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Higher-order conditioning results from a simple training procedure: Pairing two relatively neutral conditioned stimuli, A and X, allows properties separately conditioned to X (e.g., through pairing it with an unconditioned stimulus, US) to be evident during A. The phenomenon extends the range of ways in which Pavlovian conditioned responding can be expressed and increases its translational relevance. Given this relevance and the wealth of available behavioral analysis, it is a surprisingly underdeveloped territory for formal theoretical analysis. Here, we first provide a critical review of two (informal) classes of account for higher-order conditioning that reflect either: (a) processes that are analogous to Pavlovian conditioning, but involving associatively activated representations (e.g., A→US); or (b) the formation of an associative chain (e.g., A→X, and X→US). Our review first identifies fundamental theoretical and empirical challenges to both classes of account. We then develop a new computational model of higher-order conditioning that meets the challenges identified by showing: how reciprocal associations between A, X, and the US are formed and affect performance; and how the similarity of stimuli, their traces, and associatively retrieved representations modulate this process. The model generates a wealth of novel predictions, providing a platform for further empirical and theoretical analysis. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Robert A. Rescorla changed how Pavlovian conditioning was studied and interpreted. His empirical contributions were fundamental and theoretically driven. One involved testing a central tenet of the model that he developed with Allan R. Wagner. The Rescorla-Wagner learning rule uses a pooled error term to determine changes in a directional association between the representations of the conditioned stimulus (CS) and unconditioned stimulus (US). This learning rule predicts that 2 equally salient CSs (A and B) will undergo equivalent associative change when they are conditioned in compound (i.e., AB→US). Rescorla's results suggested that this was not the case (e.g., Rescorla, 2000). Here, we show that these results can be reconciled with a model that uses a learning rule with a pooled error term once that rule is applied equivalently to all of the stimuli presented on a given trial, and the resulting reciprocal associations (directly and indirectly) contribute to performance. This model, called HeiDI, integrates several features of Rescorla's research and theorizing while addressing an issue that he recognized required further analysis: how learning is translated into performance. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Exposure to two similar stimuli (AX and BX; e.g., two tastes) reduces the extent to which a conditioned response later established to BX generalizes to AX. This example of perceptual learning is more evident when AX and BX are exposed in an alternating manner (AX, BX, AX, BX,…) than when AX and BX occur in separate blocks (e.g., AX, AX,…BX, BX,…). We examined in male rats (N = 126) the impact of rapid alternation to AX and BX on generalization of a taste aversion from BX to AX. Experiment 1 showed that such alternating presentations (with 5-min intervals between AX and BX) reduced generalization relative to blocked exposure; but only as assessed by consumption levels and not by lick cluster size (an index of hedonic reactions). Experiment 1 also showed that the nature of exposure did not affect how A influenced performance to a novel conditioned taste, Y. Experiment 2 replicated the pattern of results involving the different influences of rapidly alternating and blocked exposure on generalization from BX to AX, and showed that this effect was only evident when rats received access to water during the 5-min intervals between AX and BX. These results reinforce parallels between perceptual learning effects in rats and humans, both at empirical and theoretical levels. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Associative treatments of how Pavlovian conditioning affects conditioned behavior are rudimentary: A simple ordinal mapping is held to exist between the strength of an association (V) between a conditioned stimulus (CS) and an unconditioned stimulus (US; i.e., V CS-US ) and conditioned behavior in a given experimental preparation. The inadequacy of this simplification is highlighted by recent studies that have taken multiple measures of conditioned behavior: Different measures of conditioned behavior provide the basis for drawing opposite conclusions about V CS-US across individual animals. Here, we develop a simple model involving reciprocal associations between the CS and US (V CS-US and V US-CS ) that simulates these qualitative individual differences in conditioned behavior. The new model, HeiDI (How excitation and inhibition Determine Ideo-motion), enables a broad range of phenomena to be accommodated, which are either beyond the scope of extant models or require them to appeal to additional (learning) processes. It also provides an impetus for new lines of inquiry and generates novel predictions. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

Pavlovian conditioning procedures produce marked individual differences in the form of conditioned behavior. For example, when rats are given conditioning trials in which the temporary insertion of a lever into an operant chamber (the conditioned stimulus, CS) is paired with the delivery of food (the unconditioned stimulus, US), they exhibit knowledge of the lever-food relationship in different ways. For some rats (known as sign-trackers), interactions with the lever dominate, while for others (goal-trackers), approaching the food well dominates. A formal model of Pavlovian conditioning (HeiDI) attributes such individual differences in behavior to variations in the perceived salience of the CS and US. An application of the model in which the perceived salience of the CS declines (i.e., adapts) across its duration predicts changes in these individual differences within the presentation of the CS: The sign-tracking bias is predicted to decline and goal-tracking bias is predicted to increase across the presentation of a lever. The accuracy of these predictions was confirmed through analysis of archival data from female and male rats. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

The model elaborated here adapts the influential pooled error term, first described by Wagner and Rescorla (Rescorla & Wagner, 1972; Wagner & Rescorla, 1972), to govern the formation of reciprocal associations between any pair of stimuli that are presented on a given trial. In the context of Pavlovian conditioning, these stimuli include various conditioned and unconditioned stimuli. This elaboration enables the model to deal with cue competition phenomena, including the relative validity effect, and evidence implicating separate error terms and attentional processes in association formation. The model also includes a performance rule, which provides a natural basis for (individual) variation in the strength and nature of conditioned behaviors that are observed in Pavlovian conditioning procedures. The new model thereby begins to address theoretical and empirical issues that were apparent when the Rescorla-Wagner model was first described, together with research inspired by the model over the ensuing 50 years. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

Texture is often used as a convenient stimulus dimension to study aspects of discrimination learning in rodents. However, the basis of texture discrimination learning is often left untested: Although learning could involve the whisker system, it could also be based on other senses (e.g., olfactory or visual). Here, we investigated whether mice use their whisker system to learn texture discriminations. Mice were placed in an apparatus illuminated with a dim red light, and the mice had to learn which of 2 sawdust-filled bowls contained a buried reward. The outer surfaces of the bowls were 3-D printed with different textures (grooved or smooth). Within a 60-min session, mice learned to dig in 1 bowl (e.g., grooved) rather than the other (e.g., smooth) to gain the reward. This texture discrimination and an equivalent odor discrimination were retained overnight (Experiments 1 and 2); and whisker trimming disrupted learning based on the texture of the bowls but not learning based on the odor of the sawdust in the bowls (Experiments 3 and 4). These results provide a secure basis upon which to investigate the behavioral and brain basis of texture learning in rodents. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Pavlovian conditioning procedures result in dramatic individual differences in the topography of learnt behaviors in rats: When the temporary insertion of a lever into an operant chamber is paired with food pellets, some rats (known as sign-trackers) predominantly interact with the lever, while others (known as goal-trackers) predominantly approach the food well. Two experiments examined the sensitivity of these two behaviors to changing reinforcement contingencies in groups of male and female rats exhibiting the different phenotypes (i.e., sign-trackers and goal-trackers). In both phenotypes, behavior oriented to the food well was more sensitive to contingency changes (e.g., a reversal in which of two levers was reinforced) than was lever-oriented behavior. That is, the nature of the two behaviors differed independently of the rats in which they were manifest. These results indicate that the behavioral phenotypes reflect the parallel operation of a stimulus-stimulus associative process that gives rise to food-well activity and a stimulus-response process that gives rise to lever-oriented activity, rather than the operation of a single process (e.g., stimulus-stimulus) that generates both behaviors. (PsycINFO Database Record (c) 2018 APA, all rights reserved).

Laboratory rats can exhibit marked, qualitative individual differences in the form of acquired behaviors. For example, when exposed to a signal-reinforcer relationship some rats show marked and consistent changes in sign-tracking (interacting with the signal; e.g., a lever) and others show marked and consistent changes in goal-tracking (interacting with the location of the predicted reinforcer; e.g., the food well). Here, stable individual differences in rats' sign-tracking and goal-tracking emerged over the course of training, but these differences did not generalize across different signal-reinforcer relationships (Experiment 1). This selectivity suggests that individual differences in sign- and goal-tracking reflect differences in the value placed on individual reinforcers. Two findings provide direct support for this interpretation: the palatability of a reinforcer (as measured by an analysis of lick-cluster size) was positively correlated with goal-tracking (and negatively correlated with sign-tracking); and sating rats with a reinforcer affected goal-tracking but not sign-tracking (Experiment 2). These results indicate that the observed individual differences in sign- and goal-tracking behavior arise from the interaction between the palatability or value of the reinforcer and processes of association as opposed to dispositional differences (e.g., in sensory processes, "temperament," or response repertoire). (PsycINFO Database Record, ((c) 2016 APA, all rights reserved).)

Four experiments with male rats investigated perceptual learning involving a tactile dimension (A, B, C, D, E), where A denotes 1 end of the continuum (e.g., a rough floor) and E the other (e.g., a smooth floor). In Experiment 1, rats given preexposure to A and E learned an appetitive discrimination between them more readily than those not given preexposure. Experiment 2a showed that rats preexposed to B and D acquired a discrimination between A and E more readily than those preexposed to A and E; and in Experiment 2b the same preexposure treatments had no effect on the acquisition of a discrimination between B and D. In Experiments 3a and 3b, rats given preexposure to C learned a discrimination between A and E more readily than those not given preexposure. Experiment 4 demonstrated that preexposure to a texture (e.g., B) that was adjacent to the to-be-discriminated textures (e.g., C and E) facilitated a discrimination between them relative to preexposure to their midpoint (D). These novel perceptual learning effects are interpreted as reflecting a redistribution of processing between the notional elements of the texture dimension. (PsycINFO Database Record, ((c) 2016 APA, all rights reserved).)

Decisions made by operational commanders at emergency incidents have been characterized as involving a period of information gathering followed by courses of action that are often generated without explicit plan formulation. We examined the efficacy of goal-oriented training in engendering explicit planning that would enable better communication at emergency incidents. While standard training mirrored current operational guidance, goal-oriented training incorporated "decision controls" that highlighted the importance of evaluating goals, anticipated consequences, and risk/benefit analyses once a potential course of action has been identified. In Experiment 1, 3 scenarios (a house fire, road traffic collision, and skip fire) were presented in a virtual environment, and in Experiment 2 they were recreated on the fireground. In Experiment 3, the house fire was recreated as a "live burn," and incident commanders and their crews responded to this scenario as an emergency incident. In all experiments, groups given standard training showed the reported tendency to move directly from information gathering to action, whereas those given goal-oriented training were more likely to develop explicit plans and show anticipatory situational awareness. These results indicate that training can be readily modified to promote explicit plan formulation that could facilitate plan sharing between incident commanders and their teams., ((c) 2015 APA, all rights reserved).)

In order to evaluate 1 account for the asymmetry that has been found with discriminations based on stimulus magnitude, in 5 autoshaping experiments, 2 groups of pigeons received a discrimination between 5 and 20 squares presented on a TV screen. One group received a 20+/5- discrimination, with food signaled by 20 squares but not 5 squares; the other group received the opposite discrimination, 5+/20-. The 20+/5- discrimination was acquired more readily than 5+/20- in Experiments 1, 3a, 3b, and 4. For Experiment 1, the screen was white for the intertrial interval (ITI) and the stimuli were black squares on a white background; for Experiment 3a, the screen was black for the ITI and the stimuli were black squares on a white background; and for Experiments 3b and 4, the screen was white for the ITI and the stimuli were white squares on a black background. In Experiment 2, the stimuli were black squares on a white background, but they were separated by an ITI in which 288 black squares were presented against a white background. The 20+/5- discrimination was now acquired more slowly than the 5+/20- discrimination. The asymmetry in the acquisition of the magnitude discriminations in each experiment is attributed to inhibition being associated with the stimuli present during the ITI. The generalization of this inhibition, along a dimension related to the number of squares on the screen, is then assumed to disrupt the acquisition of 1 discrimination to a greater extent than the other., ((c) 2015 APA, all rights reserved).)

The formation of an integrated memory for a pattern of stimulation could be based on the elements of that pattern becoming directly linked to one another, or by each of the elements becoming linked to a shared separate configural representation. These 2 accounts have proven difficult to discriminate between. Here, rats received exposure to four patterns of stimulation, each consisting of an auditory stimulus, a visual context, and a time of day; and we examined whether pre-training lesions to the hippocampus influenced memory for the patterns. These lesions abolished pattern memory that required configural processes (Experiments 1A and 1B) but had no effect on pattern memory that could be supported by elemental processes (Experiment 2). This dissociation provides support for the views that elemental and configural processes ordinarily support pattern memory and that rats with lesions to the hippocampus are left reliant on elemental processes. (, (PsycINFO Database Record (c) 2011 APA, all rights reserved).)

Rats with lesions of the perirhinal cortex and a control group were required to find a platform in 1 corner of a white rectangle and in the reflection of this corner in a black rectangle. Test trials revealed that these groups were able to integrate information regarding the shape of the pool and the color of its walls (black or white) to identify the correct location of the platform. A clear effect of the perirhinal cortex lesions was, however, revealed using an object recognition task that involved the spontaneous exploration of novel objects. The results challenge the view that the perirhinal cortex enables rats to solve discriminations involving feature ambiguity.

In the present study, the authors examined the role of the medial prefrontal cortex in acquired equivalence and distinctiveness of cues. Rats were placed in 4 experimental contexts (A, B, C, and D) where they received presentations of 2 auditory stimuli (X and Y). In Contexts A and B, X was paired with food and Y was not, whereas in Contexts C and D, Y was paired with food and X was not. Rats that received sham lesions and those with lesions of the medial prefrontal cortex acquired this configural discrimination equally readily. Rats then received many pellets in A but not in C. After this training, sham-lesion rats exhibited more magazine activity in B than in D (an acquired distinctiveness/acquired equivalence effect), whereas those with medial prefrontal cortex lesions did not. These results indicate that the medial prefrontal cortex is involved in the process by which experience with stimuli influences the degree of generalization between them.

Dual-process models of recognition memory in animals propose that recognition memory is supported by two independent processes that reflect the operation of distinct brain structures: a familiarity process that operates independently of the hippocampus and a context-dependent (episodic) memory process that is dependent on the hippocampus. A novel variant of an object recognition procedure was used to examine this proposal. Healthy rats showed a preference for exploring a novel object rather than a familiar object: a familiarity-dependent recognition effect. They also showed a preference for exploring a familiar object that was presented in a different spatiotemporal context rather than a familiar object that was presented either in a different spatial or temporal context: a context-dependent form of recognition that is sensitive to "what" object has been presented "where" and "when." Rats with excitotoxic hippocampal lesions showed the familiarity-dependent but not the context-dependent form of recognition. The results provide support for dual-process theories of recognition memory., (Copyright (c) 2007 APA, all rights reserved.)

The ability of rats with ibotenate lesions of the entorhinal cortex to form memories for events was assessed by using a gustatory within-event learning procedure. Rats first received exposure to 2 events, AX and BY, each composed of a pair of flavors. Following this exposure period, Flavor X alone was paired with the delivery of lithium chloride. Lesioned and control rats showed a greater aversion to A than to B and to AX than to BX. These results challenge theories that suppose that the entorhinal cortex plays a general role in forming representations of patterns of stimulation., ((c) 2005 APA)

In associative priming, rats are more likely to orient to a visual stimulus whose memory has not been recently activated (V1) than to one whose memory has been recently activated (V2). However, rats with excitotoxic hippocampal lesions are more likely to orient to the primed V2 than to the unprimed V1. This study investigated the influence of hippocampal lesions on nonassociative priming. Rats received presentations of 2 visual stimuli, V1 and V2, that had been presented more (V2, primed) or less (V1, unprimed) recently. Control rats oriented to V1 rather than to V2, whereas hippocampal rats oriented to V2 rather than to V1. These results parallel those observed in an associative priming procedure and thereby suggest that the role of the hippocampus in priming is general.

This study with rats examined the effects of excitotoxic lesions to the entorhinal cortex (EC) and hippocampus (HPC) on using extramaze and intramaze cues to navigate to a hidden platform in a water maze. HPC lesions resulted in a disruption to the use of extramaze cues, but not intramaze cues, whereas EC lesions had no effect on the use of these cues when they were encountered for the first time. However, prior navigation training in which 1 type of cue was relevant disrupted navigation with the other type in rats with EC lesions. Results show that the EC contributes to the processing of spatial information, but that this contribution is most apparent when there is a conflict between 2 sources of navigational cues in the water maze.

The prefrontal cortex in humans has been implicated in processes that underlie novelty detection and attention. This study examined the contribution of the rat medial prefrontal cortex to novelty detection using the targeting, or orienting, response (OR) as a behavioral index. Lesions to the medial prefrontal cortex (specifically the prelimbic and infralimbic cortices) influenced neither the OR to a novel visual stimulus from a localized light source (V1), nor the change in this OR over the course of a series of exposures to V1. However, after exposure to V1, the OR to a 2nd visual stimulus from the same source, V2, was more pronounced in control rats than in lesioned rats. These results suggest that the medial prefrontal cortex in the rat contributes to the process of novelty detection.

Learning about contextual cues is markedly disrupted in rats with hippocampal lesions. One analysis of this disruption is that it reflects a general failure to form associations between the elements of complex events. A straightforward prediction of this analysis is that sensory preconditioning will be disrupted by hippocampal lesions. This prediction was assessed by presenting rats with flavored solutions composed of 2 elements (A and X) before X was paired with an injection of the emetic, lithium chloride. A subsequent test revealed that rats were less willing to consume Solution A than they were to consume a control solution, B. This was true of rats with sham lesions and those with excitotoxic lesions of hippocampus. These findings fail to support the proposition that the hippocampus-dependent deficit in contextual conditioning is due to a general disruption to the process of associating the elements of complex events.

Novel behavioral assays were used to assess the role of the entorhinal cortex in modulating attention to components of stimulus compounds. In Stage 1, rats received discrimination training with compounds constructed from 3 dimensions (auditory, visual, and tactile); in each compound the combination of components from 2 dimensions (e.g., auditory and visual) were relevant to the solution of the discrimination, and the remaining dimension (e.g., tactile) was irrelevant. In Stage 2, rats received a different discrimination in which the relevant dimensions were either congruent (auditory and visual) or incongruent (auditory and tactile) with those that were relevant in Stage 1. Sham-operated rats acquired the congruent discrimination more rapidly than the incongruent discrimination--a finding indicative of a process of attentional modulation--whereas rats with excitotoxic lesions of the entorhinal cortex acquired both discriminations equally readily.

This study sought to characterize the effects of removing the nuclei of primary importance in relaying the thalamic head direction signal to the hippocampal formation (the anterior dorsal [AD] and lateral dorsal [LD] nuclei) on the performance of a variety of spatial and nonspatial tasks. The results indicate that combined excitotoxic lesions of the AD and LD nuclei produce marked deficits on a variety of spatial tasks. These tasks included T-maze alternation and the ability to locate a hidden platform set at a fixed distance and fixed direction from a beacon in a Morris water maze. Although object recognition appeared unaffected, marked impairments were found in the ability to detect when an object was placed in a novel position (object-in-place memory).

Rats received excitotoxic lesions of different subsystems within the hippocampal system--either the hippocampus proper (cornu ammonis and dentate gyrus; hippocampal lesions) or the entorhinal cortex-subicular region (entorhinal lesions). Subsequently, their activity in an operant chamber was monitored both before and after footshock had been delivered (Experiment 1). Rats with hippocampal lesions showed enhanced activity before the delivery of footshock and reduced freezing after the delivery of shock. Rats with entorhinal lesions showed control levels of activity before the delivery of footshock and control levels of freezing after the delivery of footshock. Both types of lesion impaired spatial learning in a water maze (Experiment 2). These results suggest that the deficits arising from damage to the hippocampal system in contextual and spatial learning have different origins.

Precocial chicks exposed to a stimulus subsequently approach that stimulus in preference to other, novel stimuli. Previous investigations of the neural basis of these imprinting preferences suggest that imprinting training results in the formation of two memories. The first memory is formed rapidly and is located in the intermediate and medial part of the hyperstriatum ventrale (IMHV) of the left hemisphere; the formation of the second, in another memory system, S', takes several hours and can be prevented by a lesion placed in the right IMHV soon after training. The results of the present study suggest that the functional characteristics of these memories differ. Although memories in both left IMHV and S' supported imprinting preferences (Experiments 1a and 2a), only memories in S' influenced the acquisition of a heat-reinforced discrimination in which imprinted objects served as discriminanda (Experiments 1b and 2b).

The contextual specificity of the conditioned response (CR) and latent inhibition (LI) was examined in rats with selective hippocampal lesions. Acquisition of the CR to a novel conditioned stimulus (CS) was equally rapid in control and hippocampal rats (Experiments 1 and 2), and CS preexposure disrupted acquisition, (i.e., produced LI) to an equal extent in both groups (Experiment 2). In control subjects, however, the CR established in one context transferred incompletely to a second context (Experiment 1), and LI was attenuated when CS preexposure and conditioning occurred in different contexts (Experiment 3). This context specificity of the CR and LI was not apparent in hippocampal rats--the CR and LI transferred readily from one context to another. In addition, hippocampal rats were impaired in a spatial learning task (Experiment 2) but were unimpaired in learning a Pavlovian contextual discrimination (Experiment 3). These results suggest that a common contextual retrieval process underlies the contextual dependence of the CR and of LI and that this process is mediated by the hippocampus.

The role of the hippocampus in associative learning was investigated in 3 experiments with rats as subjects. Hippocampal rats were impaired in the acquisition of conditioned responding both when food was signaled by the insertion of a lever (Experiment 1) and when the presentation of auditory or visual events served as the conditioned stimuli (Experiment 2). Experiment 2 also evaluated the suggestion that deficits in the acquisition of conditioned responding reflect the failure of hippocampal subjects to use contextual cues to retrieve associative information. This experiment showed that hippocampal rats were impaired in learning that a given stimulus was reinforced in Context A but nonreinforced in Context B. Experiment 3 demonstrated that hippocampal rats were unimpaired in learning a simple Pavlovian contextual discrimination. This pattern of results suggests that the hippocampus is involved in a higher order contextual retrieval process.