Cocaine and food deprivation: effects on food-reinforced fixed-ratio performance in pigeons.

Key pecking by 6 pigeons was maintained by a fixed-ratio 30 schedule of food presentation while body weights were 80% of free-feeding weights. Acute administration of cocaine (0.3 to 13.0 mg/kg, i.m.) dose-dependently decreased response rates. Dose-effect curves were shifted to the right when 3 of the 6 pigeons were maintained at 70% of free-feeding weights and were shifted to the left when the other 3 pigeons were maintained at 90% of free-feeding weights. Then a dose of cocaine that initially decreased response rates by more than 95% of control rates was administered before each daily session. Comparable degrees of tolerance to these rate-decreasing effects developed in the two groups. The rate at which responding recovered was relatively rapid for pigeons in the 70% free-feeding-weight group and was slower for 2 of the 3 pigeons in the 90% free-feeding-weight group. When body weights were then increased from 70% to 80% or were decreased from 90% to 80% of free-feeding weight, performance was disrupted initially only for pigeons whose weight went from 70% to 80% of free feeding. In the present experiment the degree of deprivation may have indirectly influenced the degree of tolerance that developed to cocaine's response rate-decreasing effects because it directly influenced the dose chosen to be administered chronically. The degree of deprivation appeared to have a more direct influence on the rate at which tolerance developed.     

On the effects of food deprivation and component reinforcer rates on multiple-schedule performance

Six pigeons were used to investigate the effects of varying body weight and component reinforcer rates in two-component multiple variable-interval variable-interval schedules. In Parts 1 and 3 of the experiment, unequal component reinforcer rates were arranged, and body weights were respectively increased and decreased. At 80% ad lib weight, response-rate ratios were closer to unity than reinforcer-rate ratios, but at 100% or more of ad lib weight, response-rate ratios generally equaled reinforcer-rate ratios. In Part 2, component reinforcer-rate ratios were varied over five conditions with the subjects maintained at 100% or more of their ad lib weights, and response-rate ratios matched reinforcer-rate ratios. The data thus support the empirical finding that response allocation in multiple schedules is a function of deprivation. Although this qualitative result is predicted by three models of multiple-schedule performance, only a model that assumes no direct component interaction adequately describes the data.     

The effects of cocaine on behavior maintained by timeout from avoidance.

Rats were trained on concurrent schedules under which responses on one lever postponed shock (avoidance) and responses on the other lever produced brief (2-min) periods of signaled timeout from avoidance. For 6 rats, timeout from avoidance was programmed on a variable-interval 45-s schedule that generally resulted in rates that were lower than those on the avoidance lever. For another 6 rats, timeout was arranged on a variable-ratio 15 schedule that produced higher baseline rates. Cocaine (3 to 40 mg/kg) produced large, dose-dependent increases in behavior maintained by timeout in both groups of rats. Avoidance responding was also generally increased by cocaine, but the increases were of lesser magnitude. Increases in response rates were seen across a broad range of doses on behavior maintained by either interval or ratio schedules, an outcome that was unexpected on the basis of most studies of cocaine on food-maintained behavior. These results were similar to those of previous studies of the effects of amphetamine on behavior maintained by timeout from avoidance and suggest that stimulant drugs affect behavior maintained under a shock-postponement schedule differently than they affect behavior maintained by timeout from avoidance.     

Negative reinforcement as shock-frequency reduction

Is a conditioned aversive stimulus necessary in avoidance conditioning? Or is a reduction in the rate of aversive stimulation alone sufficient to generate and maintain an avoidance response? Rats were subjected to an avoidance procedure in which shocks occurred randomly in time, but a response could reduce the overall rate of shock. Fifteen acquisition curves, obtained from 16 animals, showed both immediate and delayed, rapid and gradual increases in response rate; there was no representative acquisition curve. Response rates were directly related to the amount by which the response reduced shock frequency. In extinction, when shock rates were not affected by responding, the response total was inversely related to the amount by which the response had reduced shock frequency during prior conditioning, with as many as 20,000 extinction responses when the shock frequency reduction had been relatively small. Responding on this procedure shows that avoidance conditioning can occur without benefit of either classical exteroceptive stimuli or covert stimuli inferred from the temporal constancies of a procedure. It also shows that reduction in shock rate is alone sufficient to maintain avoidance.     

Human, free-operant avoidance of “time out” from monetary reinforcement

To assess the aversive effects of withdrawing monetary reinforcement, human subjects were exposed to a free-operant avoidance procedure in which periods of no reinforcement occurred if the subject failed to respond, and each response postponed withdrawal of reinforcement. Avoidance behavior was developed either through specific instructions about the consequence of responding or through preliminary escape-avoidance training. In all cases, rates of response were found to be a positively accelerated function of decreases in the duration by which responding postponed reinforcement withdrawal. The findings with respect to the function relating avoidance behavior to the interval of postponement were viewed as similar to those obtained when shock is used as the aversive event in free-operant avoidance conditioning.     

The rabbit as a subject in behavioral research

Domestic rabbits were exposed to a wide variety of variables employed in behavioral research. It was found that: (1) Although food could be used as a reinforcer, the long periods of severe deprivation required to reduce body weight made its use impractical. (2) Water was an efficient reinforcer in that it maintained high rates of behavior after 22 hr of deprivation. (3) Except that rates of responding were higher, fixed-ratio and variable-interval schedules of reinforcement produced patterns of behavior similar to those demonstrated by rats and pigeons. (4) Although the duration of the post-reinforcement pause was a function of the duration of the interval under fixed-interval schedules, scalloping, as defined as a gradually increasing rate of responding between reinforcement, was not evident. (5) When provided with the means to both turn on and turn off intracranial stimulation, the duration of the stimulation and the frequency with which it was turned on and off was a function of the intensity of the stimulation. (6) Electric shock could suppress behavior and maintain escape responding, but would maintain avoidance responding only in a few subjects.     

Effects of signaled and unsignaled shock on schedule-controlled lever pressing and schedule-induced licking: Shock intensity and body weight

Schedule-controlled lever pressing and schedule-induced licking were studied in rats under a multiple fixed-interval fixed-interval schedule of food reinforcement. Following acquisition of stable rates of pressing and licking, a multiple variable-time variable-time schedule of electric-shock delivery was superimposed upon the baseline schedule. In only one component of the multiple schedule, a 5-sec stimulus preceded each shock (signaled shock). In the other component shock was unsignaled. Several shock intensities (Experiment 1) and body weights (Experiment 2) were studied. Lever pressing and licking were affected similarly by experimental manipulations, although with parametric differences. Depending upon shock intensity and body weight, rates of lever pressing and licking were hardly suppressed, suppressed primarily in the unsignaled shock component (differential suppression), or markedly suppressed in both components. Differential suppression during components with signaled and unsignaled shock and conditioned suppression of responding during the preshock stimulus appeared not to be functionally related. Differential suppression depended more on the discriminability of shock-free time, and on shock intensity, body weight, and the type of response than on the “preparatory” behavior preceding shock.     

Variable-ratio schedules of timeout from avoidance: effects of d-amphetamine and morphine.

Rats were trained on concurrent schedules in which pressing one lever postponed shock and pressing the other lever produced periods of signaled timeout from avoidance on variable-ratio schedules. These procedures generated high rates of timeout-reinforced responding and provided a baseline for studying the effects of drugs on behavior maintained by different types of negative reinforcement (shock postponement vs. timeout). Morphine (2.5 to 10.0 mg/kg) reduced behavior maintained by timeout at doses that increased or had no effect on avoidance responding. In contrast, d-amphetamine (0.125 to 2.0 mg/kg) produced large increases in timeout responding at doses that had minimal effect on avoidance in rats trained on variable-interval and variable-ratio schedules. Thus, the event-dependent effects of morphine, observed in previous studies in which timeout responding was maintained at low rates by interval schedules, were replicated with high timeout rates maintained by variable-ratio schedules. The effects of d-amphetamine could also be described as "event dependent" because timeout responding was stimulated more than avoidance regardless of the maintenance schedule or baseline rate.     

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.     

A comparison of operant licking and lever pressing in the rat

Two groups of rats were trained on a signaled, free-operant, avoidance procedure to lick or to lever press in order to avoid shock while water-deprived or satiated and, in the case of licking, while ingesting deionized water, isotonic saline, or 10% sucrose. The most effective avoidance licking occurred while the rats were water-deprived and ingesting 10% sucrose. Water deprivation level had no effect on lever pressing for shock avoidance. Two other groups of rats were operantly conditioned to lick or to lever press for food pellets while waterdeprived or satiated and, in the case of licking, while ingesting deionized water or 10% sucrose. The most effective licking for food reinforcement occurred while the rats were water-deprived and ingesting 10% sucrose. Water deprivation level had no effect on lever pressing for food reinforcement. The data indicated that effective operant licking must be supported by factors related to water regulation and taste palatability.     

Effects of chlordiazepoxide and cocaine on concurrent food and avoidance-of-timeout schedules.

Five rats were trained on a concurrent schedule in which responses on one lever produced a food pellet on a random-interval 30-s schedule during 10 s of food availability associated with distinctive exteroceptive stimuli. Responses on another lever postponed for 20 s the presentation of a 50-s timeout, during which all stimuli were extinguished and the schedule contingencies on the food lever were suspended. The response rates maintained by the random-interval schedule exceeded those maintained by the avoidance contingency, but both provided a stable baseline to assess the behavioral effects of different drugs. Low doses of cocaine hydrochloride (1 and 3 mg/kg) did not affect food-reinforced responding or avoidance response rates. Intermediate doses (5.6, 10, and 13 mg/kg) produced a dose-dependent decrease in food-maintained and avoidance response rates, and both types of responding were virtually eliminated after administration of the highest doses (17 and 30 mg/kg) of cocaine. Low doses of chlordiazepoxide (1 and 3 mg/kg) increased food-maintained and avoidance response rates, and both rates decreased systematically after 10 and 30 mg/kg of this drug. The effects of cocaine and chlordiazepoxide on response rates maintained by avoidance of timeout from food presentation were unlike those reported when subjects responded to avoid shock presentation. The results of this experiment thus provide evidence to suggest that the effects of drug administration on avoidance behavior may be a function of the nature of the consequent event to be avoided.     

Conditioned suppression of an avoidance response by a stimulus paired with food

Three food-deprived Long-Evans rats were exposed to a non-discriminated shock avoidance procedure. Superimposed upon this operant avoidance baseline were periodic presentations of a conditioned stimulus that was paired with food, the unconditioned stimulus. These pairings resulted in increases in the rate of shock over that recorded when the conditioned stimulus was not present. A traditional suppression ratio failed to reveal any differential effect of the conditioned stimulus on the overall rate of avoidance responding, although all subjects showed a consistent pattern of pausing and postshock response bursts during presentations of the conditioned stimulus. When food was withheld during a final extinction phase, the conditioned stimulus ceased to occasion increases in shock rates and disruptive postshock response bursts were eliminated. An analysis of conditioned suppression procedures is proposed that stresses not only operant-Pavlovian or appetitive-aversive incompatibility, but also the manner in which the baseline schedule of reinforcement affects operant behavior changes that are elicited by the superimposed Pavlovian procedure.     

Effects of body weight on discriminative-stimulus control by phencyclidine in the pigeon.

Using a color-tracking procedure with responses reinforced under a second-order schedule, the discriminative-stimulus properties of phencyclidine were studied in pigeons maintained at 70%, 80%, or 90% of their free-feeding weights. The generalization curves for phencyclidine were similar at all three body weights. Generalization curves for pentobarbital, d-amphetamine, and saline were also unrelated to body weight. These data suggest that food deprivation may not influence the discriminative-stimulus properties of drugs in the way that it influences the reinforcing-stimulus properties of drugs. The reason may be that during discrimination training interoceptive stimuli resulting from food deprivation do not become conditioned to the stimulus properties of the drug, because the food-deprivation stimuli are paired equally often with the presence and absence of drug stimuli.     

Effects of naloxone on acquisition and extinction of jump-up avoidance behavior in rats

Two experiments were carried out to assess the effects of the opioid antagonist, naloxone, on the acquisition and extinction of shock avoidance by rats in the jump-up apparatus. In Experiment 1 naloxone pretreatment facilitated acquisition but had no effect on extinction of avoidance behavior. In the second experiment the effect of naloxone on acquisition was replicated and in addition, it was shown that naloxone enhanced freezing when a response prevention or flooding procedure was introduced. Again naloxone failed to alter the course of extinction, nor did it interact with the effects of flooding which, by itself, facilitated extinction. The results suggest that naloxone's effects are limited to increasing the functional intensity of the US, and provide further support for the dissociation between extinction of avoidance behavior and other indices of fear.     

Rats defend different body weights depending on palatability and accessibility of their food.

Normal adult rats lived on powdered diets adulterated to contain as much as 1.6% quinine sulfate, on a palatable high-fat diet, or in Skinner boxes with 45-mg Noyes pellets available on fixed-ratio (FR) schedules as high as FR 156. They maintained lower body weights over periods of months in proportion to the percentage of quinine adulteration or the fixed ratio. Rats exposed to the high-fat diet overate as much and gained weight as rapidly as rats recovering from food deprivation, and became moderately obese. Rats having become lean or obese contingent on the palatability or accessibility of their diet defended body weight by eating more in the cold, less when force-fed by gavage, and more to restore weight after food deprivation. Yet on chow they restored and defended body weights typical of rats whose diet had been confined to commercially prepared chow. These results are interpreted to be inconsistent with motivational models that rigidly distinguish drive from incentive, that treat body weight changes as evidence for failure to regulate energy balance or body weight, or that rely exclusively on deprivation of food or reduction of body weight for definitions of need for calories. Instead, caloric homeostasis in rats may incorporate ecological constraints.     

The effect of rate of delivery of response-independent shocks upon avoidance responding

Two hooded rats were trained to bar-press to avoid electric shock on a continuous avoidance schedule with response-shock and shock-shock intervals equal. The rate of delivery of response-independent shocks superimposed on this schedule was varied. The response-independent shocks led to generally higher response rates but, with responses during shock omitted, the rates decreased as the response-independent shock rate was increased. The actual shock rate received by the subjects was linearly related to the maximum potential shock rate. There was an increasing, negatively accelerated function between percentage avoidance and response rate, but there was no consistent relation between the number of shocks avoided and response rate. Response rate decreased as the potential shock rate increased, but responding was maintained even when as few as 15% of the shocks could be avoided.     

SUPPRESSIVE AND FACILITATIVE EFFECTS OF SHOCK INTENSITY AND INTERRESPONSE TIMES FOLLOWED BY SHOCK

Although response‐dependent shock often suppresses responding, response facilitation can occur. In two experiments, we examined the suppressive and facilitative effects of shock by manipulating shock intensity and the interresponse times that produced shock. Rats' lever presses were reinforced on a variable‐interval 40‐s schedule of food presentation. Shock followed either long or short interresponse times. Shock intensity was raised from 0.05 mA to 0.4 mA or 0.8 mA. Overall, shock contingent on long interresponse times punished long interresponse times and increased response rates. Shock contingent on short interresponse times punished short interresponse times and decreased response rates. In Experiment 1, raising the range of interresponse times that produced shock enhanced these effects. In Experiment 2, the effects of shock intensity depended on the interresponse times that produced shock. When long interresponse times produced shock, low intensities increased response rates. High intensities decreased response rates. When short interresponse times produced shock, high shock intensities punished short interresponse times and decreased response rates more than low intensities. The results may explain why punishment procedures occasionally facilitate responding and establish parameters for future studies of punishment.     

Some effects of methylphenidate on stimulus control of ratio avoidance behavior in the rat

After exposure to an avoidance schedule which included a warning signal, a rat was placed on a multiple schedule in which the first component was the same as before, i.e., a single response reset the response-shock interval, delaying shock, and the second component differed only in that four bar-presses were required to postpone shock. A fixed ratio requirement of four responses (FR 4) generated behavior resembling a fixed ratio requirement of one response (FR 1) since responding was controlled by the warning signal but more shocks were received. At a dosage of 2 mg/kg, methylphenidate given intraperitoneally decreased shock frequency during FR 4 periods while FR 1 behavior was not affected; at 4 mg/kg, stimulus control of avoidance responding was impaired during both components. Results at 4 mg/kg were partially confirmed by two animals exposed to an FR 4 avoidance schedule which included a warning signal but with different parameters. Response distributions showed that methylphenidate increased response rates in the absence of the warning signal, i.e., stimulus control of ratio-avoidance behavior was impaired although the increased response rates reduced shock frequency. One hour later responses again occurred more frequently during the signal than in its absence but shocks were less frequent than during control (non-drug) periods.     

Remote effects of aversive contingencies: Disruption of appetitive behavior by adjacent avoidance sessions

Disruption of ongoing appetitive behavior before and after daily avoidance sessions was examined. After baselines of appetitive responding were established under a fixed-interval 180-s schedule of food presentation, 4 rats were exposed to 40-min sessions of the appetitive schedule just prior to 100-min sessions of electric shock postponement, while another 4 rats received the 40-min appetitive sessions just following daily sessions of shock postponement. In all 8 subjects, fixed-interval response rates decreased relative to baseline levels, the effect being somewhat more pronounced when the avoidance sessions immediately followed. The disruption of fixed-interval responding was only partially reversed when avoidance sessions were discontinued. During the initial exposure to the avoidance sessions, patterns of responding under the fixed-interval schedule were differentially sensitive to disruption, with high baseline response rates generally more disturbed than low rates. These disruptions were not systematically related to changes in reinforcement frequency, which remained fairly high and invariant across all conditions of the experiment; they were also not systematically related to the response rates or to the shock rates of the adjacent avoidance sessions. The results, while qualitatively resembling patterns of conditioned suppression as typically studied, occurred on a greatly expanded time scale. As disruption of behavior extending over time, the present data suggest that some forms of conditioned suppression are perhaps best viewed within a larger temporal context.     

Free operant avoidance in rats under increased nitrogen pressures.

Rats performed on a free operant avoidance schedule with a response-shock interval of 20 sec. and a shock-shock interval of 2 sec. Avoidance response rates increased and shock frequency decreased when the rats were exposed to elevated pressures of both air and a nitrogen-oxygen mixture in a hyperbaric chamber. Increases in response rates were related to raised partial pressures of nitrogen at 89.0 psi and 111.3 psi. Conditional probabilities of interresponse times indicated that increases in response rates were not due to disruption of temporal discrimination. Increased avoidance rates under pressure suggested direct excitatory effects of high pressures of nitrogen.