Gonzalo P Urcelay

• PhD
• Professor (Assistant) at University of Nottingham

Goal-directed actions are defined by their sensitivity to the causal association between actions and outcomes, as well as the subjective value ascribed to those outcomes. When this sensitivity diminishes, actions may transition into habitual behavior. Based on recent findings from animal studies, we hypothesized that delaying outcomes relative to a...

A canonical view in the neuroscience of learning and memory literature is that failures in memory expression reflect storage failures, and hence, amnesic manipulations following training or following memory reactivation can permanently erase memory traces. In this review, we analyze extant literatures from the learning and memory domains suggesting...

Goal-directed actions are defined by their sensitivity to the causal association between actions and outcomes, as well as the subjective value ascribed to those outcomes. When this sensitivity diminishes, actions may transition into habitual behavior. Based on recent findings from animal studies, we hypothesized that delaying outcomes relative to a...

Previous studies involving birds and humans have identified spatial proximity as a source of overshadowing between landmarks in navigation. In Herrera et al. (2024), subjects were trained in an open environment to find a hidden goal with reference to a cross-shaped array of four landmarks placed at various distances from it. Critically, two of the...

Safety signals reinforce instrumental avoidance behavior in nonhuman animals. However, there are no conclusive demonstrations of this phenomenon in humans. Using human participants in an avoidance task, Experiments 1–3 and 5 were conducted online to assess the reinforcing properties of safety signals, and Experiment 4 was conducted in the laborator...

Although numerous behavioral constructs have been proposed to account for anxiety disorders, how these disorders develop within an individual has been difficult to predict. In this perspective, I selectively review clinical and experimental evidence suggesting that avoidance (i.e., safety) behavior increases beliefs of threat or fear. The experimen...

The ABA renewal effect occurs when behavior is trained in one context (A), extinguished in a second context (B), and the test occurs in the training context (A). Two mechanisms that explain ABA renewal are context summation at the test and contextual modulation of extinction learning, with the former being unlikely if both contexts have a similar a...

Cancer patients often experience anticipatory nausea and vomiting (ANV) due to Pavlovian conditioning. Both N-methyl-D-aspartate and beta-adrenergic receptors are known to mediate memory formation, but their role in the development of ANV remains unclear. This study used a conditioned context aversion (CCA) paradigm, an animal model for ANV, to ass...

A canonical view in the neuroscience of learning and memory literature is that failures in memory expression reflect storage failures, and hence amnesic manipulations following training or following memory reactivation can permanently erase memory traces. In this review, we analyse extant literatures from the learning and memory domains suggesting...

Instrumental behaviour is goal-directed when controlled by the value of the outcome and knowledge of the action-outcome contingency, otherwise it is deemed to be controlled by external stimuli (habit). It is accepted that stress modulates the control of instrumental actions in a manner that favours habitual over goal-directed control. Indeed, while...

Although numerous behavioural constructs have been proposed to account for anxiety disorders, how these develop within an individual has been difficult to predict. In this perspective, I selectively review clinical and experimental evidence suggesting that avoidance (i.e., safety) behaviour increases beliefs of threat or fear. The experimental evid...

Generalization enables individuals to respond to novel stimuli based on previous experiences. The degree to which organisms respond is determined by their physical resemblance to the original conditioned stimulus (CS+), with a stronger response elicited by more similar stimuli, resulting in similarity-based generalization gradients. Recent research...

Safety signals provide relief and have been found to reinforce instrumental avoidance behaviour in non-human animals. However, the available evidence in humans has been obtained with subjective measures and hence there are no objective demonstrations of this phenomenon in humans. Using human participants in an avoidance task, three experiments were...

Goodyear and Kamil (2004) assessed the ability of Clark’s nutcrackers to find buried food based on a cross-shaped array of landmarks at different distances from the goal. Their findings suggested that proximal landmarks overshadowed learning about distal landmarks, and this was attenuated when assessing the effect of distal landmarks on learning ab...

Three experiments explored whether weakening temporal contiguity between auditory cues and an aversive outcome attenuated cue-competition in an avoidance learning task with human participants. Overall, with strong temporal contiguity between auditory cues and the outcome during training (the offset of the predictive auditory signals concurred with...

In situations in which multiple predictors anticipate the presence or absence of an outcome, cues compete to anticipate the outcome, resulting in a loss of associative strength compared to control conditions without additional cues. Critically, there are multiple factors modulating the magnitude and direction of such competition, although in some s...

Anticipatory nausea (AN) is caused by an association between contextual cues and the experience of nausea (the side effects of chemotherapy or radiation treatment) and it develops predominantly in female patients undergoing chemotherapy. Preclinical studies in rodents show that the administration of an illness-inducing agent in the presence of nove...

The objective of the present study was to develop a model of avoidance learning and its extinction in planarians (Schmidtea mediterranea). Based on previous experiments showing conditioned place preference, we developed a procedure to investigate conditioned place avoidance (CPA) using shock as an unconditioned stimulus (US) and an automated tracki...

When multiple cues are associated with the same outcome, organisms tend to select between the cues, with one revealing greater behavioral control at the expense of the others (i.e., cue competition). However, non-human and human studies have not always observed this competition, creating a puzzling scenario in which the interaction between cues can...

In a large variety of contexts, it is essential to use the available information to extract patterns and behave accordingly. When it comes to social interactions for instance, the information gathered about interaction partners across multiple encounters (e.g., trustworthiness) is crucial in guiding one’s own behavior (e.g., approach the trustworth...

In this chapter, we summarize the history of behaviourism and discuss some critical developments that have shaped the way we currently think about behaviour. We then discuss phenomena that led researchers in the behaviourist tradition to conclude that nonhuman animals represent the consequences of their behaviour, can associate absent events which...

The chapter provides an overview of Pavlovian conditioning

Three experiments (n = 81, n = 81, n = 82, respectively) explored how temporal contiguity influences Action-Outcome learning, assessing whether an intervening signal competed, facilitated, or had no effect on performance and causal attribution in undergraduate participants. Across experiments, we observed competition and facilitation as a function...

In a signal detection theory approach to associative learning, the perceived (i.e., subjective) contingency between a cue and an outcome is a random variable drawn from a Gaussian distribution. At the end of the sequence, participants report a positive cue-outcome contingency provided the subjective contingency is above some threshold. Some researc...

Over the last 50 years, cue competition phenomena have shaped theoretical developments in animal and human learning. However, recent failures to observe competition effects in standard conditioning procedures, as well as the lengthy and ongoing debate surrounding cue competition in the spatial learning literature, have cast doubts on the generality...

Research on retrieval-induced malleability of maladaptive emotional memories has been mostly focused on the effect of drugs and extinction (i.e. post-retrieval extinction). Only a few studies addressed post-retrieval appetitive-aversive interactions. Due to the relevance that the understanding of the interactions between memory content and appetiti...

In anticipation of palatable food, rats can learn to restrict consumption of a less rewarding food type resulting in an increased consumption of the preferred food when it is made available. This construct is known as anticipatory negative contrast (ANC) and can help elucidate the processes that underlie binge-like behavior as well as self-control...

In anticipation of palatable food, rats can learn to restrict consumption of a less rewarding food type resulting in a binge on the preferred food when it is made available. This construct is known as anticipatory negative contrast (ANC) and can help elucidate the processes that underlie binge-like behavior as well as self-control in rodent motivat...

Chronic nicotine exposure reduces sensitivity to the effects of nicotine, which then results in behavioural changes and tolerance development. In the planaria, a valuable first-stage preclinical model for addictive behaviour, acute nicotine administration has been shown to steadily alter the motility of the animals, a result that has been interpret...

In planarians, as seen in rodents, natural reinforcers (sucrose) and drugs of abuse support Conditioned Place Preference (CPP), which is a form of Pavlovian learning to examine the rewarding effects of natural reinforcers and drugs of abuse. Using this preparation, we have previously observed acquisition, extinction and reinstatement of sucrose CPP...

Two experiments address the habituation of photonegative and exploratory responses in the flatworm planaria (Dugesia). Planarians possess a well-documented photonegative response; Experiment 1 showed that repeated exposures to a bright light source with short inter trial intervals (ITIs) within 1 experimental session gradually weakens the unconditi...

In this chapter, we first introduce classical conditioning from a historical perspective. We then characterize what is learned during classical conditioning, and do so while noting some practical and functional implications of classical conditioning. In the second part, we provide a succinct overview of phenomena such as cue competition and interfe...

Avoidance behaviour is a hallmark of anxiety disorders and OCD, yet there is a pressing need to understand how extinction of avoidance responses proceeds and what variables if any determine recovery from extinction. In this study, we used a within-subjects design in which participants learned to avoid a loud noise signalled by two discrete visual s...

The transition from goal-directed to habitual forms of instrumental behavior is determined by variables such as the amount of training, schedules of reinforcement, the availability of choices, and exposure to drugs of abuse. Less is known about the control of goal-directed behavior when reinforcement is delayed rather than immediate. In these exper...

There has been a recent surge of interest in avoidance behavior and potential ways to minimize it, given its critical role in psychiatric conditions such as anxiety disorders and obsessive-compulsive disorder (OCD). In this review, we summarize recent work investigating the extinction of avoidance behavior in humans and rodents. We focus on tasks,...

Excessive checking is reported in non-clinical populations and is a pervasive symptom in obsessive compulsive disorder (OCD). We implemented a free-operant task in humans, previously used in rats, wherein participants can “check” to reduce uncertainty. Participants can press an observing key to ascertain which of two main keys will, if pressed, cur...

Prével and colleagues reported excitatory learning with a backward conditioned stimulus (CS) in a conditioned reinforcement preparation. Their results add to existing evidence of backward CSs sometimes being excitatory and were viewed as challenging the view that learning is driven by prediction error reduction, which assumes that only predictive (...

Historically, most approaches to understanding learning and memory phenomena, particularly at the neurobiological level, have emphasized information processing that occurs during or soon after training (i.e., acquisition) as critical for observing learned changes in behavior. However, this view has been challenged by studies showing that at least p...

For decades, it has been observed that stress affects cognition and behaviour through a number of different mechanisms. Recently, reports in rodents and humans have shown that stress prompts habit formation, as assessed through outcome devaluation tests during extinction. However, stress also promotes resistance to extinction, so it is not clear wh...

Despite the generality and theoretical relevance of cue competition phenomena such as blocking and overshadowing, recent findings suggest that these observations may be due to some degree of publication bias, and that we lack insight into the boundary conditions of these phenomena. The present commentary does not question the existence of cue compe...

RationaleCorticostriatal circuits are widely implicated in the top-down control of attention including inhibitory control and behavioural flexibility. However, recent neurophysiological evidence also suggests a role for thalamic inputs to striatum in behaviours related to salient, reward-paired cues. Objectives Here, we used designer receptors excl...

Two experiments using rats in a contextual fear memory preparation compared two approaches to reduce conditioned fear: (1) pharmacological reconsolidation blockade and (2) reactivation-plus-extinction training. In Experiment 1, we explored different combinations of reactivation-plus-extinction parameters to reduce conditioned fear and attenuate rea...

Historically, most approaches to understanding learning and memory phenomena, particularly at the neurobiological level, have emphasized information processing that occurs during or soon after training (i.e., acquisition) as critical for observing learned changes in behavior. However, this view has been challenged by studies showing that at least p...

In the present study, excitatory backward conditioning was assessed in a conditioned reinforcement paradigm. The experiment was conducted with human subjects and consisted of five conditions. In all conditions, US reinforcing value (i.e. time reduction of a timer) was assessed in phase 1 using a concurrent FR schedule, with one response key leading...

Associative learning theories regard the probability of reinforcement as the critical factor determining responding. However, the role of this factor in instrumental conditioning is not completely clear. In fact, a wealth of evidence from instrumental free-operant experiments has shown that participants respond at a higher rate on variable ratio th...

This chapter provides a contemporary review of the existing literature on the associative basis of avoidance, synthesizing historic debate with empirical study in rodents and humans from the fields of behavioral, cognitive, and neuroscience research. The chapter outlines a consensus view of the conditions necessary for the acquisition and maintenan...

Obsessive Compulsive Disorder (OCD) has been linked to numerous cognitive theories, such as increased intolerance of uncertainty. Certainty seeking may be one process underlying excessive checking behavior. We sought to obtain objective and quantitative measurements of certainty seeking behaviors in patients in a translational task. A rodent versio...

Different mnemonic outcomes have been observed when associative memories are reactivated by CS exposure and followed by amnestics. These outcomes include mere retrieval, destabilization-reconsolidation, a transitional period (which is insensitive to amnestics), and extinction learning. However, little is known about the interaction between initial...

This protocol details a free-operant avoidance paradigm that has been developed to evaluate the relative contribution of different sources of reinforcement of avoidance behavior that may play an important role in the development and maintenance of human anxiety disorders. The task enables the assessment of the effects of safety cues that signal a p...

Objective: The purpose of this study was to determine the neural correlates of excessive habit formation in obsessive-compulsive disorder (OCD). The authors aimed to test for neurobiological convergence with the known pathophysiology of OCD and to infer, based on abnormalities in brain activation, whether these habits arise from dysfunction in the...

Safety signals provide "relief" through predicting the absence of an aversive event. At issue is whether these signals also act as instrumental reinforcers. Four experiments were conducted using a free-operant lever-press avoidance paradigm in which each press avoided shock and was followed by the presentation of a 5-sec auditory safety signal. Whe...

The associative processes that support free-operant instrumental avoidance behavior are still unknown. We used a revaluation procedure to determine whether the performance of an avoidance response is sensitive to the current value of the aversive, negative reinforcer. Rats were trained on an unsignaled, free-operant lever press avoidance paradigm i...

Although contexts play many roles during training and also during testing, over the last four decades theories of learning have predominantly focused on one or the other of two families of functions served by contexts. In this selective review, we summarize recent data concerning these two functions and their interrelationship. The first function i...

Excessive checking is a common, debilitating symptom of obsessive-compulsive disorder (OCD). In an established rodent model of OCD checking behaviour, quinpirole (dopamine D2/3-receptor agonist) increased checking in open-field tests, indicating dopaminergic modulation of checking-like behaviours. We designed a novel operant paradigm for rats (obse...

It has been suggested that, unlike pure extinction which typically results in the return of the fear response under a variety of circumstances, memory reactivation followed by extinction can attenuate the reemergence of conditioned fear. The reactivation-extinction procedure has attracted the attention of basic and clinical researchers due to its p...

Safety signals (SS) have been shown to reinforce instrumental avoidance behaviour due to their ability to signal the absence of an aversive event; however, little is known of their neural mediation. The present study investigated whether infusions of d-amphetamine in the nucleus accumbens (Nac), previously shown to potentiate responding for appetit...

Most models of human and animal learning assume that learning is proportional to the discrepancy between a delivered outcome and the outcome predicted by all cues present during that trial (i.e., total error across a stimulus compound). This total error reduction (TER) view has been implemented in connectionist and artificial neural network models...

Background Obsessive-compulsive disorder (OCD) is a psychiatric condition that typically manifests in compulsive urges to perform irrational or excessive avoidance behaviors. A recent account has suggested that compulsivity in OCD might arise from excessive stimulus-response habit formation, rendering behavior insensitive to goal value. We tested i...

Rationale Safety signals providing relief are hypothesised to possess conditioned reinforcing properties, supporting the acquisition of a new response (AnR) as seen with appetitive stimuli. Such responding should also be sensitive to the rate-increasing effects of d-amphetamine and to the anxiolytics 8-OH-DPAT and diazepam. Objectives This study te...

In this chapter, we review studies conducted in humans and other animals (mainly rats) in order to characterise current understanding on experimental extinction, and the possible implications for exposure-based therapies. Extinction learning, rather than producing unlearning (or erasure) of the original S–O association, seems to be best captured as...

The context's role in Pavlovian conditioning depends on the trial spacing during training, with massed trials revealing a function akin to that of discrete stimuli, and spaced trials revealing a modulatory function (Urcelay & Miller, Journal of Experimental Psychology. Animal Behavior Processes, 36, 268-280, 2010). Here we examined the contextual d...

Previous work has demonstrated a profound effect of N-methyl-D: -aspartic acid receptor (NMDAR) antagonism in the infralimbic cortex (IL) to selectively elevate impulsive responding in a rodent reaction time paradigm. However, the mechanism underlying this effect is unclear. This series of experiments investigated the pharmacological basis of this...

Obsessive-compulsive disorder (OCD) is a debilitating condition (prevalence 1-3%), with compulsive checking as the most commonly-reported symptom. Compulsive components of obsessive-compulsive disorder (OCD) are defined as tendencies to perform repetitive, stereotyped acts to prevent perceived negative consequences. We developed a novel test of co...

The most commonly-treated symptom of OCD is compulsive checking: “I have to go and check, but then I’m not convinced about the checking. So I have to go and check that I checked properly.” Compulsive-checking routines may extend to several hours per day, at the expense of normal function, and are linked with anxiety and intolerance of uncertainty....

Obsessive-compulsive disorder (OCD) is a debilitating condition (prevalence 1-3%), with compulsive checking the most commonly-reported symptom. Compulsive checking routines relate to security/accuracy (e.g., checking doors are locked), often for many hours per day at the expense of normal function. We developed a novel test of compulsive checking...

Obsessive-compulsive disorder (OCD) is a debilitating condition (prevalence 1-3%), with compulsive checking the most commonly-reported symptom. Compulsive checking routines relate to security/order/accuracy (e.g., checking doors are locked), often for many hours per day at the expense of normal function. Although existing animal models of compuls...

In this chapter, we consider the relevance of impulsivity as both a psychological construct and endophenotype underlying attention-deficit/hyperactivity disorder (ADHD) and drug addiction. The case for executive dysfunction in ADHD and drug addiction is critically reviewed in the context of dissociable cognitive control processes mediated by the do...

At both empirical and theoretical levels, multiple functional roles of contextual information upon memory performance have been proposed without a clear dissociation of these roles. Some theories have assumed that contexts are functionally similar to cues, whereas other views emphasize the retrieval facilitating properties of contextual information...

We examined trial spacing during extinction following a human contingency learning task. Specifically, we assessed if an expanding retrieval practice schedule (Bjork & Bjork, 1992, 2006), in which the spacing between extinction trials was progressively increased, would result in faster immediate extinction and less recovery from extinction than uni...

Three conditioned lick suppression experiments with rats examined the role of the context in the selection and integration of independently acquired interval relationships. In Experiment 1, rats were exposed to separate conditioned stimuli 1 and 2 (CS1-CS2) pairings with 2 different interval relationships, each in its own distinctive context, X or...

In this review, we address the question, central to cognition, of whether nonhuman animals such as rats are capable of extracting and extending information from a given learning situation to a new learning situation without generalizing through a physical dimension of the stimuli.This capacity underlies abstraction, which is a hallmark of human cog...

Three Pavlovian fear conditioning experiments with rats as subjects explored the effect of extinction in the presence of a concurrent excitor. Our aim was to explore this particular treatment, documented in previous studies to deepen extinction, with novel control groups to shed light on the processes involved in extinction. Relative to subjects ex...

Two conditioned suppression experiments with rats investigated the influence on latent inhibition of compounding a Pavlovian conditioned inhibitor with the target cue during preexposure treatment. Results were compared with those of subjects that received conventional latent inhibition training, no preexposure, or preexposure to the target cue in c...

In the present experiments, we examined the role of within-compound associations in the interaction of the overshadowing procedure with conditioned stimulus (CS) duration, using a conditioned suppression procedure with rats. In Experiment 1, we found that, with elemental reinforced training, conditioned suppression to the target stimulus decreased...

The present experiments addressed a fundamental discrepancy in the Pavlovian conditioning literature concerning responding to a target cue following compound reinforced training with another cue of higher salience. Experiment 1 identified one determinant of whether the target cue will be overshadowed or potentiated by the more salient cue, namely c...

We argue that the propositional and link-based approaches to human contingency learning represent different levels of analysis because propositional reasoning requires a basis, which is plausibly provided by a link-based architecture. Moreover, in their attempt to compare two general classes of models (link-based and propositional), Mitchell et al....

Studies of extinction in classical conditioning situations can reveal techniques that maximize the effectiveness of exposure-based behavior therapies. In three experiments, we investigated the effect of varying the intertrial interval during an extinction treatment in a fear-conditioning preparation with rats as subjects. In Experiment 1, we found...

Weak behavioral control (blocking) occurs when a target stimulus (X) is paired with an outcome in the presence of a well-established signal for the outcome (i.e., a blocking stimulus). Conventional Pavlovian conditioning theories explain this effect by asserting that a discrepancy between expected and experienced outcomes is necessary for learning...

Two experiments using human participants investigated whether a Pavlovian backward inhibitory treatment (nonreinforced trials in phase 1 followed by reinforced trials in phase 2; i.e., AX- followed by A+) produces a stimulus which can pass summation and retardation tests for inhibition. The rationale for conducting these experiments was that previo...

In a Pavlovian fear-conditioning preparation, we investigated the effects of combining Pavlovian and explicitly unpaired inhibition treatments. A summation test for inhibition suggested a strong tendency toward unpaired inhibition when that treatment was administered alone and found robust Pavlovian inhibition when that treatment was administered a...

In 3 experiments using rats as subjects, the authors varied trial spacing to investigate the conditions under which Pavlovian and differential inhibition are observed. Experiment 1 compared Pavlovian and differential inhibition with spaced training trials. Spaced trials resulted in only the Pavlovian inhibitor passing both summation and retardation...

Four experiments using rats in a Pavlovian lick-suppression preparation investigated the effects of combining 2 treatments known for their response-decrementing effects, namely, overshadowing and degraded contingency. Contrary to most contemporary learning theories, the extended comparator hypothesis predicts that these 2 treatments will counteract...