Operant behavior in dwarf hamsters (Phodopus campbelli): Effects of rate of reinforcement and reinforcer flavor variety

We investigated operant behavior in a novel species, the dwarf hamster (Phodopus campbelli). In two experiments, hamsters were trained to lever-press for food reinforcement. In Experiment 1, rate of reinforcement was manipulated across conditions using four variable-interval schedules of reinforcement (delivering one to eight reinforcers per min). As predicted, within-session decreases in responding were steepest on the richest schedule. In Experiment 2, lever-pressing was reinforced by either a constant or a variety of flavored food pellets. Within-session decreases in responding were steeper when the reinforcer flavor remained constant than when it was varied within the session. In both experiments, subjects hoarded most reinforcers in their cheek pouches rather than consuming them in the operant chambers. These results are incompatible with post-ingestive satiety variables as explanations for within-session decreases in operant responding and suggest that habituation to repeatedly presented reinforcers best accounts for subjects’ response patterns. Additionally, a mathematical model that describes behavior undergoing habituation also described the present results, thus strengthening the conclusion that habituation mediates the reinforcing efficacy of food.     

Mathematical principles of reinforcement and resistance to change

Although Killeen's mathematical principles of reinforcement (MPR) apply to the asymptotic rate of a free operant after extended exposure to a single schedule of reinforcement, they can be extended to resistance to change in multiple schedules via alterations in the parameter representing the activating effects of reinforcers. MPR's predictions of resistance to change in relation to reinforcer rate on variable-interval (VI) schedules are empirically correct and agree with behavioral momentum theory (BMT). However, both MPR and BMT encounter problems in accounting for the effects of delayed reinforcement on resistance to change, relative to immediate reinforcement at the same rate. Further problems are raised by differences in resistance to change between variable-ratio (VR) and variable-interval performances maintained by the same reinforcer rate. With both delayed versus immediate reinforcement and variable-ratio versus variable-interval reinforcement, differential resistance to change is negatively correlated with the log ratios of baseline response rates when reinforcer rates are equated. Cases where resistance to change varies despite equated reinforcer rates, and the correlations among behavioral measures, provide challenges and opportunities for both MPR and BMT.     

Segmentation and the pairing hypothesis

The effect of stimulus contiguity and response contingency on responding in chain schedules was examined in two experiments. In Experiment 1, four pigeons were trained on two simple three-link chain schedules that alternated within sessions. Initial links were correlated with a variable-interval 30s schedule, and middle and terminal links were correlated with interdependent variable-interval 30s variable-interval 30s schedules. The combined duration of the interdependent schedules summed to 60s. The two chains differed with respect to signaling of the schedule components: a two-stimulus chain had one stimulus paired with the initial link and one stimulus paired with both the middle and the terminal link, while a three-stimulus chain had a different stimulus paired with the each of the three links. The results showed that the two-stimulus chain maintained lower initial-link responding than the three-stimulus chain. In Experiment 2, four pigeons were exposed to three separate conditions, the two- and three-stimulus chains of Experiment 1 and a three-stimulus chain that had a 3s delay to terminal-link entry from the middle-link response that produced it. The two-stimulus chain maintained lower initial-link responding than the three-stimulus chain, as in Experiment 1, and a similar initial-link responding was maintained by the two-stimulus chain and the three-stimulus chain with the delay contingency. The results demonstrate that a stimulus noncontiguous with food can maintain responding that is sometimes greater than a stimulus contiguous with food, depending on the response contingency for terminal-link entry. The results are contrary to the pairing hypothesis of conditioned reinforcement.     

Reinforcer satiation and resistance to change of responding maintained by qualitatively different reinforcers

In previous research on resistance to change, differential disruption of operant behavior by satiation has been used to assess the relative strength of responding maintained by different rates or magnitudes of the same reinforcer in different stimulus contexts. The present experiment examined resistance to disruption by satiation of one reinforcer type when qualitatively different reinforcers were arranged in different contexts. Rats earned either food pellets or a 15% sucrose solution on variable-interval 60-s schedules of reinforcement in the two components of a multiple schedule. Resistance to satiation was assessed by providing free access either to food pellets or the sucrose solution prior to or during sessions. Responding systematically decreased more relative to baseline in the component associated with the satiated reinforcer. These findings suggest that when qualitatively different reinforcers maintain responding, relative resistance to change depends upon the relations between reinforcers and disrupter types.     

Differential resurgence and response elimination

Resurgence refers to the transient recovery of previously reinforced, but presently not reinforced, responding when more recently reinforced responding is extinguished. The primary purpose of our research was to determine how differential resurgence results from the procedures used to eliminate that responding. There were three conditions in each of five experiments. In Condition 1, key pecking by pigeons was maintained under a two-component multiple variable-interval (VI) 30-s VI 30-s schedule. In Condition 2, this pecking was eliminated in different ways across components. In Condition 3, extinction was in effect for all responses, and resurgence of key pecking was compared across components. These three conditions were repeated for most pigeons, and the procedures used to eliminate responding in Condition 2 varied across experiments. In Experiment 1, there was greater resurgence, and an earlier onset of it, after a differential-reinforcement-of-other-behavior (DRO) schedule than after a VI schedule was correlated with pecking an alternative key. Experiments 2 and 3 showed that the differential resurgence in Experiment 1 probably was not due to conditional stimulus control or the periodicity of food delivery, respectively. In Experiment 4, there was no systematic difference in resurgence after either a DRO schedule or a VI schedule correlated with treadle pressing. In Experiment 5, there was greater resurgence, and/or an earlier onset of it, after a VI schedule correlated with treadle pressing than after a VI schedule correlated with pecking an alternative key. Taken together, the results showed that the reinforcement of an alternative key-peck response was the most effective means of reducing subsequent key-peck resurgence. The relation of these results to an understanding of resurgence is discussed.     

The role of terminal-link stimuli in concurrent-chain schedules: revisited using a behavioral-history procedure

A behavioral-history procedure was used to study the function of terminal-link stimuli as conditioned reinforcers in multiple concurrent-chain schedules of reinforcement. First, three pigeons were exposed to multiple concurrent-chain schedules in which the two multiple-schedule components were correlated with a blue and a white stimulus, respectively. In each component the initial links were equal independent variable-interval (VI) 15 s schedules. A fixed-interval (FI) 10 s schedule operated on the red key in one terminal link while extinction operated on the green key in the alternative terminal link. When large preferences for the red stimulus had been established, two tests were conducted. In the terminal-link test, under new initial-link stimuli--purple and brown--an FI 10 s schedule operated for both the red and green terminal-link stimuli. In the subsequent initial-link test, the blue and white initial-link stimuli were reintroduced, and, as in the terminal-link test, FI 10s operated for both the red and the green terminal-link stimuli. In the terminal-link test, the three pigeons showed no preference for the terminal links with the red stimulus, but showed clear and consistent preferences for the red stimulus when blue and white stimuli were reintroduced as initial-link stimuli in the initial-link test. This suggests that there are multiple sources of control over initial-link response allocation in concurrent-chains, including control by both terminal- and initial-link stimuli.     

EMG Biofeedback: The Effects of CRF, FR, VR, FI, and VI Schedules of Reinforcement on the Acquisition and Extinction of Increases in Forearm Muscle Tension

Biofeedback was used to increase forearm-muscle tension. Feedback was delivered under continuous reinforcement (CRF), variable interval (VI), fixed interval (FI), variable ratio (VR), and fixed ratio (FR) schedules of reinforcement when college students increased their muscle tension (electromyograph, EMG) above a high threshold. There were three daily sessions of feedback, and Session 3 was immediately followed by a session without feedback (extinction). The CRF schedule resulted in the highest EMG, closely followed by the FR and VR schedules, and the lowest EMG scores were produced by the FI and VI schedules. Similarly, the CRF schedule resulted in the greatest amount of time-above-threshold and the VI and FI schedules produced the lowest time-above-threshold. The highest response rates were generated by the FR schedule, followed by the VR schedule. The CRF schedule produced relatively low response rates, comparable to the rates under the VI and FI schedules. Some of the data are consistent with the partial-reinforcement-extinction effect. The present data suggest that different schedules of feedback should be considered in muscle-strengthening contexts such as during the rehabilitation of muscles following brain damage or peripheral nervous-system injury.     

Effects of a parasitic nematode on male mate choice in a livebearing fish with a coercive mating system (western mosquitofish, Gambusia affinis)

Studies of choice in steady state have shown that sensitivity to reinforcement increases with increasing fixed-ratio changeover (FR CO) requirements. We assessed the generality of this finding with choice in transition. Food deliveries were programmed according to concurrent variable-interval (VI) schedules. Seven different VI pairs arranged ratios of food deliveries (left/right) of 27:1, 9:1, 3:1, 1:1, 1:3, 1:9, and 1:27 at a constant overall rate across components. Within sessions, all seven ratios were presented in random order. Each component lasted for 10 food deliveries; components were separated by 60-s blackouts. A changeover lever required 1, 2, 4, 8, 16, 32, and 64 responses to alternate between two main levers. Redeterminations to all FR COs, but 64 responses, were obtained in descending order. Choice adjusted rapidly to rapid changes in the reinforcer ratio, tracking the lever associated with the highest probability of reinforcer. Sensitivity to reinforcement increased with increasing FR CO, replicating the negatively accelerated function found in our earlier study. With successive reinforcers in components, however, sensitivity reached asymptote values sooner with the largest (8, 16, and 32 responses), than with the smallest (1, 2, and 4 responses), FR CO requirements.     

Response-cost punishment via token loss with pigeons

Two experiments were conducted to investigate punishment via response-contingent removal of conditioned token reinforcers (response cost) with pigeons. In Experiment 1, key pecking was maintained on a two-component multiple second-order schedule of token delivery, with light emitting diodes (LEDs) serving as token reinforcers. In both components, responding produced tokens according to a random-interval 20-s schedule and exchange periods according to a variable-ratio schedule. During exchange periods, each token was exchangeable for 2.5-s access to grain. In one component, responses were conjointly punished according to fixed-ratio schedules of token removal. Response rates in this punishment component decreased to low levels while response rates in the alternate (no-punishment) component were unaffected. Responding was eliminated when it produced neither tokens nor exchange periods (Extinction), but was maintained at moderate levels when it produced tokens in the signaled absence of food reinforcement, suggesting that tokens served as effective conditioned reinforcers. In Experiment 2, the effect of the response-cost punishment contingency was separated from changes in the density of food reinforcement. This was accomplished by yoking either the number of food deliveries per component (Yoked Food) or the temporal placement of all stimulus events (tokens, exchanges, food deliveries) (Yoked Complete), from the punishment to the no-punishment component. Response rates decreased in both components, but decreased more rapidly and were generally maintained at lower levels in the punishment component than in the yoked component. In showing that the response-cost contingency had a suppressive effect on responding in addition to that produced by reductions in reinforcement density, the present results suggest that response-cost punishment shares important features with other forms of punishment.     

Response-reinforcer relations and resistance to change

Behavioral momentum theory suggests that the relation between a response and a reinforcer (i.e., response-reinforcer relation) governs response rates and the relation between a stimulus and a reinforcer (i.e., stimulus-reinforcer relation) governs resistance to change. The present experiments compared the effects degrading response-reinforcer relations with response-independent or delayed reinforcers on resistance to change in conditions with equal stimulus-reinforcer relations. In Experiment 1, pigeons responded on equal variable-interval schedules of immediate reinforcement in three components of a multiple schedule. Additional response-independent reinforcers were available in one component and additional delayed reinforcers were available in another component. The results showed that resistance to disruption was greater in the components with added reinforcers than without them (i.e., better stimulus-reinforcer relations), but did not differ for the components with added response-independent and delayed reinforcement. In Experiment 2, a component presenting immediate reinforcement alternated with either a component that arranged equal rates of reinforcement with a proportion of those reinforcers being response independent or a component with a proportion of the reinforcers being delayed. Results showed that resistance to disruption tended to be either similar across components or slightly lower when response-reinforcer relations were degraded with either response-independent or delayed reinforcers. These findings suggest that degrading response-reinforcer relations can impact resistance to change, but that impact does not depend on the specific method and is small relative to the effects of the stimulus-reinforcer relation.     

Dishabituation with component transitions may contribute to the interactions observed during multiple schedules

Rates of responding by rats were usually higher during the variable interval (VI) 30-s component of a multiple VI 30-s fixed interval (FI) 30-s schedule than during the same component of a multiple VI 30-s VI 30-s schedule (Experiment 1). Response rates were also usually higher during the FI 30-s component of a multiple VI 30-s FI 30-s schedule than during the same component of a multiple FI 30-s FI 30-s schedule (Experiment 2). The differences in response rates were not observed when the components provided VI or FI 120-s schedules. These results were predicted by the idea that differences in habituation to the reinforcer between multiple schedules contribute to behavioral interactions, such as behavioral contrast. However, differences in habituation were not apparent in the within-session patterns of responding. Finding differences in response rates in both experiments violates widely-held assumptions about behavioral interactions, including that behavioral contrast does not occur for rats and that improving the conditions of reinforcement decreases, rather than increases, response rate in the alternative component.     

Effect of contingent auditory stimuli on concurrent schedule performance: an alternative punisher to electric shock

This study explored whether load auditory stimuli could be used as functional punishing stimuli in place of electric shock. Three experiments examined the effect of a loud auditory stimulus on rats’ responding maintained by a concurrent reinforcement schedule. In Experiment 1, overall response rate decreased when a concurrent 1.5 s tone presentation schedule was superimposed on the concurrent variable interval (VI) 180-s, VI 180-s reinforcement schedule. On the contrary, response rate increased when a click presentation schedule was added. In Experiment 2, the extent of the response suppression with a 1.5 s tone presentation varied as a function of the frequency of the reinforcement schedule maintaining responses; the leaner the schedule employed, the greater the response suppression. In Experiment 3, response suppression was observed to be inversely related to the duration of the tone; response facilitation was observed when a 3.0-s tone was used. In Experiments 1 and 2, a preference shift towards the alternative with richer reinforcement was observed when the tone schedule was added. In contrast, the preference shifted towards the leaner alternative when the click or longer duration stimulus was used. These results imply that both the type and duration of a loud auditory stimulus, as well as the reinforcement schedule maintaining responses, have a critical role in determining the effect of the stimuli on responding. They also suggest that a loud auditory stimulus can be used as a positive punisher in a choice situation for rats, when the duration of the tone is brief, and the reinforcement schedule maintaining responses is lean.     

Effect of signaling reinforcement on resistance to change in a multiple schedule

This study evaluated the effect of a signal on resistance to change using a multiple schedule of reinforcement. Experiment 1 presented pigeons with three schedules: a signaled delay to reinforcement schedule (a two-link chain schedule with a variable-interval 120-s initial link followed by a 5-s fixed-time schedule), an unsignaled delay schedule (a comparable two-link tandem schedule), and an immediate, zero-delay variable-interval 125-s schedule. Two separate disruption procedures assessed resistance to change: extinction and adding a variable-time 20-s schedule of reinforcement to the inter-component interval. Resistance to change tests were conducted twice, once with the signal stimulus (the terminal link of the chain schedule) present and once with it absent. Results from both disruption procedures showed that signal absence reduced resistance to change for the pre-signal stimulus. In probe choice tests subjects strongly preferred the signal stimulus over the unsignaled stimulus and exhibited no reliable preference when given a choice between the signal stimulus and immediate stimulus. Experiment 2 presented two equal signaled schedules where, during resistance to change tests, the signal remained for one schedule and was removed for the second. Resistance to change was consistently lower when the signal was absent.     

Locomotor activity in a novel environment predicts both responding for a visual stimulus and self-administration of a low dose of methamphetamine in rats

There is evidence that visual stimuli used to signal drug delivery in self-administration procedures have primary reinforcing properties, and that drugs of abuse enhance the reinforcing properties of such stimuli. Here, we explored the relationships between locomotor activity, responding for a visual stimulus, and self-administration of methamphetamine (METH). Rats were classified as high or low responders based on activity levels in a novel locomotor chamber and were subsequently tested for responding to produce a visual stimulus followed by self-administration of a low dose of METH (0.025 mg/kg/infusion) paired with the visual stimulus. High responder rats responded more for the visual stimulus than low responder rats indicating that the visual stimulus was reinforcing and that operant responding for a visual stimulus has commonalities with locomotor activity in a novel environment. Similarly, high responder rats responded more for METH paired with a visual stimulus than low responder rats. Because of the reinforcing properties of the visual stimulus, it was not possible to determine if the rats were responding to produce the visual stimulus, METH or the combination. We speculate that responding to produce sensory reinforcers may be a measure of sensation seeking. These results indicate that visual stimuli have unconditioned reinforcing effects which may have a significant role in acquisition of drug self-administration, a role that is not yet well understood.     

Stimulus change dis-habituates operant responding supported by water reinforcers

The present experiment examined whether habituation contributes to within-session decreases in operant responding for water reinforcers. The experiment asked if this responding can be dishabituated, a fundamental property of habituated behavior. During baseline, rats’ lever pressing was reinforced by water on a variable interval 15-s schedule. During experimental conditions, rats responded on the same schedule and a new stimulus was introduced for 5 min at 15, 30 or 45 min into the 60-min session. The new stimulus was extinction, continuous reinforcement or flashing lights in different conditions. Rate of responding primarily decreased within the session during baseline. Introducing a new stimulus sometimes suppressed (extinction, continuous reinforcement) and sometimes increased (flashing lights) responding while it was in effect. The new stimulus increased responding after it ended and before it was presented in the session. The results are incompatible with the idea that non-habituation satiety factors (e.g., cellular hydration and blood volume) contributed to within-session changes in responding. These satiety factors should increase with increases in consumption, decrease with decreases in consumption and remain constant with constant consumption of water. Nevertheless, all stimulus changes increased operant responding for water. These results support the idea that habituation contributes to within-session decreases in responding for water reinforcers.     

Superstitious responding and reinforcement rate under concurrent variable-interval extinction schedules

To examine superstitious responding, four pigeons key pecked under multiple concurrent variable-interval 45 s variable-interval 90 s concurrent variable-interval 90 s variable-interval 180 s schedules in the absence of a changeover delay. The two variable-interval 90 s schedules then were replaced by extinction, and key-peck responding during extinction was examined as a function of the prevailing reinforcement rate. During the first several sessions, extinction-key responding was maintained closer to baseline levels in the presence of the higher reinforcement rate, and this effect dissipated or even reversed with continued exposure to extinction. Although extinction-key responding generally decreased to near-zero levels after several sessions, in a few instances, it continued for 30 and 45 sessions. These results demonstrate how concurrent variable-interval extinction schedules can be used to investigate what often has been labeled superstitious responding.     

Psychological distance to reward: equating the number of stimulus and response segments

Psychological distance to reward, or the segmentation effect, refers to the preference for a terminal link of a concurrent-chains schedule consisting of a simple reinforcement schedule (e.g. fixed interval [FI] 30s) relative to its chained-schedule counterpart (e.g. chained FI 15s FI 15s). This experiment was conducted to examine whether the segmentation effect is due to the number of terminal-link stimulus and response segments per se. Three pigeons pecked under a concurrent-chains schedule in which identical variable-interval (VI) schedules operated in the initial links. In each session, half the terminal-link entries followed one initial-link key and the other half followed the other initial-link key. The initial-link keys correlated with the different terminal links were manipulated across conditions. In the first three conditions, each terminal link contained a chained fixed-time (FT) FT schedule, and in the final three conditions, each terminal link contained a chained FI FI schedule. In each condition, in one terminal link (alternating), the order of two key colors correlated with the different schedule segments alternated across terminal-link entries, whereas in the other terminal link (constant), the order of two other key colors was identical for each entry. With the chained FT FT schedule terminal links, there was indifference between the alternating and constant terminal links within and across pigeons, as indexed by initial-link choice proportions. In addition, terminal-link response rates were relatively low. With the chained FI FI schedule terminal links, for each pigeon, there was relatively more preference for the alternating terminal link and terminal-link response rates increased relative to conditions with the chained FT FT schedule terminal links. These data suggest that the segmentation effect is not due simply to the number of terminal-link stimulus or response segments per se, but rather to a required period of responding during a stimulus segment that never is paired with reinforcement.     

Immediate post-exposure effects of high-peak-power microwave pulses on operant behavior of Wistar rats.

Behavioral effects of high-peak-power microwave pulses on Wistar rats were studied by operant schedules. Each of twelve rats that had been trained to press a lever to receive food pellets was assigned randomly in groups of four to three different schedules of reinforcement: fixed-ratio (FR), variable-interval (VI), and differential-reinforcement-of-low-rates (DRL). After achieving a steady baseline performance, each animal was exposed for 10 min to 1.25-GHz microwave radiation at 1-MW peak-power (10-microseconds pulse width). Each pulse produced a peak whole-body SA and SAR of 2.1 J/kg and 0.21 MW/kg. Total doses (SAs) were set to 0.50, 1.5, 4.5, and 14 kJ/kg by adjusting the pulse-repetition rate. The corresponding time-averaged whole-body SARs were 0.84, 2.5, 7.6, and 23 W/kg. A microwave-transparent animal holder was used to keep the animal's body axis parallel to the E-field. Exposures at the highest dose caused an average colonic temperature rise of 2.5 degrees C and these animals failed to respond at all for about 13 minutes after the exposure. Their colonic temperatures had decreased to 1.1 degrees C, or less, above their pre-exposure (normal) temperature level when they began to respond. The FR and VI animals failed to reach their baseline levels of performance thereafter, while those on the DRL schedule displayed variable effects. No behavioral effects were found at the lower dose levels. It is concluded that the behavioral perturbations produced by pulsed microwave irradiation were thermal in nature.     

Resistance to change varies inversely with reinforcement context

We report two experiments which test whether resistance to prefeeding and satiation for a variable-interval (VI) schedule that delivers a constant rate of reinforcement varies inversely with the reinforcement rate for an alternative schedule. In Experiment 1, eight pigeons responded in a multiple schedule in which the red key was always associated with a VI 90-s schedule and the green key with either a richer (VI 18s) or leaner (VI 540s) schedule in different conditions. After baseline training in each condition, prefeeding test sessions were conducted in which 10g, 20g, 30g, 40g, and 50g food were provided one-hour prior to test. Additional baseline training was given between each test session. In Experiment 2, two groups of pigeons responded in a multiple schedule similar to Experiment 1. After baseline training, pigeons were exposed to a 5-h satiation test session in which the VI 90-s schedule was available continuously. Test sessions were conducted when pigeons were maintained at 85%, 95%, and 85% of their body weights in an ABA design. Results of both experiments showed that responding in the VI 90-s schedule that alternated with a leaner schedule during baseline was more resistant to prefeeding and satiation. These data rule out alternative explanations for results of previous studies, and confirm that resistance to change varies inversely with reinforcement context.     

Rapid acquisition of preference in concurrent schedules: effects of reinforcement amount

Four pigeons were trained on concurrent variable-interval 30-s schedules. Relative reinforcer amounts arranged across the two alternatives were varied across sessions according to a pseudorandom binary sequence [cf., Hunter, I., Davison, M., 1985. Determination of a behavioral transfer function: white-noise analysis of session-to-session response-ratio dynamics on concurrent VI schedules. J. Exp. Anal. Behav. 43, 43-59]; the ratios (left/right) were either 1/7 or 7/1. Reinforcer amount was manipulated by varying the number of 1.2s hopper presentations. Sessions ended after 30 reinforcers (15 for each alternative). After approximately 30 sessions, response ratios for all pigeons began to track the changes in amount ratio (i.e., subjects' responding showed a moderate increase in sensitivity of responding to reinforcer amount). Characteristics of responding were similar to procedures in which reinforcer rate and immediacy have been manipulated, although sensitivity estimates for amount were lower than those previously obtained with rate and immediacy. This procedure may serve as a useful method for studying the effects of certain environmental manipulations (e.g., drug administration) on sensitivity to reinforcer amount.