Discriminative vocal conditioning in rhesus monkeys: Evidence for volitional control?

Both emotional and volitional systems are invoked in explaining the control of human vocalizations, but vocal behavior of nonhuman primates is often believed to be totally under emotional control. Monkeys' poor performance on conditioning tasks, especially discriminative conditioning of vocalization (DCV), has been cited as evidence against volitional control. In Experiment 1, rhesus monkeys failed a DCV task in which food was given for vocalizations emitted during an arbitrary visual stimulus, but in Experiment 2, monkeys showed clear discriminative performance when an otherwise comparable shock-avoidance DCV procedure was used. This evidence that monkeys possess some degree of volitional control has implications for the relation between animal vocalizations and the origin of human speech.     

Effects of d-amphetamine on observing behavior in the squirrel monkey

Four squirrel monkeys were trained to press a lever, which produced stimuli indicating availability or non-availability of reinforcement for pushing a key. Food reinforcements were available for the key response at random intervals with an average rate of 1 per min. When food was available, a single lever response produced a red light behind the key. Reinforcement availabilities and red keylights remained until terminated by a reinforced key response. When reinforcement was not available, each lever response produced a 0.5-sec green light on the key. Except after lever responses, the key remained dark. Under this procedure, lever responses functioned as observing behavior in that they produced discriminative stimuli correlated with the availability or non-availability of reinforcement for key responses. The procedure generated a high rate of responding on the lever, short latencies of the key response after onset of red lights and few responses to the key in the absence of red lights. Intra-muscular d-amphetamine, in doses from 0.125 to 1.0 mg/kg, abolished both observing behavior and key responding for periods that increased as a function of dose. However, both observing and key rates were increased at the smallest dose in two subjects whose performances included responding to the key in the absence of red lights. Results are discussed in relation to previous findings regarding effects of amphetamines on operant behavior and on observing and monitoring performance.     

Discrimination between reinforced action patterns in the rat

Laboratory rats were rewarded for face-washing, rearing, or scratching by being given the opportunity to press retractable levers for food reward. Yoked control animals received the same number of lever presentations and food rewards, but did not have to face-wash, rear, or scratch to obtain the levers. The experimental animals showed increases in number of bouts of reinforced target behavior above control levels, and the total amount of time spent face-washing increased when a 1.5-sec criterion for reinforced bout length was introduced. The activities in this experiment were made to serve a discriminative as well as an instrumental function, since the cue to tell the rat which lever to press for reward when the levers were presented was the activity that the rat had engaged in to obtain lever presentation. In two separate experiments high levels of discrimination between behaviors were obtained. Discrimination was worse following scratching than after other actions, and scratching also showed relatively poor instrumental conditioning. The relationship between Pavlovian and instrumental conditioning processes in this situation is discussed.     

Roles of nucleus accumbens and basolateral amygdala in autoshaped lever pressing

Initially-neutral cues paired with rewards are thought to acquire motivational significance, as if the incentive motivational value of the reward is transferred to the cue. Such cues may serve as secondary reinforcers to establish new learning, modulate the performance of instrumental action (Pavlovian-instrumental transfer, PIT), and be the targets of approach and other cue-directed behaviors. Here we examined the effects of lesions of the ventral striatal nucleus accumbens (ACb) and the basolateral amygdala (BLA) on the acquisition of discriminative autoshaped lever-pressing in rats. Insertion of one lever into the experimental chamber was reinforced by sucrose delivery, but insertion of another lever was not reinforced. Although sucrose was delivered independently of the rats' behavior, sham-lesioned rats rapidly came to press the reinforced but not the nonreinforced lever. Bilateral ACb lesions impaired the initial acquisition of sign-tracking but not its terminal levels. In contrast, BLA lesions produced substantial deficits in terminal levels of sign-tracking. Furthermore, whereas ACb lesions primarily affected the probability of lever press responses, BLA lesions mostly affected the rate of responding once it occurred. Finally, disconnection lesions that disrupted communication between ACb and BLA produced both sets of deficits. We suggest that ACb is important for initial acquisition of consummatory-like responses that incorporate hedonic aspects of the reward, while BLA serves to enhance such incentive salience once it is acquired.     

The conditioned reinforcement of repeated acquisition

Three monkeys were trained to emit a chain of three responses on three separate levers in a set of six levers to obtain food. The chain producing food (correct chain) was changed each day. During a trial, a press on any lever produced a feedback stimulus; a press on a correct lever produced an additional distinctive stimulus; the third correct press produced a food pellet. Test sessions in which either the food or the distinctive stimuli were removed were interspersed with baseline sessions. In tests without food presentations, the subjects acquired the correct chain rapidly, with a level of accuracy comparable to baseline. Removing the distintive stimuli for either the first or second member of the correct chain greatly retarded acquisition of that member of the chain. Removing all distinctive stimuli often reduced accuracy throughout the chain to chance level, even though food was presented following each correct chain. These results were interpreted as evidence that the distinctive stimuli presented after correct responses functioned as conditioned reinforcers. Reductions in accuracy following an omitted distinctive stimulus indicated that they were also discriminative stimuli for correct responding in their presence.     

Cortical control of conditioned and spontaneous vocal behavior in rhesus monkeys

Rhesus monkeys were trained on a discriminative conditioning of vocalizations paradigm, a task that requires volitional vocal control. The monkeys' spontaneous vocal and other social behavior was also assessed. Each monkey then underwent ablation of anterior cingulate cortex, ablation of the homologus of Broca's and Wernicke's areas, or no lesion. The anterior cingulate lesions severely disrupted both conditioned and spontaneous vocal behavior, but had no effect on nonvocal social behavior or on a discriminative conditioning task requiring a nonvocal response. The lesions of the homologs of Broca's and Wernicke's areas had no effects on the behaviors studied. These findings have implications for the comparative neurology of primate vocalization and speech.     

The beginning of conversation: Early patterns of mother-infant vocal responsiveness

Microanalytic techniques were used to characterize the structure of the prespeech communication of 4-month-old infants and their mothers. Two observers continuously recorded the interactive behavior of mothers and their infants during hour-long observations in the homes of 25 families. Loglinear models were used to examine the extent to which the vocal behavior of one person was conditional upon the vocal behavior of the partner. Within the limits of this microanalytic approach, analyses indicated that patterns of mother-infant vocal exchange were structurally similar to patterns of adult conversation. Initial vocal responses were followed by suppression of vocalization, allowing the partner to join the conversation. The comparative effectiveness of vocal behavior as an elicitor of vocalization and as a response to vocalization was shown for mothers and infants relative to the other behaviors observed. Vocalization served as a modulator of visual attentiveness: When the partner was not visually attentive, vocalization elicited visual attention.     

An adjusting avoidance schedule

A shock-avoidance schedule is described in which the animal accumulated 5 sec of safe time whenever it pressed a lever. With this schedule, the animal was not differentially reinforced for long pauses between responses; and, consistent with this property of the schedule, the probability that the animal would press the lever was not related in any regular way to the amount of time that had elapsed since its preceding lever press. Other features of the performance are also described.

If the animal was given a warning stimulus whenever it came within 5 sec of a shock, it tended to spend more time in the close temporal vicinity of the shock and less time at the maximum temporal distance from shock.

     

Signal functions in discriminated avoidance behavior

Four rats were trained to lever press under a discriminated avoidance/escape schedule in which separately signalled safe and warning periods were 100 sec and 32 sec respectively. The auditory and not the visual component of the compound warning signal became associated with the discriminative control of lever pressing. Avoidance behavior also came under temporal control, in that the probability of lever pressing increased as the warning period progressed. Timing began with the onset of the warning signal rather than the offset of the safe signal. However, after the warning signal had been progressively eliminated, timing began with the offset of the safe signal. When neither signal was normally available, the temporal distribution of avoidance behavior changed markedly. Drifts in the temporal distribution of lever pressing occurred throughout the study; these were manipulated for two animals.     

Differential reinforcement of lever-press durations: Effects of deprivation level and reward magnitude

Three experiments investigated the performance of rats on a task involving differential reinforcement of lever-press durations. Experiment 1, which employed a discrete-trials procedure, manipulated deprivation level between subjects and reward magnitude within subjects. The minimum lever-press duration which would result in reward was varied from .4 to 6.4 sec. It was found that low deprivation resulted in longer mean durations and less response variability at the higher criterial values than did high deprivation. The magnitude of reward was not found to affect performance. Experiment 2 manipulated reward magnitude between subjects while holding deprivation level constant, and used the same general procedures as in Experiment 1. Small reward resulted in longer mean lever-press durations and less variability in responding than did large reward at the higher criterial values. The intertrial intervals were omitted in Experiment 3 in which deprivation level was varied between subjects and reinforcement was delivered only for response durations extending between 6.0 and 7.6 sec. Low deprivation resulted in longer mean lever-press durations and less response variability than did high deprivation, but the probability of a rewarded press duration did not differ between groups. The results overall are consistent with the hypothesis that low deprivation and small reward magnitude lead to weaker goal-approach responses and, hence, to less competition with lever holding. The deprivation and reward magnitude manipulations did not appear to influence lever holding performance by affecting the ability of animals to form temporal discriminations.     

Vocalization and looking behaviour of Down's syndrome infants.

The effect of contingent and non-contingent vocal stimulation on vocalization and looking behaviour was investigated for 10 Down's syndrome infants. There were two experimental sessions consisting of four stages, namely 1 min base-line, 1 min experimental condition, a further 1 min base-line and a final 2 min experimental condition. Maternal vocal stimulation was contingent upon infant vocalization for the experimental conditions of one session and was presented every 10 s independent of infant vocalization for the experimental conditions of the other session. Neither condition increased infant vocalization but there was a significant decrease in looking at mother during the base-line stages which followed maternal vocalization (F = 5.07, d.f. = 4, 28, P less than 0.01). It is suggested that looking behaviour may be an important indication of competence in pre-linguistic social communication for developmentally delayed infants.     

Contingency effects with maintained instrumental reinforcement

In three experiments we investigated the effect on the performance of thirsty rats of varying the instrumental contingency between lever pressing and the delivery of a saccharin reinforcer. In Experiment 1, the subjects performed more slowly in a non-contingent condition, in which the momentary probability of reinforcement was unaffected by whether or not the animals pressed, than in a contingent condition in which the reinforcer was never presented except following a lever press. This was true of performance under both random ratio and interval schedules in which the function determining the probability of reinforcement following a lever press remained the same across the contingent and non-contingent conditions. Experiment 2 demonstrated that instrumental performance was less affected when the contingency was degraded by the introduction of free reinforcers if these reinforcers were signalled. In Experiment 3, lever pressing was reinstated to some degree after non-contingent training by giving non-reinforced exposure to the operant chamber in the absence of the lever. These results suggest that free reinforcers depress instrumental behaviour through a performance mechanism engaged by their ability to support conditioning of the contextual cues.     

Effect of drugs on response-duration differentiation VII: response-force requirements

Rats were trained to press a lever for at least 1 s but for less than 1.3 s. The force required to press the lever was then increased or decreased by 10, 15, or 20 g. Increases in the force requirements for lever pressing decreased timing accuracy, but decreases in the force requirement had the opposite effect. Accuracy decreases at increasing force requirements were characterized by an increase in the relative frequency of responses that were too short to meet the reinforcement criterion. In contrast, increases in accuracy when the force requirements were decreased were characterized by increases in response durations that met the reinforcement criterion and decreases in the relative frequency of responses that were too short to produce the reinforcer. Phencyclidine (PCP) and methamphetamine produced dose-dependent decreases in accuracy that were associated primarily with increases in the relative frequency of short response durations, although methamphetamine also produced increases in long response durations at some doses. When the effects of PCP were determined with the force requirement increased by 10 g or decreased by 15 g, the cumulative response-duration distribution shifted toward even shorter response durations. When the effects of methamphetamine were determined with the force requirement on the lever increased by 10 g, the cumulative frequency distribution was shifted toward shorter response durations to about the same extent as it had been before force requirements increased; however, when the force required to press the lever was decreased by 15 g, these shifts toward shorter response durations almost completely disappeared. These results show that increases and decreases in the force requirements for lever pressing have different effects on the accuracy of temporal response differentiation.     

Orbital prefrontal cortex and guidance of instrumental behavior of rats by visuospatial stimuli predicting reward magnitude

The orbital prefrontal cortex (OPFC) is part of a circuitry mediating the perception of reward and the initiation of adaptive behavioral responses. We investigated whether the OPFC is involved in guidance of the speed of instrumental behavior by visuospatial stimuli predictive of different reward magnitudes. Unoperated rats, sham-lesioned rats, and rats with bilateral lesions of the OPFC by N-methyl-D-aspartate (NMDA) were trained in a visuospatial discrimination task. The task required a lever press on the illuminated lever of two available to obtain a food reward. Different reward magnitudes were permanently assigned to lever presses to respective sides of the operant chamber; that is, responses to one lever (e.g., the left one) were always rewarded with one pellet and responses to the other lever with five pellets. On each trial, the position of the illuminated lever was pseudorandomly determined in advance. Results revealed that OPFC lesions did not impair acquisition of the task, as the speed of conditioned responses was significantly shorter with expectancy of a high reward magnitude. In addition, during reversal, shift and reshift of lever position–reward magnitude contingencies and under extinction conditions, performance of the OPFC-lesioned and control groups did not differ. It is concluded that the OPFC in rats might not be critical for adapting behavioral responses to changes of stimulus–reward magnitude contingencies signaled by visuospatial cues.     

Effects of discrete-trial and free-operant procedures on the acquisition and maintenance of successive discrimination in rats.

Rats were trained on a successive discrete-trial discrimination between two tonal stimuli to examine the effects of availability of a lever during intertrial intervals. In the discrete-trial condition, in which a lever was removed from the chamber during intertrial intervals, 10-s trials were initiated by the presentation of both discriminative stimulus and lever. In the free-operant condition, in which a lever was present during both trials and intertrial intervals, 10-s trials were initiated only by the presentation of a discriminative stimulus. Experiment 1 employed 50-s intertrial intervals and demonstrated that discriminative performances were acquired faster and maintained better in the free-operant conditions than in the discrete-trial conditions. Experiment 2 employed 5-s intertrial intervals and showed that poor discriminative performances in the discrete-trial conditions were improved. These results indicate that the presentation of a lever to start a trial can overshadow or mask the control by a discriminative stimulus and thereby obstruct the acquisition and maintenance of discriminative performances. Furthermore, the overshadowing or masking effects are strengthened as a function of the duration of intertrial intervals.     

Positive contrast in the rat: a test of the additivity theory

Rats were trained to lever press for food on a multiple variable-interval variable-interval schedule, then shifted to a multiple variable-interval extinction schedule. For six subjects (group L), schedule components were signalled by the presence or absence of a flashing light emitted from an alternate, "signal" lever. For four subjects (group T), schedule components were signalled by two distinct auditory tones. Contrary to the predictions of an additivity theory based on the summation of response classes, contacts on the signal lever did not increase after the schedule shift in group L. However, nine of the ten subjects in the study demonstrated positive contrast effects on the operant lever. In a subsequent test for stimulus control, enhancement and suppression by the discriminative stimuli were found in these same nine subjects. An additivity theory based on the summation of excitatory processes, rather than response classes, appears to account for these data.     

An assessment of factors contributing to instrumental performance for sexual reward in the rat

Male and female rats were gonadectomized, implanted in adulthood with capsules containing either testosterone propionate (TP) or cholesterol, and were trained to lever press for access to an oestrous female. When lever press performance was tested in extinction, only the male rats implanted with TP displayed significantly higher levels of responding than controls, demonstrating that lever pressing for oestrous females as a reward is sexually dimorphic. Ejaculation patterns from a separate assessment of these rats' copulatory ability were significantly correlated with their instrumental performance. In a second experiment, "masculinized" females exposed to TP postnatally and given TP implants responded in extinction at mean levels equivalent to those exhibited by adult males that were either intact or castrated with androgen replacement. These data suggest that organizational steroid exposure perinatally affects the actual reward value assigned to oestrous females inadulthood, in combination withconsummatory sexual experience.     

Vocal changes in stutterers and nonstutterers during high- and low-pitched speech

This study was conducted to provide an indirect test of Wingate's “modified vocalization” hypothesis. In this formulation, the improved fluency that stutterers experience in various novel conditions is attributed to changes in the key correlates of stress, namely, fundamental frequency, vocal SPL, and rate.Normal speakers and stutterers read aloud in an habitual condition following instructions to read at higher- and lower-than-normal pitches. Objective measures were taken of subjects' fundamental frequency, fundamental frequency deviation, vocal SPL, and fluent reading rate. Disfluences were also counted. Findings showed that both stutterers and normals altered several features of voicing from the habitual to the two experimental conditions. Significant condition main effects emerged for fundamental frequency deviation, vocal SPL, fluent reading rate, and disfluency. The only meaningful between-group difference noted showed that the stutterers were more disfluent than the normals across all conditions. Results were interpreted as supporting Wingate's “modified vocalization” position and were discussed relative to how the vocal changes observed might act to promote fluency.     

Relationship between expressive language ability and rhythm perception, pitch perception, vocal range, and vocal midpoint among mentally retarded adults

This study investigated the relationship between expressive language ability and rhythm perception, pitch perception, vocal range, and vocal midpoint for 28 mentally retarded adults. A standardized speech test determined language scores, a standardized music test assessed rhythm and pitch perception, and vocalization determined vocal range and midpoint. Multiple correlation and multiple linear regression analyses indicated a significant correlation between expressive language ability and rhythm perception. Further research with music tests more appropriate for mentally retarded adults is recommended.     

Fixed-interval limited-hold avoidance with and without signalled reinforcement

Rats trained to lever press on a fixed-interval limited-hold avoidance schedule maintained a pattern of responding similar to that maintained by fixed-interval limited-hold schedules of positive reinforcement. But this positively accelerated pattern of behavior was maintained only when the occurrence of reinforcement was signalled by the presentation of a brief flash of light. This result suggests that the discriminative function of the reinforcer in avoidance is less pronounced than the discriminative function of the reinforcer in escape or positive reinforcement. It also suggests that the distinction between positive reinforcement and avoidance is not superfluous. Although the schedule of reinforcement is an important variable in determining the pattern of behavior, other variables, such as the nature (i.e., stimulus presentation, termination, or omission) of the reinforcer, are also potent determinants of behavior.