Extinction Context Research Articles

The role of context conditioning in the reinstatement of responding to an alcohol-predictive conditioned stimulus

Re-exposure to an unconditioned stimulus (US) can reinstate extinguished conditioned responding elicited by a conditioned stimulus (CS). We tested the hypothesis that the reinstatement of responding to an appetitive CS is driven by an excitatory association formed between the US and the context that the US was ingested in during US re-exposure. Male, Long-Evans rats were acclimated to drinking alcohol (15%, v/v) in the home-cage, then trained to associate an auditory CS with an alcohol-US that was delivered into a fluid port for oral intake. During subsequent extinction sessions, the CS was presented as before, but without alcohol. After extinction, rats were re-exposed to alcohol as in training, but without the CS (alcohol re-exposure). 24 h later at test, the CS was presented as in training, but without alcohol. First, we tested the effect of extinguishing the context-alcohol association, formed during alcohol re-exposure, on reinstatement. Conducting four context extinction sessions across four days (spaced extinction) after the alcohol re-exposure session did not impact reinstatement. However, four context extinction sessions conducted across two days (massed extinction) prevented reinstatement. Next, we conducted alcohol re-exposure in a context that either differed from, or was the same as, the test context. One alcohol re-exposure session in a different context did not affect reinstatement, however, three alcohol re-exposure sessions in a different context significantly reduced reinstatement during the first CS trial. These results partially support the view that a context-US association formed during US re-exposure drives the reinstatement of responding to an appetitive, alcohol-predictive CS.

Outcome-selective reinstatement is predominantly context-independent, and associated with c-Fos activation in the posterior dorsomedial striatum

Research from human and animal studies has found that after responding has been successfully reduced following treatment it can return upon exposure to certain contexts. An individual in recovery from alcohol use disorder, for example, might relapse to drinking upon visiting their favourite bar. However, most of these data have been derived from experiments involving a single (active) response, and the context-dependence of returned responding in situations involving choice between multiple actions and outcomes is less well-understood. We thus investigated how outcome-selective reinstatement – a procedure involving choice between two actions and outcomes – was affected by altering the physical context in rats. In Experiment 1, rats were trained over 6 days to press a left lever for one food outcome (pellets or sucrose) and a right lever for the other outcome. Then, rats received an extinction session in either the same context (A) as lever press training, or in a different context (B). Rats were tested immediately (5 min) after extinction in Context A or B such that there were four groups in total: AAA, ABB, ABA, and AAB. Reinstatement testing consisted of one food outcome being delivered ‘freely’ (i.e. unearned by lever pressing and unsignalled by cues) to the food magazine every 4 min in the following order: Sucrose, Pellet, Pellet, Sucrose. Selective reinstatement was considered intact if pellet delivery increased pressing selectively on the pellet lever, and sucrose delivery selectively increased pressing on the sucrose lever. This result (Reinstated > Nonreinstated) was observed for rats in group AAA and ABB, but not rats in groups ABA and AAB. Experiment 2 was conducted identically, except that rats received two extinction sessions over two days and tested one day later. This time, all groups demonstrated intact outcome-selective reinstatement regardless of context. Analysis of c-Fos expression in several brain regions revealed that only c-Fos expression in the posterior dorsomedial striatum (pDMS) was related to intact reinstatement performance. Overall, these results suggest that outcome-selective reinstatement is predominantly context-independent, and that intact reinstatement is related to neuronal activity in the pDMS.

Using Extinction-Renewal to Circumvent the Memory Strength Boundary Condition in Fear Memory Reconsolidation.

Reconsolidation is a process by which memories are destabilized, updated, and then restabilized. Strong memories are resistant to undergoing reconsolidation. Here, we addressed whether an overtrained fear memory could be made susceptible to reconsolidation by first extinguishing, and then renewing, the memory. Rats were trained with ten tone-footshock pairings, followed by eight days of tone extinction in the training context. The next day, rats were placed into a second context and memory for the tone was renewed/reactivated with a single tone presentation. Immediately following reactivation, rats received an injection of midazolam or vehicle. Rats were then tested for freezing to the tone in a third context. Midazolam had no effect in rats that did not undergo tone extinction, but significantly attenuated freezing to the tone in extinguished rats. Thus, rats that received tone extinction underwent tone memory reconsolidation following its renewal. In a second experiment, we administered the reactivation session and midazolam injections prior to extinction. Midazolam had no effect and rats extinguished at a rate similar to controls. These data suggest that strong emotional memories are capable of updating following weakening of memory expression through extinction.

Higher functional connectivity between prefrontal regions and the dorsal attention network predicts absence of renewal

Renewal describes the recovery of an extinguished response when extinction and recall contexts differ, demonstrating the context-dependency of extinction. The unexpected outcome change during extinction presumably directs attention to the context and promotes renewal. Accordingly, studies show that context processing for renewal is modulated by salience of and attention to context. Besides context-processing hippocampus, renewal involves ventromedial prefrontal cortex, orbitofrontal cortex and inferior frontal gyrus, which mediate response processing. Since showing renewal is a trait-like processing tendency, individuals with and without renewal may differ in resting-state functional connectivity of prefrontal regions with networks mediating attentional and salience processing.We analyzed resting-state functional MRI data from healthy participants (n = 70) of a non-fear-related contextual extinction task particularly suited for investigation of renewal. Participants without renewal exhibited significantly higher functional connectivity between prefrontal regions and bilateral intraparietal sulcus of the dorsal attention network. Functional connectivity between these regions correlated negatively with renewal level. Only in participants with renewal, the renewal level correlated positively with connectivity between left frontal eye field and several prefrontal regions. In contrast, functional connectivity of prefrontal regions with the salience network did not differ between groups.The results deliver first-time evidence for differences in resting-state functional connectivity between participants with and without renewal in non-fear-related extinction. Intraparietal-sulcus-guided top-down attentional control appears more strongly related to prefrontal activity in participants without renewal, and thus may have a role in their default processing mode of focusing on the stimulus and disregarding the context.

Emergence of complex dynamics of choice due to repeated exposures to extinction learning

Extinction learning, the process of ceasing an acquired behavior in response to altered reinforcement contingencies, is not only essential for survival in a changing environment, but also plays a fundamental role in the treatment of pathological behaviors. During therapy and other forms of training involving extinction, subjects are typically exposed to several sessions with a similar structure. The effects of this repeated exposure are not well understood. Here, we studied the behavior of pigeons across several sessions of a discrimination-learning task in context A, extinction in context B, and a return to context A to test the context-dependent return of the learned responses (ABA renewal). By focusing on individual learning curves across animals, we uncovered a session-dependent variability of behavior: (1) during extinction, pigeons preferred the unrewarded alternative choice in one-third of the sessions, predominantly during the first one. (2) In later sessions, abrupt transitions of behavior at the onset of context B emerged, and (3) the renewal effect decayed as sessions progressed. We show that the observed results can be parsimoniously accounted for by a computational model based only on associative learning between stimuli and actions. Our work thus demonstrates the critical importance of studying the trial-by-trial dynamics of learning in individual sessions, and the power of “simple” associative learning processes.

An Analysis of Extinction-Cue Features in the Reduction of Operant Behavior Relapse

The present free operant conditioning experiments with rats examined the impact of manipulating different traits of an extinction-cue on ABA renewal. All rats were trained to perform an operant response (R1) for food in context A, and a different operant response (R2) in context B. Responses were then extinguished, R1 in context B and R2 in context A. In Experiment 1, for half of the rats, both responses were extinguished in the presence of an intermittent light. For the other half, a continuous light was utilized during extinction of R1 and R2. Finally, all rats were tested in both contexts (R1 was tested in the presence of the cue in both contexts, whereas R2 was tested in the presence of the cue only in the extinction context). Returning the rats to the original conditioning context renewed the initial operant behavior. However, we found that both types of extinction-cues (intermittent and continuous) attenuated ABA renewal equally. In addition, both responses underwent extinction in Experiment 2. For half of the subjects, R1 and R2 were extinguished in the presence of a dim light, whereas the other half received a bright light as an extinction-cue for both responses. Once again, we found during testing (R1 was tested in the presence of the cue in both contexts whereas R2 was tested in the presence of the cue only in the extinction context), that both types of extinction-cues reduced ABA renewal in a similar manner. The present results suggest that in some conditions, the features of the extinction cues may not affect their renewal-attenuating properties.

Behavioral and brain mechanisms mediating conditioned flight behavior in rats

Environmental contexts can inform animals of potential threats, though it is currently unknown how context biases the selection of defensive behavior. Here we investigated context-dependent flight responses with a Pavlovian serial-compound stimulus (SCS) paradigm that evokes freeze-to-flight transitions. Similar to previous work in mice, we show that male and female rats display context-dependent flight-like behavior in the SCS paradigm. Flight behavior was dependent on contextual fear insofar as it was only evoked in a shock-associated context and was reduced in the conditioning context after context extinction. Flight behavior was only expressed to white noise regardless of temporal order within the compound. Nonetheless, rats that received unpaired SCS trials did not show flight-like behavior to the SCS, indicating it is associative. Finally, we show that pharmacological inactivation of two brain regions critical to the expression of contextual fear, the central nucleus of the amygdala (CeA) and bed nucleus of the stria terminalis (BNST), attenuates both contextual fear and flight responses. All of these effects were similar in male and female rats. This work demonstrates that contextual fear can summate with cued and innate fear to drive a high fear state and transition from post-encounter to circa-strike defensive modes.

Presynaptic GABAB receptor inhibition sex dependently enhances fear extinction and attenuates fear renewal.

Anxiety and trauma-related disorders are highly prevalent worldwide, andare associated with altered associative fear learning. Despite the effectiveness of exposure therapy, which aims to reduce associative fear responses, relapse rates remain high. This is due, in part, to the context specificity of exposure therapy, which is a form of extinction. Many studies show that fear relapses when mice are tested outside the extinction context, and this is known as fear renewal. Using Pavlovian fear conditioning and extinction, we can study the mechanisms underlying extinction and renewal. The aim of the current experiment was to identify the role of presynaptic GABAB receptors in these two processes. Previous work from our lab showed that genetic deletion or pharmacological inhibition of GABAB(1a) receptors that provide presynaptic inhibition on glutamatergic terminals reduces context specificity and leads to generalization. We therefore hypothesized that inactivation of these presynaptic GABAB receptors could be used to reduce the context specificity associated with fear extinction training and suppress renewal when mice are tested outside of the extinction context. Using CGP 36216, an antagonist specific for presynaptic GABAB receptors, we blocked presynaptic GABAB receptors using intracerebroventricular injections during various time points of extinction learning in male and female mice. Results showed that blocking these receptors pre- and post-extinction training led to enhanced extinction learning in male mice only. We also found that post-extinction infusions of CGP reduced renewal rates in male mice when they were tested outside of the extinction context. In an attempt to localize the function of presynaptic GABAB receptors within regions of the extinction circuit, we infused CGP locally within the basolateral amygdala or dorsal hippocampus. We failed to reduce renewal when CGP was infused directly within these regions, suggesting that presynaptic inhibition within these regions per se may not be necessary for driving context specificity during extinction learning. Together, these results show an important sex-dependent role of presynaptic GABAB receptors in extinction and renewal processes and identify a novel receptor target that may be used to design pharmacotherapies to enhance the effectiveness of exposure therapy.

An evaluation of a renewal-mitigation procedure for inappropriate mealtime behavior.

Renewal, the increase in behavior during extinction following context changes, may be particularly concerning during intervention for feeding disorders because context changes are often necessary for intervention generality and maintenance (Podlesnik et al.,2017). In the current study, we tested for renewal and evaluated a renewal-mitigation procedure when we transferred intervention from a therapist to a caregiver, from clinic to the home, and changed the foods the feeder presented. We used an ABA arrangement to evaluate the generality of the renewal effect with 7 participants who engaged in inappropriate mealtime behavior. Context A was functional reinforcement. Context B was function-based extinction during the control and mitigation conditions and our renewal-mitigation procedure in the mitigation condition. The renewal test was function-based extinction in Context A. We observed renewal of inappropriate mealtime behavior in 4 of 7 participants, and our renewal-mitigation procedure was effective for 4 of 4 participants.

The role of context in persistent avoidance and the predictive value of relief

Fear renewal occurs when the context changes after fear extinction; however, whether avoidance is also influenced by context changes following fear extinction is untested. Forty-two participants performed an avoidance task within a typical fear renewal procedure. During Pavlovian conditioning, two stimuli (CS+) were associated with an aversive electrical stimulus (US), while a third stimulus was not (CS-). During subsequent avoidance learning, clicking a button canceled the delivery of the US during one but not the other CS+. Fear-related levels were then reduced by removing the US and the button in a new context (fear extinction with response prevention [Ext-RP]). Next, persistence of avoidance was tested in the extinction context B (group ABB) or the original conditioning context A (group ABA). We also tested whether ratings of relief pleasantness (based on both the CS- and the avoided CS+) during avoidance and Ext-RP predicted individual levels of persistent avoidance. Results showed that persistent avoidance was higher in conditioning context A than in extinction context B, and was predicted by higher relief pleasantness during avoidance conditioning. We conclude that persistent avoidance poses a threat to the long-term success of Ext-RP, and we propose that interventions aimed at mitigating the influence of context and relief levels might prove beneficial in this regard.

Dorsal hippocampus plays a causal role in context-induced reinstatement of alcohol-seeking in rats

Drug addiction is a chronic mental disorder characterized by frequent relapses. Contextual cues associated with drug use to play a critical causal role in drug-seeking behavior. The hippocampus has been implicated in encoding drug associative memories. Here we examine whether the dorsal hippocampus mediates context-induced reinstatement of alcohol-seeking. Male Long-Evans rats were trained to self-administer alcohol in Context A. Alcohol self-administration was extinguished in a distinct context (Context B). On the test day, animals were re-exposed to the alcohol Context A or the extinction Context B. Next, to assess a causal role for the dorsal hippocampus in context-induced alcohol-seeking, on the test day, we injected cobalt chloride (CoCl2; a nonselective synapse inhibitor) or vehicle into the dorsal hippocampus, and 15 min later, rats were tested by re-exposing them to the drug-associated context. The re-exposure to the alcohol-associated Context A reinstated alcohol seeking and increased Fos-positive cells in the dorsal hippocampus neurons (CA1, CA3, and Dentate Gyrus). Pharmacological inactivation with cobalt chloride of the dorsal hippocampus attenuated the reinstatement of alcohol-seeking. Our data suggest that the dorsal hippocampus may be involved in context-induced alcohol-seeking behavior.

Reduced renewal of conditioned suppression following lesions of the dorsal hippocampus in male rats.

Extinguished responding will renew when the conditioned stimulus occurs outside the extinction context. Although studies of conditioned freezing have consistently demonstrated a role for the hippocampus in renewal, several studies have demonstrated intact renewal of conditioned suppression despite damage to the hippocampus (Frohardt, Guarraci, & Bouton, 2000; Todd, Jiang, DeAngeli, & Bucci, 2017; Wilson, Brooks, & Bouton, 1995). Because these prior studies have examined renewal when testing occurred in the original conditioning context ("Context A"), the present conditioned suppression experiments examined the role of the hippocampus when testing occurred in a context not associated with prior conditioning ("Context C"). In Experiments 1 and 2, conditioning occurred in Context A, and extinction in Context B. Renewal of conditioned suppression was observed when the extinguished conditioned stimulus (CS) was tested in Context C. However, renewal was attenuated in rats with lesions of the dorsal hippocampus (DH). Summation testing failed to detect conditioned inhibition in the extinction context, suggesting instead that the context acquired negative occasion-setting properties. Attenuated renewal was not due to an inability of DH lesioned rats to discriminate contexts (Experiment 3). These experiments thus demonstrate a role for the DH in renewal of conditioned suppression when testing occurs in a neutral context. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

Extinction contexts fail to transfer control: Implications for conditioned inhibition and occasion-setting accounts of renewal.

The renewal effect is often explained as a side effect of the extinction context acting as a negative occasion setter. Four experiments tested whether extinction contexts show the selective-transfer property of occasion setters. Experiments 1-3 used a predictive judgment task where participants rated the probability of certain foods (cues) producing gastric malaise (outcomes) in different restaurants (contexts). Experiment 4 used a behavioral suppression task where sensor lights (cues) served as signals to suppress firing responses in certain galaxies (contexts). All 4 (Experiments 1-4) addressed whether a potentially negative occasion-setting context transferred its modulatory power to an extinguished (presumably occasion set) target in the test phase of an ABC renewal design. Experiments 2-4 further assessed the possibility that the extinction context acts as a conditioned inhibitor by testing a simple excitor on a context where extinction occurred. Neither selective (occasion-setting) nor nonselective transfer (conditioned inhibition) was demonstrated. Implications for theories of renewal and occasion-setting are discussed. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

Context and renewal of habits and goal-directed actions after extinction.

Instrumental behaviors that are goal-directed actions after moderate amounts of training can become habits after more extended training. Little research has asked how actions and habits are affected by retroactive interference treatments like extinction. The present experiments begin to fill this gap in the literature. In Experiments 1a and 1b, lever pressing in rats was minimally trained (1a) or extensively trained (1b) in one context (Context A), extinguished in a second context (Context B), and then tested in the acquisition context (Context A). Exposure to both contexts was equated and controlled throughout, and the status of the behavior as action or habit was determined by reinforcer devaluation methods (taste aversion conditioning). Results confirmed that action (1a) and habit (1b) renewed with action or habit status, respectively, when they were returned to Context A. Experiments 2a and 2b then similarly tested action and habit after extinction in an ABC renewal paradigm. Here, lever pressing that was trained in Context A and extinguished in Context B renewed as action in Context C regardless of whether it had been an action or habit before extinction. The apparent conversion of habit to action during renewal testing in Context C was consistent with other results suggesting that habits converted to action when the context was changed at the start of extinction. Together, the results suggest that extinction in a second context inhibits instrumental behaviors trained as either actions or habits in a context-specific manner. They also expand on prior findings suggesting that actions transfer across contexts, and that habits do not. A change of context may be sufficient to convert a habit to goal-directed action. (PsycInfo Database Record (c) 2020 APA, all rights reserved).

A New 3-Day Standardized Eyeblink Conditioning Protocol to Assess Extinction Learning From Infancy to Adulthood.

Associative learning can be observed from the neonatal period onward, providing opportunities to examine changes in basic learning and memory abilities. One method that is suitable to study associative learning is classical eyeblink conditioning (EBC) which is dependent on the cerebellum. Extinction learning can be systematically investigated in this paradigm by varying the context during learning and extinction. Because of methodological difficulties and ethical challenges, no studies have compared extinction learning using EBC across human development. Our goal was to test feasibility of a 3-day delay EBC paradigm that can be used from infancy to adulthood. Acceptance/safety was tested especially for infancy by investigating attrition rates and parental report on infant wellbeing. On a paradigm side, we tested if the paradigm leads to successful acquisition and extinction. An air puff served as unconditional stimulus (US) and a tone as conditional stimulus (CS). On day 1 during acquisition, participants received 36 US–CS pairings in context A. On day 2, participants received 12 acquisition trials in context A to consolidate association learning, followed by 48 extinction trials (tone alone presentations) in context B. Renewal was assessed on day 3 and incorporated 12 CS alone trials presented in both the acquisition context and the extinction context. Eyeblink responses were videotaped and coded offline. The protocol was tested with 12–36-months-old infants (N = 72), adolescents (N = 8), and adults (N = 8). Concerning the acceptance/safety side, attrition ranged from 21 to 58% in infant samples due to the complex preparation of the children for the paradigm. However, attrition is equal to or lower than other infant learning paradigms. Parents of infant samples were very interested in the paradigm and reported low levels of infant stress, exhaustion, and negative feelings during the sessions. Data quality was very high, and no participant had to be excluded because of insufficient data. Concerning the paradigm side, participants showed successful acquisition and extinction as a group. The procedure is ethically sound, feasible, tolerated by many infants, and acceptable among parents. The data show successful acquisition and extinction rates, making the paradigm a valuable tool for investigating developmental changes in extinction learning over the lifespan.

Adding or removing context components equally disrupts extinction in human predictive learning

In two human predictive learning experiments, we investigated the impact of adding or removing context components on extinction performance toward a cue. In each experiment, participants initially received repeated pairings of a cue and an outcome in a context composed of two distinctive components (context AB). Initial training was followed by a series of trials in which the cue was no longer followed by the outcome. This extinction treatment was conducted in the presence of a different pair of distinctive context components (context CD). During a final test, we observed that changing the extinction context CD disrupted extinction performance toward the cue regardless of whether the context was changed by adding or removing context components. We discuss implications of our results for theories of associative learning.

Contextual reinstatement promotes extinction generalization in healthy adults but not PTSD

For episodic memories, reinstating the mental context of a past experience improves retrieval of memories formed during that experience. Does context reinstatement serve a similar role for implicit, associative memories such as fear and extinction? Here, we used a fear extinction paradigm to investigate whether the retrieval of extinction (safety) memories is associated with reactivation of the mental context from extinction memory formation. In a two-day Pavlovian conditioning, extinction, and renewal protocol, we collected functional MRI data while healthy adults and adults with PTSD symptoms learned that conditioned stimuli (CSs) signaled threat through association with an electrical shock. Following acquisition, conceptually related exemplars from the CS category no longer signaled threat (i.e., extinction). Critically, during extinction only, task-irrelevant stimuli were presented between each CS trial to serve as “context tags” for subsequent identification of the possible reinstatement of this extinction context during a test of fear renewal the next day. We found that healthy adults exhibited extinction context reinstatement, as measured via multivariate pattern analysis of fMRI data, in the medial temporal lobe that related to behavioral performance, such that greater reinstatement predicted CSs being rated as safe instead of threatening. Moreover, context reinstatement positively correlated with univariate activity in the ventromedial prefrontal cortex and hippocampus, regions which are thought to be important for extinction learning. These relationships were not observed in the PTSD symptom group. These findings provide new evidence of a contextual reinstatement mechanism that helps resolve competition between the retrieval of opposing associative memories of threat and safety in the healthy adult brain that is dysregulated in PTSD.

Stress inoculation in mice induces global resilience

Each year, more than half a billion people in the world are affected by stress-related health disorders. Consequently, there is an urgent need for new insights to guide interventions designed to increase stress resilience. Studies of humans and various animals have uncovered the process of stress inoculation, in which exposure to mild stressors enhances subsequent stress resilience. Here, we investigate whether stress inoculation-induced resilience in mice consistently occurs across a multiplicity of different stress contexts (tests). C57BL/6 J adult male mice were randomised either to stress inoculation training (n = 36) or to a non-inoculated, but handled control condition (n = 36). Thereafter, indications of coping and resilience were assessed during (i) acute social defeat in a context similar to that used for stress inoculation training, and (ii) fear conditioning and learned extinction in a novel context. Stress inoculation effects were also assessed during (iii) tail-suspension and (iv) open-field tests that each represent milder stressors. Stress-inoculated mice showed more active defence behaviour during acute social defeat, higher sociability before and after defeat, and greater indications of learned extinction of conditioned fear compared to non-inoculated control mice. Stress-inoculated mice also responded with diminished tail-suspension immobility and open-field defecation. Results suggest that stress inoculation protects against various stressors that differ in quality and relative intensity. Stress inoculation research in mice may serve as the basis for mechanistic studies of global resilience in humans.

Discerning the diets of sweep-feeding eurypterids: assessing the importance of prey size to survivorship across the Late Devonian mass extinction in a phylogenetic context

AbstractEurypterids are generally considered to comprise a mixture of active nektonic to nektobenthic predators and benthic scavenger-predators exhibiting a mode of life similar to modern horseshoe crabs. However, two groups of benthic stylonurine eurypterids, the Stylonuroidea and Mycteropoidea, independently evolved modifications to the armature of their anterior appendages that have been considered adaptations toward a sweep-feeding life habit, and it has been suggested the evolution toward sweep-feeding may have permitted stylonurines to capture smaller prey species and may have been critical for the survival of mycteropoids during the Late Devonian mass extinction. There is a linear correlation between the average spacing of feeding structures and prey sizes among extant suspension feeders. Here, we extrapolate this relationship to sweep-feeding eurypterids in order to estimate the range of prey sizes that they could capture and examine prey size in a phylogenetic context to determine what role prey size played in determining survivorship during the Late Devonian. The mycteropoidCyrtoctenuswas the most specialized sweep-feeder, with comblike appendage armature capable of capturing mesoplankton out of suspension, while the majority of stylonurines possess armature corresponding to a prey size range of 1.6–52 mm, suggesting they were suited for capturing small benthic macroinvertebrates such as crustaceans, mollusks, and wormlike organisms. There is no clear phylogenetic signal to prey size distribution and no evolutionary trend toward decreasing prey sizes among Stylonurina. Rather than prey size, species survivorship during the Late Devonian was likely mediated by geographic distribution and ability to capitalize on the expanding freshwater benthos.

Renewal of conditioned tentacle lowering by circadian contextual cues in snails Cornu aspersum

Previous experiments using tentacle lowering conditioning in terrestrial snails Cornu aspersum have shown extinction and recovery of the conditioned response (CR) as a consequence of both inserting a delay between the extinction and test (spontaneous recovery) and of re-exposing the animal to the unconditioned stimulus after extinction (reinstatement). Two experiments that examined recovery of the CR due to a change in context (renewal effect) were carried out to continue this line of research. In Experiment 1, subjects received conditioning with an odour (CS) followed by extinction in the presence of another odour (CS + C), before being exposed to the original one (CS). In Experiment 2, conditioning and extinction of an odour CS took place in the presence of different circadian contextual cues (hour of the day and presence of light). The results showed that a return to the original context of conditioned training, after the extinction in a different context, either defined by an odour (Experiment 1) or by circadian cues (Experiment 2), produce a recovery of the CR compared to suitable control groups. These results can be interpreted as an instance of ABA renewal effect and they provide information about psychological mechanisms involved in extinction processes.