EXPERIMENTS CONCERNING THE TRANSFER OFA LEARNED RESPONSE AND ACTIVITY LEVEL BY BRAIN HOMOGENATES

L agerspetz , K. M. J., R aitis , P., T irri , R. & L agerspetz , K. Y. H. Experiments concerning the transfer of a learned response and activity level by brain homogenates. Scand.J. Psychol ., 1968, 9, 225–229,— The learning of a conditioned avoidance response caused an increase in the open-field activity of male albino mice. Pseudo-conditioning had an opposite effect. The transfer of brain homogenate from the trained animals did not facilitate the acquisition of the avoidance response by the recipient animals. Differences in the open-field activity were not transmitted with the brain homogenates. However, the constancy of the open-field activity scores was decreased by the homogenate injection, and also by severe electric shocks.     

Negative reinforcement as shock-frequency reduction

Is a conditioned aversive stimulus necessary in avoidance conditioning? Or is a reduction in the rate of aversive stimulation alone sufficient to generate and maintain an avoidance response? Rats were subjected to an avoidance procedure in which shocks occurred randomly in time, but a response could reduce the overall rate of shock. Fifteen acquisition curves, obtained from 16 animals, showed both immediate and delayed, rapid and gradual increases in response rate; there was no representative acquisition curve. Response rates were directly related to the amount by which the response reduced shock frequency. In extinction, when shock rates were not affected by responding, the response total was inversely related to the amount by which the response had reduced shock frequency during prior conditioning, with as many as 20,000 extinction responses when the shock frequency reduction had been relatively small. Responding on this procedure shows that avoidance conditioning can occur without benefit of either classical exteroceptive stimuli or covert stimuli inferred from the temporal constancies of a procedure. It also shows that reduction in shock rate is alone sufficient to maintain avoidance.     

Avoidance extinction in goldfish

Independent groups of goldfish trained to avoid shock in a shuttlebox situation were presented with several extinction procedures in which the relationships between the conditioned stimulus and shock were altered and/or response contingencies removed. Random shock presentations, equivalent to the number of shocks received during avoidance acquisition, resulted in response decrements similar to those obtained when the conditioned stimulus was presented alone. Pairing the conditioned stimulus with shock on every trial, however, served to maintain response levels. When response-contingent punishment was superimposed upon these Pavlovian pairings, performance was facilitated slightly although punishment alone resulted in somewhat faster response reduction than that produced by exposure only to the conditioned stimulus. Extinction of avoidance responding produced by exposure to the conditioned stimulus alone was dependent on the total duration of exposure and independent of both number of stimulus onsets and response prevention. These experiments demonstrated that, in general, the procedures used to reduce avoidance responding in rats were equally effective for goldfish, with one exception: the introduction of a Pavlovian contingency following avoidance acquisition, making the previously avoidable shock unavoidable, maintained response probabilities near previously established levels.     

Limbic networks and associative learning: II. Entorhinal contributions to spontaneous alternation,passive avoidance,and taste aversion learning

The hippocampus plays an important role in learning and memory, but the precise nature of that involvement remains uncertain. Transection of the perforant path, a primary input pathway to the hippocampus, has been shown to produce changes in reaction to novelty and acquisition of active avoidance; the nature and magnitude of these changes vary with lateral or medial perforant path damage. In a series of experiments on adult rats, the role of these pathways in spontaneous alternation, exploration, acquisition and extinction of conditioned responses, passive avoidance, and conditioned taste aversion was investigated. Lateral transection reduced exploration while medial transection facilitated acquisition of an active avoidance response; no effects were observed on any other measure. Results are discussed in terms of what perforant path damage might reveal regarding the interactions of the hippocampus with other brain regions.     

Limbic networks and associative learning: II. Entorhinal contributions to spontaneous alternation, passive avoidance, and taste aversion learning

The hippocampus plays an important role in learning and memory, but the precise nature of that involvement remains uncertain. Transection of the perforant path, a primary input pathway to the hippocampus, has been shown to produce changes in reaction to novelty and acquisition of active avoidance; the nature and magnitude of these changes vary with lateral or medial perforant path damage. In a series of experiments on adult rats, the role of these pathways in spontaneous alternation, exploration, acquisition and extinction of conditioned responses, passive avoidance, and conditioned taste aversion was investigated. Lateral transection reduced exploration while medial transection facilitated acquisition of an active avoidance response; no effects were observed on any other measure. Results are discussed in terms of what perforant path damage might reveal regarding the interactions of the hippocampus with other brain regions.     

MODIFICATION OF SHUTTLE BOX TO IMPROVE RATE OF AVOIDANCE LEARNING IN RATS

W allgren , H. & S avolainen , S. Modification of shuttle-box to improve rate of avoidance learning in rats. Scand. J. Psychol ., 1962, 3 , 78–80.—Acquisition by rats of a conditioned avoidance response seems to be retarded in an ordinary shuttle-box by the necessity to re-enter the compartment where shocks have previously been received. To avoid this, a four-compartment box was introduced in which the animals can proceed in one direction. Rats were given either continuous trials until nine avoidance responses were made during 10 consecutive trials, or 30 trials per day until 27 correct responses were made during one session. With both methods of training, the rate of learning was approximately 40 per cent more rapid in the four-compartment box than in the ordinary shuttle-box.     

Effectiveness of the counterconditioning procedure in eliminating avoidance behavior

Data were obtained on the effectiveness of three forms of counterconditioning—each differing in the degree to which competing responses were introduced. Counterconditioning was also compared with an extinction procedure in eliminating an instrumental avoidance response. Only the counterconditioning procedure which evoked and reinforced another conditioned response during extinction of the avoidance response, was significantly more effective than a traditional extinction procedure in achieving a decrement in avoidance responding and this was only on the first session on which such procedures were in effect. When the opportunity to perform the competing response was removed, only Ss in this group evidenced a significant increase in avoidance responding.

All procedures were equally effective over sessions in eliminating avoidance behavior. A delay in effectiveness occurred to the degree to which a procedure introduced a competing response. It was in terms of the immediacy of response reduction and achievement of a very low rate of responding with very few responses being performed, that the counterconditioning procedure was superior to the extinction procedure.

The implication of these results for behavior theory and therapy were elaborated.     

Pavlovian processes and response competition as determinants of avoidance response-prevention effects

Two experiments investigated the facilitation of avoidance extinction by exposure to lengthy (5-sec) shock during avoidance response prevention. In Experiment 1, animals exposed to light only or to light-shock pairings during response prevention showed equal facilitation of extinction relative to shock-only animals or to animals receiving no response prevention. Preshock rearing, directly antagonistic to the avoidance response, developed for shocked animals during response prevention and persisted during extinction for light-shock animals. Immediately before extinction, half of each group was permitted a single escape from a light-shock compound by means of the response previously required for avoidance. The only effect was upon the extinction performance of light-shock animals. Rearing was eliminated and extinction responding increased to a level far above that for any of the other animals. Experiment 2 demonstrated that the shock-only treatment affected the extinction performance and rearing of nonescape and escape animals in a manner entirely equivalent to the effects of the light-shock treatment of Experiment 1, provided stimulus conditions (light absent) were the same for all experimental phases. Thus, lengthy shock during avoidance response prevention simultaneously leads to the acquisition of competing behavior and enhances control by a warning signal or contextual stimuli over the avoidance response. Implications for the CS-only response-prevention treatment and the transfer of aversive control are discussed.     

Acquisition and extinction of signaled avoidance as a function of intermittent reinforcement

Signaled, shuttle-box avoidance responding in female rats of the Fischer₃₄₄ strain was examined as a function of four separate contingencies of intermittent reinforcement. In Experiment 1, when avoidance responses during acquisition were reinforced 25% of the time with prompt CS termination, animals responded equally often during acquisition and significantly more often during extinction than animals who received such reinforcement on a 100% schedule. Similar results were found under a trace procedure in Experiment 2 when avoidance responses were reinforced 25% of the time with informational feedback stimuli. In contrast, during Experiment 3, when animals were shocked on only 25% of the trials on which they failed to respond, the level of avoidance responding during both acquisition and extinction was significantly less than it was when animals were shocked on a 100% schedule. Comparable results were found in Experiment 4 when avoidance responses during acquisition averted shock on only 25% of the trials. Thus, intermittent reinforcement contingencies involving response-contingent feedback stimuli and shock have differential effects on avoidance responding during both acquisition and extinction trials under the signaled avoidance procedure.     

Role of pituitary-adrenocortical system in mediating avoidance behavior of rats with septal lesions.

Three experiments examined the hypothesis that the effects of septal lesions and systemic injections of scopolamine on avoidance acquisition could be attributed to the effects of either of these treatments on adrenocorticotropic hormone (ACTH) secretion. Septal lesions and scopolamine facilitated 2-way conditioned avoidance response acquisition, and the lesions retarded passive avoidance acquisition. However, neither the injections of dexamethasone, a synthetic glucocorticoid which inhibited ACTH secretion as did septal lesions, nor injections of ACTH which mimicked the facilitatory effects of scopolamine on basal ACTH secretion, affected avoidance in these paradigms. Thus, the main hypothesis was not supported. The finding that scopolamine did not affect passive avoidance indicates that a cholinergic system may not be involved in mediating the suppressive effects of punishment.     

Conscious knowledge of CS-UCS contingency information affects extinction retrieval of conditioned disgust responses: Findings from an online de novo disgust conditioning task

ObjectiveThe present study aimed to establish and develop an online de novo conditioning paradigm for the measurement of conditioned disgust responses. We further explored the effects of explicit instructions about the CS-UCS contingency on extinction learning and retrieval of conditioned disgust responses.MethodThe study included a sample of 115 healthy participants. Geometric figures served as conditioned stimuli (CS) and disgust-evoking pictures as unconditioned stimuli (UCS). During disgust conditioning, the CS+ was paired with the UCS (66% reinforcement) and the CS- remained unpaired; during extinction and retrieval, no UCS was presented. Half of the participants (n = 54) received instructions prior to the disgust extinction stating that the UCS will not be presented anymore. 1-2 days or 7-8 days later participants performed a retrieval test. CS-UCS contingency, disgust and valence ratings were used as dependent measures.ResultsSuccessful acquisition of conditioned disgust response was observed on the level of CS-UCS contingency, disgust and valence ratings. While some decline in valence and disgust ratings during the extinction stage was observed, contingency instructions did not significantly affect extinction performance. Retrieval one week later revealed that contingency instructions increased the discrimination of the CSs.ConclusionsExtinction of conditioned disgust responses is not affected by explicit knowledge of the CS-UCS contingencies. However, contingency instructions prior to extinction seem to have a detrimental effect on long-term extinction retrieval.     

Brain asymmetry and human electrodermal conditioning

Two experiments are reviewed that demonstrate effects of brain laterality on human classical conditioning. Pictures of facial emotional expressions were used as conditioned stimuli (CSs) together with shock as unconditioned stimlus (UCS). Bilateral electrodermal responses were recorded as dependent measures. In the first experiment, one group was conditioned to an angry face, and one group to a happy face. During extinction, the face-CSs were presented to the right hemisphere on half of the trials and to the left hemisphere on the other half of the trials. Results showed that the right hemisphere was superior in showing persisting effects of learning, and especially to the angry CS+. In the second experiment, lateralized presentations of the angry and happy faces were made during acquisition, with foveal presentations during extinction. Once again, the angry face elicited greater skin conductance responses (SCRs) during extinction in the group that had this stimulus presented to the right hemisphere during acquisition. It is concluded that emotional conditioning is differentially regulated by the two hemispheres of the brain.     

Selective and protracted effect of nifedipine on fear memory extinction correlates with induced stress response

Memory extinction, defined as a decrease of a conditioned response as a function of a non-reinforced conditioned stimulus presentation, has high biological and clinical relevance. Extinction is not a passive reversing or erasing of the plasticity associated with acquisition, but a novel, active learning process. Nifedipine blocks L-type voltage gated calcium channels (LVGCC) and has been shown previously to selectively interfere with the extinction, but not the acquisition, of fear memory. We studied here the effect of retrograde and anterograde shifts of nifedipine application, with respect to an extinction training, on the extinction of fear conditioning. Subcutaneous injection of 30 mg/kg nifedipine, at least up to 4 h before the extinction session, significantly impaired extinction, as did intraperitoneal injection of 15 mg/kg nifedipine, at least up to 2 h before extinction training. However, the injection of nifedipine also induced a strong and protracted stress response. The pharmacokinetics of nifedipine suggest that it was mainly this stress response that triggered the specific inhibition of extinction, not the blockade of LVGCC in the brain. Our results support recent findings that stress selectively interferes with the extinction, but not the acquisition, of fear memory. They also indicate that a pharmacological approach is not sufficient to study the role of brain LVGCC in learning and memory. Further research using specific genetically modified animals is necessary to delineate the role of LVGCC in fear memory extinction.     

Safety behavior can hamper the extinction of fear of movement-related pain: An experimental investigation in healthy participants

Excessive fear of movement-related pain (FMRP), and its associated avoidance behavior, is considered a major risk factor for disability in chronic musculoskeletal pain. The current study aimed to investigate whether engaging in safety behavior, conceptualized as an avoidance response, hampers the extinction of FMRP. In a differential conditioning paradigm, we used joystick movements as conditioned stimuli (CSs) and a painful electrocutaneous stimulus as the unconditioned stimulus (US). In the Safety group, participants received the opportunity to avoid the pain-US by pressing a safety button during the extinction phase, whereas in the Control group, this option was not included. In a subsequent test phase, this safety button was no longer available. In two experiments, results demonstrate successful acquisition and extinction. Retrospective FMRP ratings in both experiments revealed a return of fear of pain in the test phase in the Safety group, but not in the Control group. In Experiment 1, mean eyeblink startle reflex amplitudes partly corroborated the self-report findings on fear of pain. The present results suggest that performing safety behavior during cognitive-behavioral interventions, i.e., exposure, might increase the risk of a return of FMRP.     

Effects of naloxone on acquisition and extinction of jump-up avoidance behavior in rats

Two experiments were carried out to assess the effects of the opioid antagonist, naloxone, on the acquisition and extinction of shock avoidance by rats in the jump-up apparatus. In Experiment 1 naloxone pretreatment facilitated acquisition but had no effect on extinction of avoidance behavior. In the second experiment the effect of naloxone on acquisition was replicated and in addition, it was shown that naloxone enhanced freezing when a response prevention or flooding procedure was introduced. Again naloxone failed to alter the course of extinction, nor did it interact with the effects of flooding which, by itself, facilitated extinction. The results suggest that naloxone's effects are limited to increasing the functional intensity of the US, and provide further support for the dissociation between extinction of avoidance behavior and other indices of fear.     

Extinction of conditioned sexual responses in male Japanese quail (Coturnix japonica): role of species-typical cues

The authors examined how a conditioned stimulus (CS) that included species-typical cues affected the acquisition and extinction of conditioned sexual responses in male quail (Coturnix japonica). Some subjects were conditioned with a CS that supported sexual responses and included a taxidermic head of a female quail. Others were conditioned with a similar CS that lacked species-typical cues. Pairing the CSs with access to live females increased CS-directed behavior, with the head CS eliciting significantly more responding than the no-head CS. Responding to the head CS persisted during the 42-day, 126-trial extinction phase; responses to the no-head CS extinguished. Responding declined when the cues were removed or the subjects were sexually satiated. Possible functions and mechanisms of these effects are discussed.     

Effects of UCS intensity and postpeak acquisition trials on classical conditioning of the SCR

The present study examined the effects of UCS intensity and number of postpeak acquisition trials on classical conditioning and extinction of the SCR. A 2 × 3 design was employed in which subjects received either a 1, 2, or 4 mA shock UCS and either two or 16 acquisition trials beyond the peak CR. While conditioning was demonstrated during acquisition, there was no relationship between strength of conditioning and intensity of UCS. The phenomenon of stronger resistance to extinction following fewer acquisition trials (e.g., two past the peak CR) than with many (e.g., 16 past the peak CR) was demonstrated only for the groups that were conditioned with the 4 mA UCS. Resistance to extinction varied positively with UCS intensity, but only for the subjects who received two postpeak acquisition trials. Sixteen trials beyond the peak CR resulted in the UCS intensity having little or no effect on resistance to extinction.     

Deficits in acquisition and extinction of conditioned responses in mGluR7 knockout mice

Metabotropic glutamate receptor 7 (mGluR7) is expressed in brain regions implicated in emotional learning and working memory, and previous behavioral experiments indicated contributions of mGluR7 to various complex behaviors. In the present study, we investigated the specific effects of mGluR7 deletion on a variety of conditioning paradigms that model crucial neurocognitive and psychopathological behavioral phenomena. Null-mutant mGluR7^−/− mice displayed defects during scheduled appetitive conditioning, acquisition and extinction of appetitive odor conditioning, extinction of response suppression-based conditioned emotional responding (CER), acquisition of discriminative CER, and contextual fear conditioning. mGluR7^−/− animals were slower to acquire the association between a conditioned stimulus and a positive or negative reinforcer, but eventually reached similar performance levels to their wildtype littermates. Notably, extinction learning of conditioned responses was slower in mGluR7^−/− compared to wildtype animals. The observed delays in the acquisition of complicated stimulus associations across conditioning procedures may suggest a critical role for mGluR7 in neurocognitive functions and psychopathology.     

Excitatory conditioning of individual Drosophila melanogaster

Testing individual animals from a heterogenic population of Drosophila melanogaster, we demonstrate conditioning of the proboscis extension reflex. The presentation of paired (conditioning) stimuli produced (a) an increase in the average number of conditioned responses over trials, (b) measured differences in performance levels among individual subjects, and (c) a sex difference, with more males conditioned than females, and those that did did so more quickly. The presentation of unpaired (control) stimuli produced significantly lower average levels of acquisition responding and a change in the distribution of individual response patterns. Neither central excitatory state nor sensitization induced by the conditioned or unconditioned stimuli directly affected the conditioned response, whereas unconditioned stimulus preexposure adversely affected performance levels. Presenting the unpaired (extinction) stimuli after conditioning produced less of a decline in responding than did an extinction procedure with removal of the unconditioned stimulus. With the ability to identify individual differences in acquisition and extinction patterns, and given the relatively large number of individuals that can be tested simultaneously on the automated stimulation apparatus, it is now possible to make precise behavioral measurements on samples large enough for the behavior-genetic analysis of D. melanogaster with conditioning as the phenotype.     

Pavlovian phenomena in conditioned acceleration—Stimulus summation

Four naive male rats were separately fear-conditioned to two stimuli (light and sound) by pairing these stimuli with shocks. During Sidman avoidance these unreinforced stimuli elicited increases in response rates (conditioned acceleration). In a multiple schedule with light-shock and sound-shock reinforcements in the first component, the rats were tested for summation effects in the second (avoidance) component by presenting the light twice, the sound twice and the compound twice. They were then extinguished by eliminating the shocks in the first component. The effects of compounding were evidenced by: (1) a sizeable and reliable amplitude increase to the compound stimulus during conditioning; (2) a smaller and less consistent decrease in latencies during conditioning in all four subjects; and (3) a greater resistance to extinction for the compound stimulus as measured by mean amplitudes for all four subjects and on six of seven extinction sessions. Latency data during extinction were inconclusive. An additional unexpected finding was what appeared to be a two-factor secondary extinction of the avoidance itself as a result of Pavlovian extinction. The responding in the first four minutes of avoidance was inhibited to such an extent that the number of shocks received during this period increased 700 per cent from the last three conditioning sessions to the first three extinction sessions.