Some types of conditioned inhibitors carry collateral excitatory associations

Two experiments using summation tests in conditioned suppression with rats examined whether conditioned inhibitors generated by five different conditioning procedures have an associative structure which includes collateral excitatory associations. The existence of collateral conditioned excitation was inferred from an increase in the amount of manifest conditioned inhibition after a conditioned inhibitory stimulus (CS-) extinction treatment. The five CS-s evaluated were differential, explicitly unpaired, conditional, backward, and trace. With abbreviated conditioning (Experiment 1), the differential and explicitly unpaired CS-s exhibited conditioned inhibitory properties; the conditional, backward, and trace CS-s did not. Repeated extinction presentations of the CS- in the absence of the unconditioned stimulus unmasked conditioned inhibition to the conditional, backward, and trace CS-s revealing moderate degrees of conditioned inhibition for all five CS-s. After more extensive conditioning (Experiment 2), the explicitly unpaired and conditional CS-s were strongly inhibitory and the differential and trace CS-s were moderately inhibitory. The backward CS- was not inhibitory. Repeated presentations of the CS- in extinction unmasked conditioned inhibition to trace and backward CS-s. All five CS-s were now nearly equally inhibitory. That the measured inhibitory power of backward, trace, and conditional (after few trials) CS-s can be modulated by a CS- extinction treatment suggests that they have similar associative structures and carry, in addition to inhibitory associations, collateral excitatory associations that mask the expression of conditioned inhibition.     

Stimulus generalization of excitation and inhibition

Five sign-tracking experiments with pigeons investigated the breadth of stimulus generalization of excitatory and inhibitory conditioning. Using a compound stimulus test procedure, these experiments found evidence for a narrower generalization of associative strength in excitation than in inhibition. They also found a narrower associative gradient with a more strongly conditioned excitatory stimulus, although this did not seem to account for the difference between exciters and inhibitors. These results confirm earlier findings comparing generalization after acquisition and extinction and raise various theoretical issues about generalization.     

Stimulus generalization of excitation and inhibition

Five sign-tracking experiments with pigeons investigated the breadth of stimulus generalization of excitatory and inhibitory conditioning. Using a compound stimulus test procedure, these experiments found evidence for a narrower generalization of associative strength in excitation than in inhibition. They also found a narrower associative gradient with a more strongly conditioned excitatory stimulus, although this did not seem to account for the difference between exciters and inhibitors. These results confirm earlier findings comparing generalization after acquisition and extinction and raise various theoretical issues about generalization.     

Neuronal manifestation of two-way connections in conditioning

Neuronal characteristics of two-way conditional connections were studied in chronic experiments with alert cats during different types of alimentary conditional reflexes. Single unit and multi-unit activities were chronically recorded in the visual and motor cortex and the lateral hypothalamic nucleus. Cross-correlational analysis of neuronal impulse trains was used to characterize the organization of neuron groups in brain structures involved in the formation of a conditional reflex. The experimental data showed an increase in the number of neuronal two-way connections after learning in all three investigated coordinations: visual-motor, visual-hypothalamic, and motor-hypothalamic. With visual-motor interneuronal coordination, the strengthening of two-way connections was due to the enhancement of connections from the motor to the visual neuron (backward connections); with visual- and motor-hypothalamic coordination, the number of interneuronal connections was equal in both directions. In all investigated coordinations, the analysis of the temporal parameters of the interactions between the neurons of different groups showed a dependence of the conditioning procedure on delays of up to 30 ms and a dependence of the motivational state on the interval range of 90–100 ms. The polyfunctional and polycomponent character of two-way neuronal connections in conditioning can be inferred from these data. Evaluation of the activity of neurons of different brain structures suggests the specific organization of intracortical and cortical-subcortical integrity in learning.     

Extinguishing conditioned inhibition in flavour-aversion learning: Effects of repeated testing and extinction of the excitatory element

Three conditioned inhibition experiments using an A+/AX- design are reported in which lithium-mediated excitatory conditioning occurred to distinctive environmental stimuli and inhibitory conditioning to a vinegar flavour. Increased vinegar preferences were observed (i.e., conditioned inhibition) in each experiment, and these preferences extinguished with repeated testing as well as following extinction of the excitatory element. Vinegar preferences could be reinstated through reconditioning of the extinguished excitatory stimulus. These experiments speak to the status of inhibitory responding as a “slave” process to conditioned excitation.     

An inhibition effect in the temporal constrains of attentional selection: the Backward Blink

The study aims at investigating the dynamics of visual information processing during sequential control of attentional selection. By combining two different traditional paradigms we show for the first time that during rapid selection of visual targets, encoding of a new target stimulus is facilitated by suppression of the preceding relevant stimulus category (Backward Blink; BB). Such inhibition would operate to reduce interference from previously stored information, in order to facilitate the instantiation of a new attentional episode. Results suggest that the same underlying inhibitory mechanism might contribute to two different attentional effects, specifically, the switch cost and the Attentional Blink, suggesting a general inhibitory mechanism of attentional control, with broad implications for understanding how the brain perceives any task-relevant stimulus.     

Computer simulation of some of the motor circuits underlying Parkinson’s disease

Many researchers have published functional models of brain structures involved in Parkinson’s disease. These models include the primary structures of the basal ganglia and related cortical areas, and have hypothesized normal and dysfunctional states. Simple computer simulation of the hypothesized excitatory and inhibitory interactions among brain structures can be used to evaluate these models. A computer simulation can be programmed to simulate normal function, disease function, and treatment effects described in published models.     

Inhibitory stimulus control following errorless discrimination learning

Three generalization procedures were used to investigate inhibitory stimulus control following discrimination learning with few errors. Three groups of pigeons acquired a discrimination between a green stimulus (the positive stimulus) and a vertical or horizontal line (the negative stimulus) through differential autoshaping followed by multiple schedule presentation of the two stimuli with gradually increasing stimulus durations. Genereralization testing was along a line-tilt continuum. For one group, the test involved a resistance-to-reinforcement procedure in which responses to all line tilts were reinforced on a variable-interval schedule. For a second group, also tested with the resistance-to-reinforcement procedure, the lines were superimposed on the green field that formerly served as the positive stimulus. A third group was tested in extinction with the combined stimuli. Control groups had no discrimination training but responding to green was nondifferentially reinforced. The control subjects responded more to all line tilts during testing than did the comparable experimental subjects, indicating that the negative stimulus had become an inhibitory stimulus. Both resistance-to-reinforcement groups revealed inhibitory gradients around the negative stimulus, but the gradient for the extinction group was relatively flat. These data are consistent with others that modify Terrace's early conclusion concerning the failure of inhibition to develop during errorless training.     

Neuron circuits: The self-exciting neuron

Conditions are determined under which a single self-exciting neuron can reach a state of permanent excitation, in the case that the neuron develops both an excitatory and an inhibitory substance as well as in the case that it develops only a single, excitatory substance. In case both substances are developed it is possible to have periodic solutions, a possibility which does not arise when the circuit consists of neurons, or of a neuron, developing only an excitatory substance.     

Identity-based inhibitory processing during focused attention

Recent data suggest that the operating principles governing inhibition of distractor-related activity may deviate from dominant models describing inhibitory processing during selective attention. Here we aimed to gain a better understanding of these data in order to determine whether they actually defy premises of current models. In addition to providing evidence against noninhibitory accounts of the data (see especially Experiment 6), the results support three main novel findings that challenge current theories. First, the data provide evidence that inhibition overpowered excitation from ongoing external input (Experiments 1–4), which suggests that inhibitory control processes are more powerful than current models indicate. Second, negative effects emerged even when targets appeared alone (Experiment 5), which suggests that selection does not play an essential role in triggering inhibitory processing. Third, relatively early distractor-related activity was affected, which supports a role for inhibition prior to action control (Experiment 3). These findings suggest a need to revise current models describing inhibition of distracting information during selective attention.     

Central inhibition-some refractory observations

The observation that the response to the second of two stimuli is delayed if a response has to be made to the first has led to the development of a theory of a central refractory state during which incoming stimuli cannot be elaborated. In the experiment reported here the two stimulus-response situations have been made as independent as possible, and under these conditions it is shown that this theory cannot be maintained in its present form. The concept that the central integrating mechanism readily becomes blocked by nay single stimulus is dismissed as nonproven and uneconomic. The present findings, however, confirm previous observations that some interference between the two situations may occur but indicate that this is not necessarily maximal immediately after the presentation of the first stimulus. It is tentatively suggested that the phenomena of central inhibition can be interpreted in terms of the interaction between the excitatory and inhibitory significance of the stimuli and the internal anticipatory set.     

Backward inhibitory learning in honeybees: a behavioral analysis of reinforcement processing

One class of theoretical accounts of associative learning suggests that reinforcers are processed according to learning rules that minimize the predictive error between the expected strength of future reinforcement and its actual strength. The omission of reinforcement in a situation where it is expected leads to inhibitory learning of stimuli indicative for such a violation of the prediction. There are, however, results indicating that inhibitory learning can also be induced by other mechanisms. Here, we present data from olfactory reward conditioning in honeybees that show that (1) one- and multiple-trial backward conditioning results in conditioned inhibition (CI); (2) the inhibition is maximal for a 15-sec interval between US and CS; (3) there is a nonmonotonic dependency on the degree of CI from the US-CS interval during backward pairing; and (4) the prior association of context stimuli with reinforcement is not necessary for the development of CI. These results cannot be explained by models that only minimize a prediction error. Rather, they are consistent with models of associative learning that, in addition, assume that learning depends on the temporal overlap of a CS with two processes, a fast excitatory and a slow inhibitory one, both evoked by a reinforcer. The fmdings from this behavioral analysis of reinforcement processing are compared with the known properties of an individual, identified neuron involved in reinforcement processing in the bee brain, to further understand the mechanisms underlying predictive reward learning.     

Pavlovian conditioned inhibition of fear during shuttlebox avoidance behavior

Three experiments explored the development and function of conditioned inhibition of fear during the acquisition and maintenance of shuttlebox avoidance behavior. The development of inhibition to an exteroceptive feedback stimulus was found to be a function of the number of successive avoidance responses to which the animal had been trained and of the duration of the intertrial interval, a parameter shown also to affect the rate of acquisition of avoidance learning. Master animals who learned the instrumental avoidance response, and yoked animals who did not, showed equivalent inhibitory fear conditioning in each experiment. The results of one experiment suggest that conditioned inhibition plays no important role in “protecting” fear conditioned to the discrete warning signal during avoidance maintenance. These data indicate that feedback stimuli develop their inhibitory properties by a Pavlovian process and that certain aspects of their function may, therefore, be readily understood within the framework of mediational two-process learning theory.     

Superconditioning from a reduced reinforcer

In four autoshaping experiments pigeons received conditioned inhibition training of the form A++, AB-, where ++ is a strong reinforcer and - is nonreinforcement. Subsequent AB+ training, in which + is a moderate reinforcer, resulted in enhanced conditioning of A, relative to an A stimulus receiving no treatment, one receiving A+ treatment, and one receiving A++ treatment. This enhancement of conditioning to Aconstitutes a demonstration of “superconditioning”. The presence of the inhibitory B sufficiently enhanced the reinforcing power of the moderate reinforcer (+) that it was able to further increase the excitation controlled by A. This occurred even though A had previously been paired with a stronger reinforcer (++). Superconditioning was also observed when A and B were extinguished prior to treatment with + or were originally neutral stimuli followed by + in the presence of an inhibitor trained elsewhere.     

Context Specificity of Excitation and Inhibition in Ambiguous Stimuli

Two experiments examined the susceptibility of excitation and inhibition to contextual change when the conditioned stimulus (CS) was ambiguous using an appetitive conditioning paradigm. In Experiment 1, excitation to a CS was lost with a context switch when inhibition had been learned to the CS in a prior feature-negative (FN) discrimination. Control groups that had received either more or less excitatory conditioning in the absence of inhibitory pretraining showed no loss. In Experiment 2 inhibition was lost with a context switch in a group that had received excitatory and then inhibitory conditioning with the CS. No loss was observed in a group that had not received excitatory pretraining. The results suggest that contexts are especially likely to control performance to ambiguous CSs when they can modulate the second of two learned associations. Results are discussed in terms of their implications for occasion setting and modern conditioning theory.     

Neurophysiologic mechanisms of internal inhibition

The experimental results obtained in the authors’ laboratory as a result of multiple recording of slow biopotentials, the recording of neuronal activity and of mathematical modeling, are reviewed. The authors conclude that the elaboration of internal inhibition is followed, and determined to a great extent, by the restriction in conduction of excitations due to the increase of inhibitory hyperpolarization and discordance in the periodicity of slow potentials, reflecting oscillations in excitability of neuronal populations in the cortex and other brain structures.     

Some determinants of inhibitory stimulus control

Interspersed reinforcement and extinction during discrimination learning generate a U-shaped gradient of inhibition about the stimulus correlated with extinction. The present work showed that extinction is not a necessary determinant of inhibitory stimulus control. In Exp. I, a reduction in the rate of reinforcement, through a shift from a multiple variable-interval 1-min variable-interval 1-min schedule to a multiple variable-interval 1-min variable-interval 5-min schedule, resulted in a post-discrimination line orientation gradient of inhibition about the stimulus correlated with the variable-interval 5-min schedule. In Exp. II, the rates of reinforcement, correlated with a pair of stimuli, were held constant during a shift from a multiple variable-interval 1-min variable-interval 1-min schedule to a multiple variable-interval 1-min differential-reinforcement-of-low-rate schedule. Inhibitory stimulus control about the stimulus correlated with the differential reinforcement of low rate was obtained. In both experiments, a reduction in the rate of responding during one stimulus and behavioral contrast during the other stimulus preceded the observation of inhibitory stimulus control.     

Mobility,extraversion, and neuroticism

Mobility, a property of Pavlovian higher nervous activity, was measured using an induction task. Negative induction, following presentation of an excitatory stimulus, is an index of mobility of excitation and positive induction, following presentation of an inhibitory stimulus is an index of mobility of inhibition. Mobility scores from 27 male and 37 female first-year psychology students, age 17 to 42 years were related to extraversion and neuroticism derived from Eysenck’s Personality Inventory (Form A). Mobility of excitation and mobility of inhibition were found to be unrelated. Individual differences in the magnitude and latency of induction indicate that individuals can be typed according to low, medium, or high mobility of either nervous process. No relationship was found between neuroticism and mobility, and no relationship was found between extraversion and mobility of excitation. However, a significant negative relationship was found between extraversion and mobility of inhibition, providing a link between Pavlovian properties of the nervous system and Eysenck’s personality dimension of extraversion.     

刺激前alpha振荡对视知觉的影响

人类对感觉阈限附近的视觉刺激的知觉不总是一致的。为探究这种视知觉不一致的现象及其神经机制, 一些研究者关注刺激前脑内自发alpha神经振荡(8~13 Hz)对视知觉的影响。近年来的研究发现, 刺激前alpha振荡能量的降低能提高被试的探测击中率, 但不能提高知觉精确度; 而刺激前alpha振荡的相位能预测被试能否成功探测刺激。刺激前alpha能量被认为调控了视皮层的基础活动强度; alpha能量的降低反映了皮层基础活动的增强, 进而提高了对较弱刺激的探测率。刺激前alpha相位则被认为调控了皮层兴奋和抑制的时间; 大脑在刺激呈现时的不同状态(兴奋/抑制)决定了最终的知觉结果。     

Differential brain responses when applying criterion attribute versus family resemblance rule learning

Subsystems of category learning have been identified on the basis of general domains of content (e.g., tools, faces). The present study examined categories from the standpoint of internal structure and determined brain topography associated with expressing two fundamentally different category rule structures (criterion attribute, CA, and family resemblance, FR). CA category learning involves processing stimuli by isolated features and classifying by properties held by all members. FR learning involves processing stimuli by integral wholes and classifying on overall similarity among members without sharing identical features. fMRI BOLD response to CA and FR categorization was measured with pseudowords as stimuli. Category knowledge for both tasks was mastered prior to brain imaging. Areas of activation emerged unique to the structure of each category and followed from the nature of the rule abstraction procedure. CA categorization was implemented by strong target monitoring and expectation (medial parietal), rule maintenance in working memory, feature selection processes (inferior frontal), and a sensitivity to high frequency components of the stimulus such as isolated features (anterior temporal). FR categorization, consistent with its multi-featural nature, involved word-level processing (left extrastriate) that evoked articulatory rehearsal (medial cerebellar). The data suggest category structure is an important determinant of brain response during categorization. For instance, anterior temporal structures may help attune visual processing systems to high frequency components to support the learning of criterial, highly predictive rules.