Precentral cortex unit activity during the M-wave and contingent negative variation in behaving squirrel monkeys.
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Biomedical subjects
Publications and source records attributed to E S Boyd.
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Typical M-waves and CNVs were produced in squirrel monkey frontal and rostral parietal cortex by light cues, as well as by tone cues. With light cues, M-waves were about 50% as large as with tone cues, whereas CNVs were of about the same magnitude with either type of cue. Typical M-waves and CNVs were also produced when avoidance of tail-shock was substituted for food as a reinforcer. The magnitude of both ERPs appeared to follow an inverted-U function with increasing shock intensity. The use of tail-shock as a reinforcer generated more complexities both in ERP changes and in behavior than did food. In some animals, at high shock intensities, ERP increased, rather than decreased, as each session progressed. In some animals, a combination of food and avoidance of tail-shock produced a decrease, rathan than an increase, in ERPs and did not improve behavioral performance. Furthermore, animals would not perform effectively on a CNV paradigm with punishment for aborting the trial. It was also demonstrated that increased stress (immobilization of head or tail) resulted in a reduced magnitude of both the M-wave and the CNV. This may account for some of the complexities encountered with avoidance of tail-shock as a reinforcer.
A typical CNV paradigm, with food as reward, evokes in the squirrel monkey's post-arcuate and post-central cortices both M-waves in response to the cues and what appears to be a CNV in the interstimulus interval. Both wave forms appear to be generated locally in cortex and to be more closely related temporally to the cues than to the animal's behavioral response. The CNV, like the M-wave, appears to reflect the animal's level of interest in obtaining the reward. It is smaller when the animal misses the cues than when he responds correctly. It decreases as food pellets are consumed. It varies with the level of interest of the animal in a particular type of food pellet, as judged from the animal's behavior. At some recording sites in some animals it appears to reflect both interest in the reward and interest in manipulating the environment to obtain the reward. It can be maintained for at least 2 sec, but differs from the human CNV in that, with the paradigm used, it is not well resolved by the imperative cue. Averaging evoked activity with the animal's motor response as reference point reveals another negative wave form which just precedes the motor response.
The effects of low doses of nabilone, chlorpromazine, pentobarbital, meprobamate, diazepam, chlordiazepoxiOde and d-amphetamine on behavioral responding to cues signalling the availability of food rewards, and on the M-wave, an evoked cortical potential previously reported to reflect the conditioned incentive value of the cues were determined in the squirrel monkey. Nabilone and chlorpromazine simultaneously depressed both the M-wave and behavior. Pentobarbital, meprobamate, diazepam and chlordiazepoxide could depress the M-wave without depressing behavior. This effect was most marked with diazepam. The only augmentation of the M-wave observed was the d-amphetamine, and this occurred in only one of five animals. The benzodiazepines were the only drugs to augment behavioral output. However, diazepam occasionally increased the number of cures responded to while concomitantly decreasing both total behavioral output and the amplitude of the M-wave. It is concluded that the M-wave cannot directly reflect the incentive value of the cue, but must rather reflect something that tends to parallel this value.
As squirrel monkeys learned that a tone cue signalled the availability of a food reward there appeared in pre- and post-central cortex a surface negative waveform in response to the cue. This waveform disappeared, concomitantly with the animal's behavioral response, during extinction and pseudoconditioning. It was independent of changes in cue intensity sufficient to markedly alter earlier components of evoked activity and it disappeared when evoked activity was averaged with the behavioral response, rather than the cue, as reference point. It was concluded that this evoked potential reflects cortical activity somewhere in the stimulus-response sequence between the recognition of the cue and the decision to act.
It was shown that the late negative wave evoked in squirrel monkey cortex by tone cues signalling the availability of reward was smaller when the animal did not respond to the cue and often disappeared if the animal ignored many cues in succession. It decreased in amplitude as a test session progressed, and the rate of this decrease increased if the rate of reinforcement was increased. In some animals the size of the evoked wave was smaller if the animal was not fasted before a test session. In some animals the average size of a significant positive correlation with the number of cues responded to and a negative correlation with the animal's average reaction time. It is concluded that the evoked wave is probably a reflection of the conditioned incentive value of the cue.
Changing animals from a schedule on which they had to bar-press after a tone cue to get a food pellet, to a schedule on which the food pellet was presented automatically after the tone cue and the manipulandum was not available, had no effect on the latency, amplitude, or shape of the M-wave evoked by the tone cue. The amplitude of the evoked wave did, however, decrease more rapidly, as a test session progressed, with the latter schedule than with the former schedule. Failure to fast animals before test sessions markedly reduced the amplitude of the M-wave. Variation in the type of food pellet used caused marked variations in the amplitude of the evoked response. In all cases, there was a high degree of correlation between M-wave amplitude and a behavioral indication of the animal's interest in the reward. It was concluded that the M-wave reflects the conditioned incentive value of the cue, or in other words, the level of interest of the animal in the reward.
A surface-negative wave, evoked by tone cues, appeared in monkey post-arcuate cortex as the monkey learned that the cue signaled the availability of reward. This evoked activity was depressed, concomitantly with changes in the animal's behavioral responding, by doses of delta9-tetrahydrocannabinol (delta9-THC) as low as 0.032 mg/kg and of pentobarbital as low as 4 mg/kg. Pentobarbital tended to increase the latency of the evoked wave, an effect not seen with delta9-THC.
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