[Action of lysergic acid diethylamide on the operant conditioning activity of rats].
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Operant conditioning of the H-reflex produces plasticity at several sites in the spinal cord, including the motoneuron. This study assessed whether this spinal cord plasticity is accompanied by changes in motor unit contractile properties. Thirty-one adult male Sprague-Dawley rats implanted for chronic recording of triceps surae electromyographic activity and H-reflex elicitation were exposed for at least 40 days to HRup or HRdown training, in which reward occurred when the H-reflex was greater than (12 HRup rats) or less than (12 HRdown rats) a criterion value, or continued under the control mode in which the H-reflex was simply measured (7 HRcon rats). At the end of H-reflex data collection, rats were anesthetized and the contractile properties of 797 single triceps surae motor units activated by intraaxonal (or intramyelin) current injection were determined. Motor units were classified as S, FR, Fint, or FF on the basis of sag and fatigue properties. Maximum tetanic force and twitch contraction time were also measured. HRdown rats exhibited a significant increase in the fatigue index of fast-twitch motor units. This resulted in a significant decrease in the percentage of Fint motor units and a significant increase in that of FR motor units. HRup conditioning had no effect on fatigue index. Neither HRup nor HRdown conditioning affected maximum tetanic force or twitch contraction time. These data are consistent with the hypothesis that conditioning mode-specific change in motoneuron firing patterns causes activity-dependent change in muscle properties.
Two issues concerning sensorimotor EEG operant conditioning, or biofeedback, as a therapeutic modality for the treatment of seizure disorders are the focus of this review. The first relates to the question of whether relevant physiological changes are associated with this procedure. This question is addressed through review of an extensive neurophysiological literature that is likely unfamiliar to many clinicians but that documents both immediate and sustained functional changes that are consistent with elevation of seizure thresholds. The second focuses on the clinical efficacy of this method and whether it should carry the designation of "experimental". This designation is challenged through an assessment of over 25 years of peer-reviewed research demonstrating impressive EEG and clinical results achieved with the most difficult subset of seizure patients.
Xanthan gum production under several operational conditions has been studied. Temperature, initial nitrogen concentration and oxygen mass transfer rate have been changed and average molecular weight, pyruvilation and acetylation degree of xanthan produced have been measured in order to know the influence of these variables on the synthesised xanthan molecular structure. Also, xanthan gum solution viscosity has been measured, and rheological properties of the solutions have been related to molecular structure and operational conditions. The Casson model has been employed to describe the rheological behaviour. The parameter values of the Casson model, tau(0) and K(c), have been obtained for each polysaccharide synthesised under different operational conditions. Both pyruvilation and acetylation degrees and average molecular weight of xanthan increase with fermentation time at any operating conditions. Xanthan molecules with the highest average molecular weight have been obtained at 25 degrees C. Nevertheless, at this temperature acetate and pyruvate radical concentration are lowest. Nitrogen concentration in broth does not show any clear influence over xanthan average molecular weight, although with high nitrogen source concentration xanthan with low pyruvilation degree is produced.
In the operant conditioning experiment described by HEFFERLINE et al. (1959) a subject terminates or postpones an aversive noise stimulation by producing a particular myoelectrical activity (operant response : OR). These authors observe an increase of OR occurrence frequency in subjects ignoring the aim of the experiment (naive subjects), and they come to the conclusion that this method is efficient. Nevertheless, the validity of this conclusion is questioned by the lack of control experiments. Hence, this work has to be restudied. We have submitted 13 subjects to a control experiment. Subjects were placed under conditions similar to those used by HEFFERLINE et al., but without any contingency relating the myoelectric activity with the aversive stimulus. The results of these open-loop experiments showed that in a given subject, OR occurrence frequency may vary in large proportions as a function of time. In HEFFERLINE's naive subjects, the increase of occurrence frequency remained completely within the limits of spontaneous variations observed in our control subjects. Therefore, this increase is not significant. We have submitted 18 subjects to a conditioning experiment. 14 subjects among them produced more OR than those of the control subjects during the closed-loop experiment. We conclude that these 14 subjects have been actually conditioned.
Operant conditioning of the sensorimotor rhythm of the human electroencephalogram with time-outs contingent on epileptiform activity reduces epileptic seizure rates in patients whose seizures are not well controlled by medication. A comparison of this procedure with time-out training alone demonstrates that operant conditioning of the sensorimotor rhythm is neither necessary nor sufficient for seizure reduction.
EEG operant conditioning in an alumina-gel monkey model (N = 14) to decrease EMG, to increase 9 Hz or decrease 9 Hz, and to increase 23 Hz (18 Hz and 26 Hz in 2 pilot monkeys), respectively, was not consistently beneficial in reducing seizures. The data suggested: (1) that desynchronization of the EEG by reinforcing 18-26 Hz decreased in some animals the extent and severity of seizures while increasing seizure frequency; (2) that attending during conditioned EMG suppression reduced seizures somewhat; and (3) that the operant conditioning setting became stressful to the majority of monkeys under certain conditions, precipitating status epilepticus, gastrointestinal disturbances, and shock, which culminated in the death of 3 animals. Certain precautions are discussed in the therapeutic application of this technique to epileptic patients.
The marine mollusc Aplysia californica exhibits a wide range of nonassociative and associative forms of learning. Recently, we found that the learning repertoire of Aplysia includes operant conditioning (Cook & Carew, 1986, 1989b). The behavior we examined is a naturally occurring, side-to-side head-waving response used by Aplysia in seeking food, obtaining a foothold, and egg laying. Aplysia can be operantly conditioned to reduce head-waving to one side of their body if such a response results in exposure to bright uniform-field illumination, which the animals find aversive. An essential step toward achieving a mechanistic understanding of operant conditioning is to identify and characterize the reinforcement pathway used during the learning. Toward this end, we wished to determine which of the peripheral visual pathways in Aplysia are critical for performance of the operant task. Previous experiments indicated that photic input from the optic and rhinophore nerves functionally inhibited motor neurons that participate in the operant response (head-waving), while photic input from the oral veil nerves excited these same motor neurons (Cook & Carew, 1989c). These findings suggested the hypothesis that one or both of these pathways could play an important role in mediating reinforcement during training. To explore this possibility we operantly trained animals that had received chronic bilateral transections of either the optic and rhinophore nerves or the oral veil nerves C1-C3 (in conjunction with transection of the optic and rhinophore nerves). We found that operant conditioning was not disrupted by ablation of input from the eyes and rhinophores. By contrast, ablation of input from the oral veil (together with that from the eyes and rhinophores) abolished operant conditioning. Thus, the oral veil nerves play a critical modulatory role in operant conditioning of head-waving. This observation further suggested that photic input from the oral veil is conveyed to the CNS via the oral veil nerves. In a final experiment we confirmed that stimulation of the oral veil with light evokes increased afferent activity in the oral veil nerves C1-C2. These results support the idea that the oral veil nerves contain processes that are critical components of the reinforcement pathway for operant conditioning of head-waving.
Sympathetic adrenergic nervous activity during operantly conditioned hypertension was evaluated by assessing the effects of specific alpha-(phentolamine or phenoxybenzamine) and beta- (propranolol) adrenergic blockers in baboons reinforced for increasing diastolic pressure in daily, 12-h sessions. In the first 10 min of control (no blockade) sessions, mean heart rate increased 24 bpm (21%) above the value for the 10 min immediately prior to the beginning of the sessions; systolic pressure increased 27 mm Hg (22%) and diastolic pressure increased 24 mm Hg (31%). Beta-blockade eliminated the tachycardia but did not attenuate the increased blood pressure. Alpha-blockade did not attenuate the increased blood pressure significantly either. Combined alpha- and beta-blockade did significantly attenuate the increase in diastolic pressure, but consistent, significant increases in systolic pressure (17 mm Hg, 17%) and diastolic pressure (16 mm Hg, 26%) still occurred. The results support the participation of the sympathetic adrenergic nervous system in producing operantly conditioned blood pressure changes, but the results are also consistent with the additional participation of nonadrenergic factors in operantly conditioned hypertension.
Aplysia can readily exhibit operant conditioning of their head-waving response when bright light is used as aversive reinforcement (Cook and Carew, 1986). In the first paper of this series (Cook and Carew, 1989a), we showed that the electromyographic (EMG) activity of a discrete band of neck muscles, the lateral columellar muscles (LCMs) of Aplysia is significantly correlated with the component of head-waving (the horizontal component) that is modified during operant conditioning. In the present paper, we asked whether the EMG activity of the LCMs themselves could also be contingently modified, using the same procedures that produce operant conditioning of the behavioral response. Differential EMG from the LCMs was recorded in freely behaving animals with chronically implanted muscle cuff electrodes. Animals receiving aversive reinforcement (bright light) that was contingent upon specific patterns of LCM activity readily learned to alter their differential EMG output. Like operant conditioning of the head-waving response, this operant modification of LCM activity was rapidly acquired and was specific to the contingencies of reinforcement. These results show that a restricted group of muscles, the LCMs, exhibit the essential features of the head-waving system observed at the behavioral level: (1) their activity is significantly correlated with head-waving behavior, and (2) the LCMs are capable of operant modification of their output. Thus, this restricted response system provides a useful preparation for examining the neural mechanisms of operant conditioning of head-waving in Aplysia.
In this study, we demonstrate neural changes that occurred during operant conditioning of the aerial respiratory behavior of Lymnaea stagnalis. Aerial respiration in Lymnaea occurs at the water interface and is achieved by opening and closing movements of its respiratory orifice, the pneumostome. This behavior is controlled by a central pattern generator (CPG), the neurons of which, as well as the motoneurons innervating the pneumostome, have previously been identified and their synaptic connections well characterized. The respiratory behavior was operantly conditioned by applying a mechanical stimulus to the open pneumostome whenever the animal attempted to breathe. This negative reinforcement to the open pneumostome resulted in its immediate closure and a significant reduction in the overall respiratory activity. Electrophysiological recordings from the isolated CNSs after operant conditioning showed that the spontaneous patterned respiratory activity of the CPG neurons was significantly reduced. This included reduced spontaneous activity of the CPG interneuron involved in pneumostome opening (input 3 interneuron) and a reduced frequency of spontaneous tonic activity of the CPG interneuron [right pedal dorsal 1 (RPeD1)]. The ability to trigger the patterned respiratory activity by electrical stimulation of RPeD1 was also significantly reduced after operant conditioning. This study therefore demonstrates significant changes within a CPG that are associated with changes in a rhythmic homeostatic behavior after operant conditioning.
In this study, we operantly conditioned the aerial respiratory behaviour of the freshwater snail Lymnaea stagnalis. Aerial respiration in Lymnaea stagnalis is accomplished by the spontaneous opening and closing of its respiratory orifice, the pneumostome, at the water surface. Weak tactile stimulation of the pneumostome area, when the pneumostome is open, evoked only the pneumostome closure response, which is one aspect of the escape-withdrawal reflex. Pneumostome stimulation resulted in its closure and the termination of aerial respiratory activity. A contingent tactile stimulation paradigm was used to operantly condition the animals. Stimulation of the pneumostome whenever the animal attempted to breathe resulted in significantly fewer attempts to open the pneumostome as training progressed. The latency of the first breath (subsequent to stimulation), the number of breaths and the total breathing time were measured before and after each training period. Significant, quantifiable changes in these behavioural parameters were observed only in the operant conditioning group animals. Control animals receiving tactile stimulation to their pneumostome not contingent upon pneumostome opening movements (yoked controls) or those that were physically prevented from surfacing to breathe (hypoxic controls), did not exhibit significant changes in these behavioural parameters. Our data provide the first direct evidence for operant conditioning of respiration in any animal.
An operant conditioning study was carried out on Wistar rat pups of brain-growth-spurt age to assess the plasticity of apical dendritic branching and the numerical spine densities of CA3 pyramidal neurons of the hippocampus. In rapid Golgi-stained material we studied the effects of operant conditioning commencing at 16 days of age and terminating on the 23rd day of age. The data of the learning group (L) were compared with those of the sham experimental control group (E) and of the control group reared under standard conditions (S). The results revealed a significant (ANOVA F test) increase in the number of branching points of the dendrites of the learning group compared with the other groups. The grand averages of spine densities also showed an overall increase in the learning group over the sham-experimental and standard groups. Segmental comparisons revealed a more interesting observation that significant changes in the spine densities of the learning and sham-experimental groups occurred in some segments of dendrites but not in all. A few segments showed moderate spine decreases also, compared with the control group. The spine differences of segments were mainly in the stalked types. Excrescences which are spines on which mossy fibers are known to synapse increased in both learning and sham-experimental groups, but not significantly (ANOVA). The dendritic branching and spines of nonmossy inputs of certain dendritic segments significantly increased after the learning experience in the hippocampus of the growing brain.
Feeding behavior in Aplysia can be modified by operant conditioning in which contingent reinforcement is conveyed by the esophageal nerve (E n.). A neuronal analogue of this conditioning in the isolated buccal ganglia was developed by using stimulation of E n. as an analogue of contingent reinforcement. Previous studies indicated that E n. may release dopamine. We used a dopamine antagonist (methylergonovine) to investigate whether dopamine mediated the enhancement of motor patterns in the analogue of operant conditioning. Methylergonovine blocked synaptic connections from the reinforcement pathway and the contingent-dependent enhancement of the reinforced pattern. These results suggest that dopamine mediates at least part of the neuronal modifications induced by contingent reinforcement.
Operant conditioning of the H-reflex, the electrical analog of the spinal stretch reflex, in freely moving rats is a relatively simple model for studying long-term supraspinal control over spinal cord function. Motivated by food reward, rats can gradually increase (i.e., up-condition) or decrease (i.e., down-condition) the soleus H-reflex. Earlier work showed that corticospinal tract transection prevents acquisition and maintenance of H-reflex down-conditioning while transection of other major spinal cord tracts does not. This study explores the effects on acquisition of up-conditioning of the right soleus H-reflex of mid-thoracic transection of: the right lateral column (LC, five rats) (containing the rubrospinal, vestibulospinal, and reticulospinal tracts); the entire dorsal column (DC, six rats) [containing the main corticospinal tract (CST) and the dorsal ascending tract (DA)]; the CST alone (five rats); or the DA alone (seven rats). After initial (i.e., control) H-reflex amplitude was determined, the rat was exposed for 50 days to the up-conditioning mode in which reward was given when the H-reflex was above a criterion value. H-reflex amplitude at the end of up-conditioning was compared to initial H-reflex amplitude. An increase > or =20% was defined as successful up-conditioning. In intact rats, H-reflex amplitude at the end of up-conditioning averaged 164% (+/-10%, SE), and 81% were successful. In the present study, LC and DA rats were similar to intact rats in final H-reflex amplitude and percent successful. In contrast, results for DC and CST rats were significantly different from those of intact rats. In the six DC rats, final H-reflex amplitude averaged 105% (+/-3)% of control and none was successful; and in the five CST rats, final H-reflex amplitude averaged 94% (+/-3)% and none was successful. The results indicate that the main CST, located in the dorsal column, is essential for H-reflex up-conditioning as it is for down-conditioning, while the dorsal column ascending tract and the ipsilateral lateral column (containing the main rubrospinal, vestibulospinal, and reticulospinal tracts) do not appear to be essential.