Intercurrent and reinforced behavior under multiple spaced-responding schedules

Lever pressing in rats was reinforced with food under a multiple spaced-responding schedule. A lever, food cup, and drinking tube were mounted in a running wheel so that lever pressing, running, and licking could be recorded. Running and licking had no scheduled consequences. Lever pressing was reinforced under a multiple schedule with three spaced-responding components and an extinction component. Each component was associated with a different auditory stimulus. Spaced-responding components reinforced only lever presses terminating interresponse times equal to or greater than 10, 20, or 60 sec, respectively. Rates of lever pressing, reinforcement, and licking all decreased as schedule parameter increased. Efficiency of spaced responding, as measured by reinforcements per response, also decreased. Rate of wheel running either increased or increased and then decreased with increasing schedule parameter. Individual running rates differed substantially. Neither licking nor running rate correlated with individual differences in efficiency. Analysis of conditional probabilities among the several response classes showed that, as the schedule requirement increased, the probability of running after a lever press increased and the probability of licking after a lever press decreased. After reinforcement, one subject always pressed the lever next. In the other subjects, the conditional probability of lever pressing, given reinforcement, increased while the probability of licking, given reinforcement, decreased with increasing schedule requirement. Results are discussed in relation to the concepts of schedule-induced and mediating behavior.     

The generality of selective observing

Four rats obtained food pellets by poking a key and 5-s presentations of the discriminative stimuli by pressing a lever. Every 1 or 2 min, the prevailing schedule of reinforcement for key poking alternated between rich (either variable-interval [VI] 30 s or VI 60 s) and lean (either VI 240 s, VI 480 s, or extinction) components. While the key was dark (mixed-schedule stimulus), no exteroceptive stimulus indicated the prevailing schedule. A lever press (i.e., an observing response), however, illuminated the key for 5 s with either a steady light (S+), signaling the rich reinforcement schedule, or a blinking light (S-), signaling the lean reinforcement schedule. One goal was to determine whether rats would engage in selective observing (i.e., a pattern of responding that maintains contact with S+ and decreases contact with S-). Such a pattern was found, in that a 5-s presentation of S+ was followed relatively quickly by another observing response (which likely produced another 5-s period of S+), whereas exposure to S- resulted in extended breaks from observing. Additional conditions demonstrated that the rate of observing remained high when lever presses were effective only when the rich reinforcement schedule was in effect (S+ only condition), but decreased to a low level when lever presses were effective only during the lean reinforcement component (S- only condition) or when lever presses had no effect (in removing the mixed stimulus or presenting the multiple-schedule stimuli). These findings are consistent with relativistic conceptualizations of conditioned reinforcement and extend the generality of selective observing to procedures in which the experimenter controls the duration of stimulus presentations, the schedule components both offer intermittent food reinforcement, and rats serve as subjects.     

Is induction produced by upcoming food-pellet reinforcement the outcome of an increase in overall activity or in operant responding?

Researchers have demonstrated that rats reliably increase their rates of pressing a lever for 1% liquid-sucrose reinforcement if they will soon have the opportunity to press a lever for food-pellet reinforcement. In the present experiments, the authors investigated if this increase in response rates occurred because the upcoming food pellets produced an increase in all behaviors (i.e., general arousal) or an increase in only the specific operant response (i.e., lever pressing). The results of Experiments 1 and 2 showed that the appearance of induction in rats' lever pressing for 1% sucrose reinforcement when food-pellet reinforcement was upcoming did not coincide with increases in the frequency of running in a wheel or making a nonreinforced nose-poke response. On the other hand, in Experiment 3, the authors found the appearance of induction coincided with increase nonreinforced lever presses on an adjacent lever. These results shed doubt on the idea that induction is a result of a general increase in all activity, and suggest instead that the increase in responding that occurs during induction is limited to the operant response.     

Techniques for establishing schedules with wheel running as reinforcement in rats.

In three experiments, access to wheel running was contingent on lever pressing. In each experiment, the duration of access to running was reduced gradually to 4, 5, or 6 s, and the schedule parameters were expanded gradually. The sessions lasted 2 hr. In Experiment 1, a fixed-ratio 20 schedule controlled a typical break-and-run pattern of lever pressing that was maintained throughout the session for 3 rats. In Experiment 2, a fixed-interval schedule of 6 min maintained lever pressing throughout the session for 3 rats, and for 1 rat, the rate of lever pressing was positively accelerated between reinforcements. In Experiment 3, a variable-ratio schedule of 20 or 35 was in effect and maintained lever pressing at a very stable pace throughout the session for 2 of 3 rats; for 1 rat, lever pressing was maintained at an irregular rate. When the session duration was extended to successive 24-hr periods, with food and water accessible in Experiment 3, lever pressing settled into a periodic pattern occurring at a high rate at approximately the same time each day. In each experiment, the rats that developed the highest local rates of running during wheel access also maintained the most stable and highest rates of lever pressing.     

Running and responding reinforced by the opportunity to run: effect of reinforcer duration.

The present study investigated the effect of reinforcer duration on running and on responding reinforced by the opportunity to run. Eleven male Wistar rats responded on levers for the opportunity to run in a running wheel. Opportunities to run were programmed to occur on a tandem fixed-ratio 1 variable-interval 30-s reinforcement schedule. Reinforcer duration varied across conditions from 30 to 120 s. As reinforcer duration increased, the rates of running and lever pressing declined, and latency to lever press increased. The increase in latency to respond was consistent with findings that unconditioned inhibitory aftereffects of reinforcement increase with reinforcer magnitude. The decrease in local lever-pressing rates, however, was inconsistent with the view that response strength increases with the duration of the reinforcer. Response rate varied inversely, not directly, with reinforcer duration. Furthermore, within-session data challenge satiation, fatigue, and response deprivation as determinants of the observed changes in running and responding. In sum, the results point to the need for further research with nonappetitive forms of reinforcement.     

Topography of signal-centered behavior in the rat: Effects of deprivation state and reinforcer type

In a series of three experiments, groups of food-deprived and water-deprived rats were given pairings of a retractable lever (CS⁺) with response-independent deliveries of either solid or liquid reinforcers. In Experiment 1 food-deprived rats given a solid-pellet reinforcer differentially tended to sniff, paw, mouth, and bite the CS⁺ lever more often than a lever that was not paired with food (CS⁻), whereas food-deprived rats given a liquid reinforcer tended to differentially sniff, paw, and lick the CS⁺ lever. 23½-hour water-deprived rats given liquid reinforcers showed very little CS⁺ contact. In Experiment 2 increasing the severity of water deprivation from 23½ to 47½ hours significantly increased CS⁺ contact. In Experiment 3, subjects that were simultaneously food and water deprived and given a water reinforcer failed to exhibit differential CS⁺ contact, but subjects that were simultaneously food and water deprived and given a food reinforcer did acquire differential CS⁺-contact behavior. These results suggest that (a) even under a single motivational state the nature of signal-centered behavior can be determined by type of reinforcer, (b) although water reinforcement produces less signal contact than food reinforcement, this can be facilitated with more severe water-deprivation levels, and (c) high CS-contact rates using food reinforcement are not simply a product of reductions in body weight with food deprivation.     

Effects of pre-operant running and sucrose concentration on operant wheel running on a fixed interval schedule of reinforcement

Prior research proposed that temporal control over the pattern of operant wheel running on a fixed interval (FI) schedule of sucrose reinforcement is a function of automatic reinforcement generated by wheel running and the experimentally arranged sucrose reinforcement. Two experiments were conducted to assess this prediction. In the first experiment, rats ran for different durations (0, 30, 60, and 180 min) prior to a session of operant wheel running on a FI 120-s schedule. In the second experiment, the concentration of sucrose reinforcement on a FI 180-s schedule was varied across values of 0, 5, 15, and 25%. In Experiment 1, as the duration of pre-operant running increased, the postreinforcement pause before initiation of running lengthened while wheel revolutions in the latter part of the FI interval increased. In Experiment 2, wheel revolutions markedly increased then decreased to a plateau early in the FI interval. Neither manipulation increased temporal control of the pattern of wheel running. Instead, results indicate that operant wheel running is regulated by automatic reinforcement generated by wheel activity and an adjunctive pattern of running induced by the temporal presentation of sucrose. Furthermore, the findings question whether the sucrose contingency regulates wheel running as a reinforcing consequence.     

Effects of magnitude of food reinforcement on free-operant response rates

In Experiment 1 rats were trained to press a lever on a variable-ratio schedule of food presentation and were then exposed to progressively increasing magnitudes of food reinforcement. Response running rates (rates exclusive of the postreinforcement pause) were found to increase as a function of increasing reinforcement magnitudes. The effect of reinforcement magnitude on response rates inclusive of the postreinforcement pause, however, was less pronounced. Increases in the magnitude of reinforcement were also found to increase the length of the postreinforcement pause. Rats in Experiment 2 were trained to respond on a chained differential-reinforcement-of-low-rate variable-ratio schedule, and were exposed to increasing magnitudes of reinforcement as in Experiment 1. Response running rates increased in the variable-ratio component but decreased in the other component of the schedule. The results are discussed with reference to incentive accounts of reinforcement and the action of reinforcement on the response units generated by the operative contingencies.     

DISCRIMINATION OF VARIABLE SCHEDULES IS CONTROLLED BY INTERRESPONSE TIMES PROXIMAL TO REINFORCEMENT

In Experiment 1, food‐deprived rats responded to one of two schedules that were, with equal probability, associated with a sample lever. One schedule was always variable ratio, while the other schedule, depending on the trial within a session, was: (a) a variable‐interval schedule; (b) a tandem variable‐interval, differential‐reinforcement‐of‐low‐rate schedule; or (c) a tandem variable‐interval, differential‐reinforcement‐of‐high‐rate schedule. Completion of a sample‐lever schedule, which took approximately the same time regardless of schedule, presented two comparison levers, one associated with each sample‐lever schedule. Pressing the comparison lever associated with the schedule just presented produced food, while pressing the other produced a blackout. Conditional‐discrimination accuracy was related to the size of the difference in reinforced interresponse times and those that preceded it (predecessor interresponse times) between the variable‐ratio and other comparison schedules. In Experiment 2, control by predecessor interresponse times was accentuated by requiring rats to discriminate between a variable‐ratio schedule and a tandem schedule that required emission of a sequence of a long, then a short interresponse time in the tandem's terminal schedule. These discrimination data are compatible with the copyist model from Tanno and Silberberg (2012) in which response rates are determined by the succession of interresponse times between reinforcers weighted so that each interresponse time's role in rate determination diminishes exponentially as a function of its distance from reinforcement.     

Transfer of persistence to the acquisition of a new behaviour

Two experiments tested whether the degree of effort required for the reinforcement of one behaviour would affect the acquisition of a second behaviour. In the first experiment, rats were placed in a conditioning chamber and: (a) were required to press a lever for food pellets on a fixed ratio schedule, (b) received free presentation of the pellets, or (c) did not receive pellets. Next, all rats were rewarded for a new behaviour, round trips across the length of a runway. As predicted, the fixed-ratio group had the greatest shuttle rate. In the second experiment, two groups were required to press a lever, and the number of presses per pellet was varied. For two other groups not required to press the lever, the amount of food presented per approach to the feeder was varied. The greater required number of lever presses and the lesser number of pellets per approach to the feeder produced the higher subsequent shuttle rates. Two alternative explanations were compared: the degree of accustomed effort per reinforcer becomes a learned component of behaviour, or high effort increases the habituation of frustration-produced disruptive responses.     

Operant hoarding: a new paradigm for the study of self-control.

In the first of four experiments, rats were exposed to a modified multiple continuous reinforcement-extinction schedule during 15-min daily sessions. In one condition (saves condition) with the cuelight on, a single lever press produced a food pellet, briefly extinguished the cuelight, and started a clock. Saves (additional lever presses with interresponse times less than 1 s) produced an additional food pellet, briefly extinguished the cuelight, and restarted the interresponse time clock. The cuelight was extinguished 1 s after the last lever press and remained off during a 10-s period of extinction, during which no food pellets were delivered. In the other condition (savings account condition), the contingencies were the same except that the cuelight was extinguished and was not reilluminated after the initial lever press, and the delivery of all food pellets in the reinforcement component was delayed until the onset of extinction. In both conditions, rats made saves, but mean saves (total saves divided by the number of reinforcement components) were slightly reduced in the savings account condition. In Experiment 2, using six equally spaced 15-min sessions per day on alternate days, saves were either followed immediately with food and brief cuelight offset (saves condition) or were not reinforced at all. Mean saves were much greater when saves were reinforced. In Experiment 3, during 5-min daily sessions, saves earned a single pellet (savings account condition) or a number of pellets equal to the ordinal number of the lever press (interest condition). Rats made fewer mean saves, with little change in the food rate, when saves earned interest. In Experiment 4, the rats earned all their food in the operant situation during 24 daily 5-min sessions, these separated by 55-min intersession intervals during which no food was available; otherwise, the conditions were the same as in Experiment 3. In Experiment 4, the shift to interest for saves led to an increase in mean daily mean saves (total daily mean saves divided by the number of daily sessions) as well as to an increase in the number of food pellets delivered in each session. The results are discussed in terms of self-control and behavioral economics.     

Response persistence under ratio and interval reinforcement schedules

In Experiment 1, rats were exposed to progressive-ratio schedules of food reinforcement while other rats were exposed simultaneously to yoked-interval schedules that arranged equivalent interreinforcer intervals but required only a single response at the end of the interval for food delivery. In Experiment 2, a within-subject yoked-control procedure was employed in which pigeons were exposed to alternating sessions (one per day) of progressive-ratio schedules and yoked-interval schedules as described above. In both experiments, responding under the yoked-interval schedule persisted beyond the point at which responding under the progressive-ratio schedule had ceased. The progressive-ratio schedules controlled break-and-run distributions, and the yoked-interval schedules controlled more even distributions of responses in time. Response rates decreased and postreinforcement pauses increased over time within individual sessions under both schedules. The results suggest that responding maintained by interval schedules is more persistent than that maintained by ratio schedules. The limitations and implications of this conclusion are discussed in the context of other investigations of response strength and behavioral momentum.     

Effects of cocaine on fixed-interval responding reinforced by the opportunity to run

Rate-dependent drug effects have been observed for operant responding maintained by food, water, heat, light onset, electrical brain stimulation, shock-stimulus termination, and shock presentation. The present study sought to determine if the effects of cocaine on lever pressing maintained by the opportunity to run could also be described as rate dependent. Seven male Wistar rats were trained to respond on levers for the opportunity to run in a wheel. The schedule of reinforcement was fixed-interval 60 s, and the reinforcing consequence was the opportunity to run for 60 s. On this schedule, overall rates of responding were low, usually below six presses per minute, and pauses frequently exceeded the 60-s interval. Despite these differences, an overall scalloped pattern of lever pressing was evident for each rat. Doses of 1, 2, 4, 8, and 16 mg/kg cocaine were administered 10 min prior to a session. Only at the 16 mg/kg dose did the responding of the majority of rats change in a manner suggestive of a rate-dependent drug effect. Specifically, lower response rates at the beginning of the intervals increased and higher rates at the end of the intervals decreased, as indicated by the fact that slopes from the regression of drug rates on control rates decreased. These data provide tentative support for the generalization of rate-dependent effects to operant responding maintained by wheel running. Differences in the baseline performance maintained by wheel running compared to those for food and water point to the need for further experimentation before this effect can be firmly established.     

Food anticipation and lever-directed activities in rats

Three experiments are described which elaborate some of the conditions under which rats will contact and manipulate a periodically presented retractable lever. Experiment 1 demonstrated that (i) initial manipulative oral and manual contact with the lever was facilitated if the rat had previous experience of food delivery in the experimental chamber; (ii) persistence in contacting the lever on successive presentations was a function of whether food continued to be presented in the experimental environment; and (iii) food satiation significantly reduced the tendency of the rat to contact the lever even though an expectancy of food had previously been established under conditions of food deprivation. Experiment 2 suggested that the tendency to approach and contact the lever was in part a function of the local moment-to-moment conditional probability of food delivery. Experiment 3 found that the probability of contacting the lever was higher during presentation of an auditory CS signaling a high rate of food delivery than during stimuli signaling no food at all. These results are interpreted as suggesting that the food-signaling aspects of an appetitive CS and that CS's ability to generate signal-directed behaviors are experimentall separable properties.     

Economic substitutability of electrical brain stimulation, food, and water.

Concurrent variable-ratio schedules of electrical brain stimulation, food, and water were paired in various combinations as reinforcement of rats' lever presses. Relative prices of the concurrent reinforcers were varied by changing the ratio of the response requirements on the two levers. Economic substitutability, measured by the sensitivity of response ratio to changes in relative price, was highest with brain stimulation reinforcement of presses on both levers and lowest with food reinforcement of presses on one lever and water reinforcement of presses on the other. Substitutability with brain stimulation reinforcement of presses on one lever and either food or water reinforcement for presses on the other was about as high as with brain stimulation for presses on both levers. Electrical brain stimulation for rats may thus serve as an economic substitute for two reinforcers, neither of which is substitutable for the other.     

Determinants of reinforcer accumulation during an operant task.

Responses by rats on an earn lever made available food pellets that were delivered to a food cup by responses on a second, collect, lever. The rats could either collect and immediately consume or accumulate (defined as the percentage of multiple earn responses and as the number of pellets earned before a collect response) earned pellets. In Experiment 1, accumulation varied as a function of variations in the earn or collect response requirements and whether the earn and collect levers were proximal (31 cm) or distal (248 cm) to one another. Some accumulation occurred under all but one of the conditions, but generally was higher when the earn and collect levers were distal to one another, particularly when the earn response requirement was fixed-ratio (FR) 1. In Experiment 2, the contributions of responses and time to accumulation were assessed by comparing an FR 20 earn response requirement to a condition in which only a single earn response was required at the end of a time interval nominally yoked to the FR interval. When 248 cm separated the earn and collect levers, accumulation was always greater in the FR condition, and it was not systematically related to reinforcement rate. In Experiment 3, increasing the earn response requirement with a progressive-ratio schedule that reset only with a collect response increased the likelihood of accumulation when the collect and earn levers were 248 cm apart, even though such accumulation increased the next earn response requirement. Reinforcer accumulation is an understudied dimension of operant behavior that relates to the analysis of such phenomena as hoarding and self-control, in that they too involve accumulating versus immediately collecting or consuming reinforcers.     

Reinforcement of inhibition

A differential-reinforcement-of-other-behavior (DRO) schedule with trials and delayed reinforcement was investigated. Periodically a wheel was briefly available to rats, followed six seconds later by brief availability of a bar. Variable-ratio food reinforcement of wheel turns was adjusted to give 95% turns. After variable-ratio-five reinforcement of bar presses produced 100% pressing, then separate ratio schedules were used for presses following turns (turn presses) and presses following nonturns (nonturn presses). Increasing nonturn-press reinforcements decreased turns, even though total reinforcements increased. Reversal by decreasing nonturn-press reinforcements raised turns, though with hysteresis. Thus food reinforcement increased nonturns even though delayed six to ten seconds after nonturns, a delay that greatly reduces response reinforcement. Those and other results indicate that the turn decrease was not due to reinforcement of competing responses. Evidence against other alternatives, and the reduction of responding by increased reinforcement, indicate that the term inhibition is appropriate for the phenomenon reinforced. Response-specific inhibition appears appropriate for this particular kind, since its effects are more specific to particular responses than Pavlovian conditioned-inhibition. Response-specific inhibition seems best considered a behavioral output comparable to responses (e.g., both reinforcible) but with important properties different from responses (e.g., different reinforcement-delay gradients).     

Preadaptation to the feeding schedule does not eliminate activity-based anorexia in rats

To test whether activity-based anorexia (ABA) still occurs after preadaptation to the feeding schedule, 20 rats were first exposed to a feeding schedule of one 90-min meal per day until adaptation occurred (measured by maintenance of stable body weight). Then, during ABA training, half the rats (wheel group) were confined in running wheels except during the daily meal, and half (cage group) were not. Wheel running suppressed feeding—that is, food intake in the wheel group was less than that in the cage group. Also, the rats in the wheel group lost weight, whereas those in the cage group did not. Wheel running increased over days. Thus, the defining characteristics of ABA were evident in rats that were not subjected to ABA training until after they had become well adapted to the feeding schedule. These findings support the view that the suppression of feeding produced by wheel running triggers the vicious circle of ABA. They also cast doubt on the hypothesis that activity-induced interference with adaptation to the feeding schedule plays a key role in causing ABA.     

Instrumental performance on negative schedules

Three experiments examined the performance of rats pressing a lever for food reinforcement on a schedule in which high rates of response resulted in lowered rates of reinforcement (i.e. a schedule with a negative component). In Experiment 1, rats responded on a variable interval (VI) schedule with a conjoint component such that every 30 responses a reinforcement programmed by the VI schedule was cancelled. These subjects generally emitted a lower response rate than rats responding on a VI schedule yoked to the former subjects with respect to the delivery of reinforcement, although response rate differences were sometimes not large. Similar response-rate effects were obtained in Experiment 2 using a within-subject yoking procedure. In Experiment 3, reinforced interresponse times were matched on negative and VI schedules yoked in terms of reinforcement rate, and the response rate emitted in these conditions were similar. These results give support to theories of instrumental conditioning that stress the strengthening and shaping properties of reinforcement.     

Resistance to the response-decrementing effects of response-independent reinforcement produced by delay and non-delay schedules of reinforcement

In two experiments, animals were initially exposed to response-dependent schedules of food before exposure to response-independent reinforcement matched for overall rate and temporal distribution of reinforcers to the preceding condition. In Experiment I, response decrements during the response-independent phase were smaller after delayed reinforcement training than after a comparable immediate reinforcement schedule, for both doves and rats. In Experiment II variable-interval and variable-ratio schedules, both with either immediate or delayed reinforcement, were used with rats. Both the delayed reinforcement schedules produced resistance to subsequent response-independent reinforcement, but response decrements were larger after either of the immediate reinforcement conditions. It was concluded that the critical factor in response maintenance under response-independent reinforcement was the type of response-reinforcer contiguities permitted under the response-dependent schedule rather than perception of response-reinforcer “contingencies”. If the response-dependent schedule was arranged so that behaviours other than a designated operant (key pecking or lever pressing) could be contiguous with food, responding was maintained well under response-independent schedules.