Acquisition of lever-press responding in rats with delayed reinforcement: A comparison of three procedures

The present study examined the acquisition of lever pressing in rats under three procedures in which food delivery was delayed by 4, 8, and 16 seconds relative to the response. Under the nonresetting delay procedure, food followed the response selected for reinforcement after a specified interval elapsed; responses during this interval had no programmed effect. Under the resetting procedure, the response selected for reinforcement initiated an interval to food delivery that was reset by each subsequent response. Under the stacked delay procedure, every response programmed delivery of food t seconds after its occurrence. Two control groups were studied, one that received food immediately after each lever press and another that never received food. With the exception of the group that did not receive food, responding was established with every procedure at every delay value without autoshaping or shaping. Although responding was established under the resetting delay procedure, response rates were generally not as high as under the other two procedures. These findings support the results of other recent investigations in demonstrating that a response not previously reinforced can be brought to strength by delayed reinforcement in the absence of explicit training.     

Control of responding by sounds of different quality: an evolutionary analysis.

Two experiments investigated the acquisition of discriminations between two acoustic stimuli of different quality (noise bursts vs. a 2-kHz pulsed signal) when features of the everyday environment were incorporated into the experiments. In Experiment 1, rats were trained, using food, to press a lever. Throughout all sessions, 5-s trials of noise bursts (the random stimulus) were presented, after variable intertrial intervals, through a remote speaker mounted outside the experimental enclosure. The noise burst occurred randomly with respect to reinforcement of lever pressing and had no programmed relationship to the animal's behavior. When lever pressing was established, the 2-kHz signal was presented through a speaker adjacent to the response lever according to a different set of variable intertrial intervals. A response in the presence of the 2-kHz signal terminated the trial and was reinforced. The 2-kHz signal acquired control of responding within the first few trials, whereas the random stimulus exerted no control of responding. In Experiment 2, rats were trained to press the lever in the presence of the 2-kHz signal presented through the adjacent speaker on a variable intertrial interval. After 14 sessions, 5-s trials of noise bursts (random stimulus) were presented through the remote speaker on the second variable intertrial interval. The random stimulus initially elicited exploratory behavior, which then rapidly declined. Subsequently, the random stimulus exerted no or weak control of responding. The introduction of the random stimulus had no effect on responding in the presence of the adjacent stimulus. In Experiments 3 and 4 the random stimulus was presented through the adjacent speaker, and the stimulus correlated with reinforcement was presented through the remote speaker. In both experiments, there was persistent control of responding by the random stimulus and slow development of control by the stimulus correlated with reinforcement. In Experiment 5, both stimuli were presented through the adjacent speaker. There was persistent control of responding by the random stimulus.     

Effects of reinforcement rate and delay on the acquisition of lever pressing by rats.

The acquisition of lever pressing by naive rats, in the absence of shaping, was studied as a function of different rates and unsignaled delays of reinforcement. Groups of 3 rats were each exposed to tandem schedules that differed in either the first or the second component. First-component schedules were either continuous reinforcement or random-interval 15, 30, 60 or 120 s; second-component schedules were fixed-time 0, 1, 3, 6, 12, or 24 s. Rate of responding was low under continuous immediate reinforcement and higher under random-interval 15 s. Random interval 30-s and 60-s schedules produced lower rates that were similar to each other. Random-interval 120 s controlled the lowest rate in the immediate-reinforcement condition. Adding a constant 12-s delay to each of the first-component schedule parameters controlled lower response rates that did not vary systematically with reinforcement rate. The continuous and random-interval 60-s schedules of immediate reinforcement controlled higher global and first-component response rates than did the same schedules combined with longer delays, and first-component rates showed some graded effects of delay duration. In addition, the same schedules controlled higher second-component response rates in combination with a 1-s delay than in combination with longer delays. These results were related to those from previous studies on acquisition with delayed reinforcement as well as to those from similar reinforcement procedures used during steady-state responding.     

Force and rate relations in responding during variable-interval reinforcement

Four rats responded on one-minute variable-interval schedules with several variations in peak-force of response required for food reinforcement. Measures of peak force and rate were taken for the responses, which were the downward exertions of force against a static force-transducing operandum. The analysis distinguished responses, a generic class of measured behavior, from criterion responses, an operationally specified subclass required for reinforcement. Absolute rate of response showed no systematic change, but the rate of responses meeting a newly required criterion of peak-force invariably increased through changes in the absolute rate of response, the relative-frequency distributions of peak force, or some combination of both. The relative frequency of responses meeting an elevated force criterion during variable-interval reinforcement exceeded that maintained with the same criterion with continuous reinforcement. The requirement of more effortful responding for reinforcement does not necessarily reduce response rate. Conformity of the behavior to the requirement for reinforcement is the salient effect.     

Social reinforcement of operant behavior in rats: a methodological note.

An apparatus was developed to study social reinforcement in the rat. Four Long-Evans female rats were trained to press a lever via shaping, with the reinforcer being access to a castrated male rat. Responding under a fixed-ratio schedule and in extinction was also observed. Social access was found to be an effective reinforcer. When social reinforcement was compared with food reinforcement under similar conditions of deprivation and reinforcer duration, no significant differences were observed.     

Control of responding by the location of sound: role of binaural cues.

In auditory localization experiments, where the subject observes from a fixed position, both relative sound intensity and arrival time at the two ears determine the extent of localization performance. The present experiment investigated the role of binaural cues in a different context, the sound-position discrimination task, where the subject is free to move and interact with the sound source. The role of binaural cues was investigated in rats by producing an interaural imbalance through unilateral removal of the middle auditory ossicle (incus) prior to discrimination training. Discrete trial go-right/go-left sound-position discrimination of unilaterally incudectomised rats was then compared with that of normal rats and of rats with the incus of both sides removed. While bilateral incus removal affected binaural intensity and arrival times, the symmetry of sound input between the two ears was preserved. Percentage of correct responses and videotaped observations of sound approach and exploration showed that the unilateral rats failed to localize the sounding speaker. Rats with symmetrical binaural input (normal and bilaterally incudectomised rats) accurately discriminated sound position for the duration of the experiment. Previously reported monaural localization based upon following the intensity gradient to the sound source was not observed in the unilaterally incudectomised rats of the present experiment. It is concluded that sound-position discrimination depends upon the use of binaural cues.     

Second-order schedules of token reinforcement: effects of varying the schedule of food presentation

In the initial link of a complex schedule, one discriminative stimulus was presented and lever pressing produced tokens on fixed-ratio schedules. In the terminal link, signalled by a second discriminative stimulus, deposits of the tokens produced food. With two rats, the terminal link was presented after each sixth component schedule of token reinforcement was completed. With the other two rats, the terminal link was presented following the first component schedule completed after a fixed interval. During the terminal link, each token deposit initially produced food. The schedule of food presentation was subsequently increased such that an increasing number of token deposits in the terminal link was required for each food presentation. Rates of lever pressing in the initial link were inversely related to the schedule of food presentation in the terminal link. These results are similar to those of experiments that have varied schedules of food presentation in chained schedules. Rates and patterns of responding controlled throughout the initial link were more similar to those ordinarily controlled by second-order brief-stimulus schedules than to those controlled by comparable extended chained schedules.     

The effects of d-amphetamine on the temporal control of operant responding in rats during a preshock stimulus.

The operant behavior of six rats was maintained by a random-interval schedule of reinforcement. Three-minute periods of noise were superimposed on this behavior, each period ending with the delivery of an unavoidable shock. Overall rates of responding were generally lower during the periods of noise than in its absence (conditioned suppression). These suppressed response rates also exhibited temporal patterning, with responding becoming less frequent as each noise period progressed. The effects of d-amphetamine on this behavioral baseline were then assessed. In four animals the relative response rates during the noise and in its absence suggested that the drug produced a dose-related decrease in the amount of conditioned suppression. However, this effect was often due to a decrease in the rates of responding in the absence of the preshock stimulus, rather than to an increase in response rates during the stimulus. Temporal patterning in response rates during the preshock stimulus was abolished, an effect that was interpreted in terms of rate-dependent effect of d-amphetamine. This study thus extends rate-dependent analyses of the effects of amphetamines to the patterns of operant behavior that occur during a preshock stimulus, and which have been discussed in terms of the disrupting effects of anxiety on operant behavior.     

Relationship between response rate and reinforcement frequency in variable-interval schedules: III. The effect of d-amphetamine.

Four rats were exposed to variable-interval schedules specifying a range of different reinforcement frequencies. The effects of two doses of d-amphetamine (1.6 and 3.2 mumol/kg) upon performance maintained under each schedule were examined. In the case of each rat, the response rates observed under control conditions (no injection or injection of the vehicle alone) were increasing, negatively accelerated functions of reinforcement frequency, the data conforming closely to Herrnstein's (1970) equation. In each rat, d-amphetamine (3.2 mumol/kg) significantly reduced the value of the constant Rmax, which expresses the theoretical maximum response rate. In each rat, the value of KH, which expresses the reinforcement frequency needed to obtain the half-maximal response rate, was also reduced, although this only achieved statistical significance in the case of one rat. When the proportional change in response rate in the presence of the drug was plotted against the response rate under control conditions on double logarithmic co-ordinates, linear functions of negative slope were obtained; in each rat the slope was steeper and the value of the control response rate at which d-amphetamine exerted no effect was lower in the case of the higher dose (3.2 mumol/kg) than in the case of the lower dose (1.6 mumol/kg).     

The effects of schedule history and the opportunity for adjunctive responding on behavior during a fixed-interval schedule of reinforcement.

The effects of schedule history and the availability of an adjunctive response (polydipsia) on fixed-interval schedule performance were investigated. Two rats first pressed levers under a schedule of food reinforcement with an interresponse time greater than 11 s, and 2 others responded under a fixed-ratio 40 schedule. All 4 were then exposed to a fixed-interval 15-s schedule. Water was continuously available under these conditions, but after responding became stable on the fixed-interval schedule, access was experimentally manipulated. With water freely available, subjects did not display characteristic fixed-interval response rates and patterns (i.e., scalloping or break-and-run). Instead, they exhibited predictable, stable patterns of behavior as a function of their schedule histories: Subjects with the interresponse-time history exhibited low response rates, and those with the fixed-ratio history exhibited high rates. Manipulating the amount of water available resulted in marked changes in response rates for rats with the interresponse-time history but not for those with the fixed-ratio history. The results illustrate the multiple causation of behavior by its previous and current schedules of reinforcement and other concurrent factors.     

Effects of food deprivation and reinforcement magnitude on conditioned suppression

In Experiment I, the responding of rats lever pressing on a variable-interval schedule for sucrose solution was partially suppressed by a variable duration conditioned stimulus followed by shock. When food deprivation was increased, response rates during and before the conditioned stimulus increased monotonically. Varying the concentration of sucrose across blocks of sessions or from session to session in a semi-random sequence had little effect on response rates either before or during the conditioned stimulus. With a fixed sequence of increasing concentrations across a five-session block, increased concentration produced much more rapid increases in response rates before than during the conditioned stimulus. In Experiment II, rats were presented with the same sequence of increasing concentrations across a five-session block. When tested at 80% body weight, response rates increased rapidly as concentration increased, but at 100%, body-weight rates increased only slightly. The effect of a change in body weight in Experiment II thus mimicked the effect of the conditioned stimulus in the latter part of Experiment I. These findings support the view that the effect of a pre-aversive conditioned stimulus is similar to that of a change in food deprivation, but unlike that of a change in reinforcement magnitude.     

The roles of stimulus control and reinforcement frequency in modulating the behavioral effects of d-amphetamine in the rat.

The behavioral effects of d-amphetamine have been shown to be modulated by stimulus control, with less impairment of performance occurring when control is great. When the fixed-consecutive-number schedule is used (on which at least a specified consecutive number of responses must be made on one operandum before a single response on another will produce a reinforcer), response rate tends to be invariant but reinforcement frequency is not. This study asks whether the differences in reinforcement frequency that usually accompany changes in stimulus control could themselves be responsible for the performance differences. Two versions of the fixed-consecutive-number schedule of reinforcement were combined into a multiple schedule within which stimulus control was varied but differences in reinforcement frequency were minimized by omitting some reinforcer deliveries during the component that usually had the higher reinforcement frequency. In one component, a compound discriminative stimulus was added with the eighth consecutive response on the first lever; a single response on the second lever was then reinforced. In the other component, no such stimulus was presented. With no added stimulus, large decreases occurred in the number of runs satisfying the minimum requirement for reinforcement at doses of drug that produced only minimal changes when an added stimulus controlled behavior. Thus, increased stimulus control diminishes the behavioral changes produced by d-amphetamine even when the possible contribution by baseline reinforcement rate is minimized.     

Simultaneous auditory discrimination.

Stimuli in many visual stimulus control studies typically are presented simultaneously; in contrast the stimuli in auditory discrimination studies are presented successively. Many everyday auditory stimuli that control responding occur simultaneously. This suggests that simultaneous auditory discriminations should be readily acquired. The purpose of the present experiment was to train rats in a simultaneous auditory discrimination. The apparatus consisted of a cage with two response levers mounted on one wall and a speaker mounted adjacent to each lever. A feeder was mounted on the opposite wall. In a go-right/go-left procedure, two stimuli were presented on each trial, a wide-band noise burst through one speaker and a 2-kHz complex signal through the other. The stimuli alternated randomly from side to side across trials, and the stimulus correlated with reinforcement for presses varied across subjects. The rats acquired the discrimination in 400 to 700 trials, and no response position preference developed during acquisition. The ease with which the simultaneous discrimination was acquired suggests that procedures, such as matching to sample, that require simultaneous presentation of stimuli can be used with auditory stimuli in animals having poor vision.     

Relationship between response rate and reinforcement frequency in variable-interval schedules: the effect of the concentration of sucrose reinforcement

Four rats were exposed to variable-interval schedules specifying a range of different reinforcement frequencies, using sucrose of two different concentrations and distilled water as the reinforcer. With sucrose, the rates of responding of all four rats were increasing negatively accelerated functions of reinforcement frequency, the data conforming closely to Herrnstein's equation; this was also true of the data from three of the four rats when distilled water was used as the reinforcer. The values of both constants in Herrnstein's equation were related to the sucrose concentration: the asymptotic response rate decreased, and the reinforcement frequency corresponding to the half-maximal response rate increased, with decreasing sucrose concentration.     

Effects of D-amphetamine on response acquisition with immediate and delayed reinforcement.

The present study examined in 8-hour sessions the effects of d-amphetamine (1.0, 5.6, and 10 mg/kg) on the acquisition of lever-press responding in rats that were exposed to procedures in which water delivery was delayed by 0, 8, or 16 seconds relative to the response that produced it. Both nonresetting- and resetting-delay conditions were studied. Although neither shaping nor autoshaping occurred, substantial levels of operative-lever responding developed under all conditions in which responses produced water. The lowest dose (1.0 mg/kg) of d-amphetamine either had no effect on or increased operative-lever pressing, whereas higher doses typically produced an initial reduction in lever pressing. Nonetheless, overall rates of operative-lever pressing at these doses were as high as, or higher than, those observed with vehicle. Thus, response acquisition was observed under all reinforcement procedures at all drug doses. In the absence of the drug, most responding occurred on the operative lever when reinforcement was immediate. Such differential responding also developed under both nonresetting- and resetting-delay procedures when the delay was 8 seconds, but not when it was 16 seconds. d-Amphetamine did not affect the development of differential responding under any procedure. Thus, consistent with d-amphetamine's effects under repeated acquisition procedures, the drug had no detrimental effect on learning until doses that produced general behavioral disruption were administered.     

KNOWING BEFORE DOING: DISCRIMINATION BY RATS OF A BRIEF INTERRUPTION OF A TONE

Eight rats' lever presses were reinforced after an interruption in a tone, provided the lever had not been pressed before the tone interruption. After a few sessions, long before the animals reliably refrained from lever pressing before the interruption, the latencies of postinterruption presses (time from the termination of the interruption to the moment of the lever press) dissociated into two classes: short ones for to-be-rewarded presses, and long ones for presses in the other trials, which contained no reward because one or more lever presses had occurred before the interruption. Thus discrimination of impending reinforcement in the current trial occurred before there was evidence of sensitivity to reinforcement in the reinforcement-producing aspect of behavior. This finding is related to Shimp's (1981) contention that the temporal properties of recent behavior are reinforceable, if remembered. The present finding shows that learning to discriminate whether one's behavior has met a contingency, and learning to carry out this behavior, need not go together, implying that memory of temporal properties is probably a necessary but not a sufficient condition for learning the latter.     

The effects of reinforcement frequency and response requirements on the maintenance of behavior.

Six rats responded under fixed-interval and tandem fixed-interval fixed-ratio schedules of food reinforcement. Basic fixed-interval schedules alternated over experimental conditions with tandem fixed-interval fixed-ratio schedules with the same fixed-interval value. Fixed-interval length was varied within subjects over pairs of experimental conditions; the ratio requirement of the tandem schedules was varied across subjects. For both subjects with a ratio requirement of 10, overall response rates and running response rates typically were higher under the tandem schedules than under the corresponding basic fixed-interval schedules. For all subjects with ratio requirements of 30 or 60, overall response rates and running response rates were higher under the tandem schedules than under the corresponding basic fixed-interval schedules only with relatively short fixed intervals. At longer fixed intervals, higher overall response rates and running rates were maintained by the basic fixed-interval schedules than by the tandem schedules. These findings support Zeiler and Buchman's (1979) reinforcement-theory account of response strength as an increasing monotonic function of both the response requirement and reinforcement frequency. Small response requirements added in tandem to fixed-interval schedules have little effect on reinforcement frequency and so their net effect is to enhance responding. Larger response requirements reduce reinforcement frequency more substantially; therefore their net effect depends on the length of the fixed interval, which limits overall reinforcement frequency. At the longest fixed intervals studied in the present experiment, reinforcement frequency under the tandem schedules was sufficiently low that responding weakened or ceased altogether.     

Control rate of response or reinforcement and amphetamine's effect on behavior.

The roles of control response rate and reinforcement frequency in producing amphetamine's effect on operant behavior were evaluated independently in rats. Two multiple schedules were arranged in which one variable, either response rate or reinforcement frequency, was held constant and the other variable manipulated. A multiple differential-reinforcement-of-low-rate seven-second yoked variable-interval schedule was used to equate reinforcement frequencies at different control response rates between multiple-schedule components. Amphetamine increased responding under the variable-interval component. In contrast, amphetamine decreased responding equivalently between components of a multiple random-ratio schedule that produced similar control response rates at different reinforcement frequencies. The results provide experimental support to the rate-dependency principle that control rate of responding is an important determinant of amphetamine's effect on operant behavior.     

Patterns of responding within sessions.

Rates of responding changed systematically across sessions for rats pressing levers and keys and for pigeons pressing treadles and pecking keys. A bitonic function in which response rates increased and then decreased across sessions was the most common finding, although an increase in responding also occurred alone. The change in response rate was usually large. The function relating responding to time in session had the following general characteristics: It appeared early in training, and further experience moved and reduced its peak; it was flatter for longer sessions; and it was flatter, more symmetrical, and peaked later for lower than for higher rates of reinforcement. Factors related to reinforcement exerted more control over the location of the peak rate of responding and the steepness of the decline in response rates than did factors related to responding. These within-session changes in response rates have fundamental theoretical and methodological implications.     

Food and intracranial stimulation responding suppressed with regular-interval shock.

Attenuation of conditioned suppression during intracranial stimulation was compared with that during food reinforcement. Response rates controlled by food and by brain stimulation were equalized on a multiple schedule by adjusting the stimulating current. When foot shock was delivered during timeout periods separating response components, responding for food was significantly more suppressed than responding for brain stimulation. When components were shortened from 10 to 2 minutes, responding maintained by either food or brain stimulation showed a similar temporal pattern of suppression preceding each shock, but responding in the component involving food remained significantly more suppressed. Explanations for the attenuated suppression during brain stimulation based on neural disruption, stimulus blocking, and analgesic properties were questioned. The increased responding during brain stimulation seemed to reflect greater response strength relative to food reinforced responding.