results add spatial extinction and spontaneous recovery to the list of findings supporting the assertion that extinction involves new learning that attenuates the originally acquired response, and that original learning of the spatial relationship between paired events survives extinction. (PsycINFO Database Record

Occasion setting during a spatial-search task with pigeons.

A spatial task was used to investigate if a stimulus could set the occasion for responding to a landmark. Pigeons were trained with a positive occasion setter (OS; a colored background display) signaling the contingency between a landmark (LM; visual patterned stimulus) and the location of a rewarded response. The two most common tests of an OS (transfer tests and post-training extinction of the OS) were then conducted. In Experiment 1, two occasion setting pairs were trained (+←XA/YB→+/A-/B-) with unique spatial relationships to a reinforced goal location. Transfer tests (XB- and YA-) revealed more responding to a landmark when paired with the same OS from training (e.g., XA) than on transfer tests, which was greater still than landmark-only trials (A-). Three pigeons demonstrated good spatial control of responding by the LM on transfer tests. In Experiment 2, the contingency and spatial relationship (e.g., left or right) between LM A and the goal were signaled by the OS (+←XA/YA→+/+←ZB/C→+/A-/B-). LM C was trained without an OS to assess the role of training history during transfer. Transfer tests again indicated an OS could facilitate responding and the LM controlled the location of responding. Training history affected spatial control, but not facilitation, by LM C. Lastly, post-training extinction of X had no effect on facilitation or spatial control during subsequent XA trials. These experiments are the first to evaluate conditional control of spatial information by landmarks using both of the standard tests for occasion setting.

Blocking between landmarks during 2-D (touchscreen) and 3-D (ARENA) search tasks with pigeons.

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.

Feature-positive discriminations during a spatial-search task with humans.

During feature-positive operant discriminations, a conditional cue, X, signals whether responses made during a second stimulus, A, are reinforced. Few studies have examined how landmarks, which can be trained to control the spatial distribution of responses during search tasks, might operate under conditional control. We trained college students to search for a target hidden on a computer monitor. Participants learned that responses to a hidden target location signaled by a landmark (e.g., A) would be reinforced only if the landmark was preceded by a colored background display (e.g., X). In Experiment 1, participants received feature-positive training (+←YB/ XA→+/A-/B-) with the hidden target to the right of A and to left of B. Responding during nonreinforced transfer test trials (XB-/YA-) indicated conditional control by the colored background, and spatial accuracy indicated a greater weighting of spatial information provided by the landmark than by the conditional cue. In Experiments 2a and 2b, the location of the target relative to landmark A was conditional on the colored background (+←YA/ XA→+/ ZB→+/ +←C /A-/B-). At test, conditional control and a greater weighting for the landmark's spatial information were again found, but we also report evidence for spatial interference by the conditional stimulus. Overall, we found that hierarchical accounts best explain the observed differences in response magnitude, whereas spatial accuracy was best explained via spatial learning models that emphasize the reliability, stability, and proximity of landmarks to a target.

Task complexity modifies the search strategy of rats.

Human and non-human animals exhibit a variety of response strategies (e.g., place responding) when searching for a familiar place or evading predators. We still know little about the conditions that support the use of each strategy. We trained rats to locate a hidden food reward in a small-scale spatial search task. The complexity of the search task was manipulated by reducing the number of search locations (25, 4, and 2) within an open-field apparatus and by comparison to a path-based apparatus (plus-maze). After rats were trained to reliably locate the hidden food, each apparatus was shifted to gauge whether rats were searching at the location of the goal relative to extramaze cues (i.e., place responding), or searching in the direction of the goal relative to a combination of intramaze and extramaze cues (i.e.,directional responding). The results indicate that the open field supported place responding when more than two response locations were present, whereas, the four-arm plus-maze supported strong directional responding. These results extend prior research into the role of task demands on search strategy, as well as support the use of the four-choice open field as an analog to the Morris water task for future studies targeting the neural underpinnings of place responding.

Need to train your rat? There is an App for that: a touchscreen behavioral evaluation system.

The increasing demand for highly automated and flexible tasks capable of assessing visual learning and memory in nonhuman animals has led to the exciting development of a wide array of prefabricated touchscreen-equipped systems. However, the high cost of these prefabricated systems has led many researchers to develop or modify their own preexisting equipment. We developed a freely downloadable App, the Touchscreeen Behavioral Evaluation System (TBES) for use in conjunction with an iPad (Apple, Cupertino, California) as an alternative to prefabricated touchscreen systems. TBES allows for stimulus presentation and data collection on an iPad. The touchscreen technology offered by the iPad is attractive to researchers due to its affordability, reliability, and resistance to false inputs. We highlight these, as well as the feasibility and procedural flexibility of TBES, in an effort to promote our system as a competitive alternative to those currently available.

Explorations of object and location memory using fMRI.

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.

Testing visual short-term memory of pigeons (Columba livia) and a rhesus monkey (Macaca mulatta) with a location change detection task.

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.

Visual discrimination learning with an iPad-equipped apparatus.

Visual discrimination tasks are commonly used to assess visual learning and memory in non-human animals. The current experiments explored the suitability of an iPad (Apple, Cupertino, California), as a low-cost alternative touchscreen for visual discrimination tasks. In Experiment 1, rats were trained with patterned black-and-white stimuli in a successive non-match to sample procedure. Rats successfully interacted with the iPad but failed to learn to withhold responding on trials in which the sample matched the comparison. Experiment 2 used the same patterned stimuli, but the procedure was simplified to a successive discrimination procedure and we explored the use of procedures known to facilitate discrimination learning. Rats that received training with differential outcomes and a differential reinforcement of other behavior schedule successfully acquired the task. In Experiment 3, the same rats were tested in a simultaneous discrimination task and we explored the use of a correction and non-correction method during acquisition. Rats that failed to learn the discrimination in the previous experiment, improved while trained with the correction method. These experiments support the use of the iPad in visual discrimination tasks and inform future studies investigating learning and memory within a touchscreen-equipped (iPad or other) apparatus.

Factors that influence negative summation in a spatial-search task with pigeons.

A variant of the standard conditioned inhibition procedure was used to evaluate landmark-based spatial search in a touchscreen preparation. Pigeons were given compound trials with one landmark (A) positioned in a consistent spatial relationship to a hidden goal and another landmark (B) positioned randomly with respect to A and the hidden goal (AB+). On half of the non-reinforced inhibitory trials, A was paired with landmark X (AX-) and on the remaining trials B was paired with Y (BY-). All subjects were also given reinforced trials with a transfer excitor (T+). During conditioned inhibition training, subjects showed no change in overall responding during AX- trials but did show a decrease in the number of pecks to the goal location signaled by A. During non-reinforced summation tests with landmark T, X had a greater suppressive effect than did Y on overall responding but the percentage of pecks at the goal did not differ unless X was positioned near the expected goal signaled by T. These data demonstrate that the effectiveness of a stimulus trained as an inhibitor is dependent on the strength of the association between its training excitor (A) and the US, as well as, the spatial arrangement of stimuli during testing.

WITHDRAWN: fMRI correlates of visual working memory: What vs. where.

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.

Visual short-term memory compared in rhesus monkeys and humans.

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.

Overshadowing between landmarks on the touchscreen and in arena with pigeons.

The role of generalization decrement in spatial overshadowing was evaluated using a landmark-based spatial search task in both a touchscreen preparation (Experiment 1a) and in an Automated Remote Environmental Navigation Apparatus (ARENA, Experiment 1b). A landmark appeared as a colored circle among a row of eight (touchscreen) or six (ARENA) potential locations. On overshadowing trials, Landmark X was located two positions away from a hidden goal, while another landmark, A, was in the position between X and the goal. On control trials, Landmark Y was positioned two locations away from the goal but without a closer landmark. All subjects were then tested with separate trials of A, X, Y, and BY. Testing revealed poor spatial control by X relative to A and Y, thereby replicating the spatial overshadowing effect. Spatial control by Y was similar when tested in compound with novel landmark (BY) and on trials of Y alone. Thus, overshadowing in a small-scale environment does not appear to be due to a process of generalization decrement between training and testing.