The sensory basis of avoidance responding in the rat : Relative dominance of auditory or visual warning signals and safety signals

Three experiments determined the relative degree of stimulus control exerted by the elements of a compound auditory-visual stimulus when this stimulus was presented in various temporal relationships to a barpress avoidance response. When the compound discriminative stimulus for barpress avoidance responding consisted of onset of white noise and onset of either light or darkness, the white noise always exerted dominant control. When the compound discriminative stimulus consisted of offset of white noise and onset of light, neither element controlled avoidance responding. On the other hand, when a barpress avoidance response produced a compound feedback stimulus consisting of onset of white noise and onset of either light or darkness, the visual element always exerted dominant control. When the compound feedback stimulus consisted of offset of white noise and onset of light, both elements exerted stimulus control.     

Reversibility of single-incentive selective associations.

Rats were trained to press a lever in the presence of a tone-light compound stimulus and not to press in its absence. In each of two experiments, schedules were designed to make the compound a conditioned punisher for one group and a conditioned reinforcer for the other. In Experiment 1, one group's responding produced food in the presence of the compound but not in its absence. The other group's responding terminated the compound stimulus, and food was presented only in its absence. When tone and light were later presented separately, light controlled more responding than did tone in the former group, but tone gained substantial control in the latter. The same effects were also observed within subjects when the training schedules were switched over groups. In Experiment 2, two groups avoided shock in the presence of the compound stimulus. In the absence of the compound, one group was not shocked, and the other received both response-independent and response-produced shock. When tone and light were presented separately, the former group's responding was mainly controlled by tone, but the latter group's responding was almost exclusively controlled by light. These effects were also observed within subjects when the training schedules were switched over groups. Thus, these single-incentive selective association effects (appetitive in Experiment 1 and aversive in Experiment 2) were completely reversible. The schedules in which the compound should have been a conditioned reinforcer consistently produced visual control, and auditory control increased when the compound should have become a conditioned punisher. Currently accepted accounts of selective associations based on affinities between shock and auditory stimuli and between food and visual stimuli (i.e., stimulus-reinforcer interactions) do not adequately address these results. The contingencies of reinforcement most recently associated with the compound and with its absence, rather than the nature of the reinforcer, determined whether auditory or visual stimulus control developed.     

Visual dominance in the pigeon

In Experiment 1, three pigeons were trained to obtain grain by depressing one foot treadle in the presence of a 746-Hertz tone stimulus and by depressing a second foot treadle in the presence of a red light stimulus. Intertrial stimuli included white light and the absence of tone. The latencies to respond on auditory element trials were as fast, or faster, than on visual element trials, but pigeons always responded on the visual treadle when presented with a compound stimulus composed of the auditory and visual elements. In Experiment 2, pigeons were trained on the auditory-visual discrimination task using as trial stimuli increases in the intensity of auditory or visual intertrial stimuli. Again, pigeons showed visual dominance on subsequent compound stimulus test trials. In Experiment 3, on compound test trials, the onset of the visual stimulus was delayed relative to the onset of the auditory stimulus. Visual treadle responses generally occurred with delay intervals of less than 500 milliseconds, and auditory treadle responses generally occurred with delay intervals of greater than 500 milliseconds. The results are discussed in terms of Posner, Nissen, and Klein's (1976) theory of visual dominance in humans.     

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.     

Control of responding by location of auditory stimuli: rapid acquisition in monkey and rat

Monkeys require a considerably larger number of trials to bring responding under the control of the location of an auditory stimulus than cats, rats, and bats with the same experimental procedures. The present experiment sought to determine the conditions necessary for rapid acquisition of control of responding by location of noise and tone bursts in the monkey. Monkeys were run in an enclosure that contained four loudspeakers and four manipulanda. Two conditions were used in training. In the adjacent condition, a stimulus (noise or tone burst) was presented through one or other of two speakers and a response on the manipulandum adjacent to the speaker was reinforced with food. In the nonadjacent condition, a stimulus was presented through one of two speakers and a response on a manipulandum remote from the speaker was reinforced with food. Acquisition of control was measured by change in the percentage of reinforced responses during training. In the adjacent condition, responding came under control of location within zero to three sessions. In nonadjacent conditions, the animals required 14 to 20 sessions to come under control of location. These latter numbers are comparable to those reported in the literature for localization discrimination in monkeys.     

Overshadowing in operant conditioning as a function of discriminability

Pigeons were trained in an operant, go/no-go, discrimination by successive presentations of discrete, positive and negative trials. In separate groups, the rate of discriminaion learning based on an auditory cue, on visual cues of different discriminability, and on combined auditory and visual cues was determined. The auditory cue was tone/no-tone, the visual cue was a difference in brightness level on the key. Following discrimination training, stimulus control was tested in extinction by presenting tone or notone at each of four key-brightness levels.Learning was more rapid with two cues than with either cue singly, demonstrating summation. The contribution made by the tone cue to the learning of the discrimination decreased with increasing discriminability of the light cue. The control exerted by the tone cue also decreased (i.e., the tone cue was overshadowed) with increasing discriminability of the light cue. In certain cases, the tone cue decreased control by brightness, showing that in two-cue discriminations each cue may reduce the control exerted by the other.     

Associative explanations of discriminative inhibition effects

In two experiments rats were trained to discriminate between trials signalled by a tone during which responding was reinforced, and trials signalled by the tone in compound with a light. During tone-light trials reinforcers were still delivered, but there was no contingency between responding and reinforcement. Subsequent transfer tests showed that the light had acquired the ability to suppress operant responding. Various explanations of this phenomenon are discussed. It is concluded that the data are inconsistent with traditional accounts of discriminative control.     

Nicotine trained as a negative feature passes the retardation-of-acquisition and summation tests of a conditioned inhibitor

Nicotine functions as a negative feature in a Pavlovian discriminated goal-tracking task. Whether withholding of responding to the conditional stimulus (CS) reflects nicotine functioning as a conditioned inhibitor is unknown. Accordingly, the present research sought to determine whether nicotine trained as a negative feature passed the retardation-of-acquisition and summation tests, thus characterizing it as a pharmacological (interoceptive) conditioned inhibitor. In the retardation test, rats received either nicotine (0.4 mg/kg) or chlordiazepoxide (5 mg/kg) negative feature training in which the drug state signaled when a 15-sec light CS would not be paired with sucrose; light was paired with sucrose on intermixed saline sessions. Following acquisition of the discrimination, both groups received nicotine CS training in which sucrose was intermittently available on nicotine but not intermixed saline sessions. Acquisition of conditioned responding to the nicotine CS was slower in the nicotine negative feature group than in the chlordiazepoxide negative feature group. In the summation test, rats were assigned to either the nicotine negative feature group or a pseudoconditioning control. In this control, the light CS was paired with sucrose on half the nicotine and half the saline sessions. Both groups also received excitatory training in which a white noise CS was paired with sucrose. The summation test consisted of presenting the white noise in conjunction with nicotine. Conditioned responding evoked by the white noise was decreased in the negative feature but not the pseudoconditioning group. Combined, the results provide the first evidence that an interoceptive stimulus (nicotine) can become a conditioned inhibitor.     

Comparison of Single Unit Responses to Tone, Light, and Compound Conditioned Stimuli during Rabbit Classical Eyeblink Conditioning

Unit recordings and lesion studies have implicated the cerebellum as an essential site for the acquisition and maintenance of the conditioned eyeblink response. The current study looked at the neural characteristics of conditioned stimulus (CS) processing in the interpositus nucleus of the cerebellum after training New Zealand white rabbits (Oryctolagus cuniculus) in one of two conditioning paradigms: (a) compound conditioning (CMP), a compound CS consisting of light and tone paired with an air puff unconditioned stimulus (US); or (b) stimulus compounding (ALT), alternating blocks of tone CS and light CS trials paired with the air puff US. Single unit responses were recorded during five sessions after the animals had reached an asymptotic level of responding. Animals were tested for behavioral and neural responses to CS alone trials that included tone alone, light alone, and compound tone-light trials. For the CMP group, the compound CS elicited 80 to 90% conditioned eyeblink responses (CRs), whereas the individual tone and light CSs elicited only 40 to 50% CRs. For the ALT group, all three CSs (tone, light, and compound) elicited very high levels of responding of at least 80% CRs. For the CMP group, there were roughly equal numbers of cells responding to all of the CSs. This includes cells that responded exclusively to one, and only one, of the three stimuli and also those cells that responded to combinations of two or more. Cells from the ALT group were far more likely to respond exclusively to only one of the CSs. Both the behavioral and physiological results suggest that the compound tone-light stimulus was processed as a distinct stimulus, separate from the component tone and light. These results are discussed in the context of multisensory processing.     

Summation: Further assessment of a configural theory

Rats received Pavlovian conditioning in which food was signalled by a visual stimulus, A+, an auditory stimulus, B+, and a compound composed of different visual and auditory stimuli, CD+. Test trials were then given with the compound AB. Experiments 1 and 2A revealed stronger responding during AB than during CD. In Experiment 2B, there was no evidence of a summation of responding during AB when A+ B+ training was conducted in the absence of CD+ trials. A further failure to observe abnormally strong responding during ABwas found in Experiment 3 for which the training trials with A+ B+ CD+ were accompanied by trials in which C and D were separately paired with food. The results are explained in terms of a configural theory of conditioning, which assumes that responding during a compound is determined by generalization from its components, as well as from other compounds to which it is similar.     

Reevaluation of things past

The ability of rats to reevaluate previously presented information in light of subsequently provided information was evaluated using a CER (conditioned emotional response) procedure. In Experiment 1, rats suppressed responding to a compound light + tone stimulus that was repeatedly paired with shock. Groups of rats were then presented with only one element of the compound (the tone), either presented alone or paired with shock, for 15 days. During this 15-day period, two control groups received trials with the shock alone or neither stimulus nor shock. All of the rats were then tested with the other element of the compound (the light). Rats that had received the tone paired with shock during the intervening training continued to suppress to the light, whereas rats that had received the tone alone showed rapid extinction of the CER to light. The control groups showed that this effect was not due to the number of shock presentations received. A subsequent experiment also demonstrated that these results were not due to nonspecific stimulus effects. Apparently, a subsequent change in the associative strength of one element results in a similar change to the other element of a previously established compound stimulus.     

Stimulus control and compounding with ambient odor as a discriminative stimulus on a free-operant baseline

Previous experiments have demonstrated that the simultaneous presentation of independently established discriminative stimuli can control rates of operant responding substantially higher than the rates occasioned by the individual stimuli. This "additive summation" phenomenon has been shown with a variety of different reinforcers (e.g., food, water, shock avoidance, cocaine, and heroin). Discriminative stimuli previously used in such studies have been limited to the visual and auditory sensory modalities. The present experiment sought to (1) establish stimulus control on a free-operant baseline with an ambient olfactory discriminative stimulus, (2) compare olfactory control to that produced with an auditory discriminative stimulus, and (3) determine whether compounding independently established olfactory and auditory discriminative stimuli produces additive summation. Rats lever pressed for food on a variable-interval schedule in the presence of either a tone or an odor, with comparable control developed to each stimulus. In the absence of these stimuli responding was not reinforced. During stimulus compounding tests, the tone-plus-odor compound occasioned more than double the responses occasioned by either the tone or odor presented individually. Thus, the current study (1) established stimulus control with an ambient olfactory discriminative stimulus in a traditional free-operant setting and (2) extended the generality of stimulus-compounding effects by demonstrating additive summation when olfactory and auditory discriminative stimuli were presented simultaneously.     

Selective attention and stimulus-reinforcer interactions in the pigeon

The study supplies further evidence that non-associative effects and temporal-spatial similarities between certain combinations of cue and consequence cannot explain all instances of stimulus-reinforcer interactions. Pigeons were trained to press a treadle in the presence of a discriminative compound stimulus either to avoid shock or to obtain a food reinforcer. The compound stimulus was composed of diffuse tone and light cues which had identical temporal patterns of onset, duration and offset. With the avoidance schedule the auditory cue acquired more control than the visual cue; however, when food was the reinforcer, the visual cue exerted more control. This pattern of stimulus control on the appetitive schedule did not change if random shocks were also added, even though these shocks were equal in density to the food presentations and equal in magnitude to those used for the avoidance schedule. Other changes in the appetitive procedure, such as making the tone spatially contiguous with food and removing the light in the food hopper, also failed to alter the relative control by the different cues. Prior training with a food reinforcer did not produce any change in the relative control by the two cues when the birds were retrained on the shock-avoidance schedule. The results suggest that some frequently stated alternatives to selective associability are not adequate explanations of this instance of a stimulus-reinforcer interaction.     

SELECTIVE ASSOCIATIONS PRODUCED SOLELY WITH APPETITIVE CONTINGENCIES: THE STIMULUS-REINFORCER INTERACTION REVISITED

In studies reporting stimulus-reinforcer interactions in traditional conditioning paradigms, when a tone-light compound was associated with food the light gained stimulus control, but when the compound was paired with shock avoidance the tone gained control. However, the physical nature of the reinforcerrelated events (food vs. shock) presented in the presence of the tone-light compound was always confounded with the conditioned hedonic value of the compound's presence relative to its absence. When the compound was paired with shock, its presence was negative relative to its absence (which was shock-free). In contrast, when the compound was paired with food, its presence was positive relative to its absence (which was food-free). The present experiment dealt with this confounding effect by conditioning a tone-light compound to be positive or negative, relative to its absence, solely with food reinforcement. One group of rats received food for responding in the presence of the tone-light compound and no food in its absence. The other group also responded in the presence of the compound, but received food only in its absence. These rats were trained on a chained schedule in which responding in the presence of the tone-light compound produced a terminal link signaled by the absence of the compound; responding ceased in the terminal link because it delayed food delivery. In a test session to assess stimulus control by the elements of the compound, tone and light were presented separately under extinction conditions. Rats that had been exposed to a positive correlation between food and the compound emitted almost double the responses in the presence of the light as in the presence of the tone. In comparison, rats that had been exposed to a negative correlation emitted only two thirds as many responses in the presence of the light as in the presence of the tone. Because this selective association was produced using only food, it appears that the contingencies under which a reinforcer is presented, rather than (or as well as) its physical properties, can generate the selective associations previously attributed to “stimulus-reinforcer interactions.” This could mean that regardless of the class of reinforcer that ultimately maintains responding (appetitive or aversive), the contingency-generated hedonic value of the compound stimulus may influence the dominant modality of stimulus control.     

Spontaneous configuring to a tone-light compound using appetitive training

Rats were trained to bar-press for water in the presence of a simultaneous compound tone-light stimulus using a discrete-trial procedure. Different groups were given between 300 and 3900 trials. When subsequently tested in extinction, all rats showed consistent levels of responding to the compound stimulus but responding to the tone and light components declined as a function of number of training trials. This is a demonstration of spontaneous configuring. Two other experiments showed that with similar procedures generalization from a component of the training stimulus to other values on the same continuum decreases with increased training with the compound stimulus. It was concluded that configuring and generalization from a single stimulus involve the same processes.     

Compounding discriminative stimuli controlling free-operant avoidance

The performances of three rats were stabilized on a multiple schedule that maintained responding by a free-operant avoidance schedule during independent presentations of tone and light. The simultaneous absence of these stimuli signalled shock-free periods and controlled response cessation. Subsequently, test sessions were administered consisting of independent presentations of each stimulus and these stimuli compounded (tone-plus-light). During an extinction test, additive summation was observed to the compounded stimuli, i.e., more responses were emitted to the compound than to either tone or light. During a series of 28 maintenance-test sessions in which the shock schedule remained operative, the compounded stimuli produced a generally enhanced response rate and fewer pauses terminating with shock than either single stimulus condition. These results extend the generality of free-operant additive summation to responding maintained by aversive control. In addition, a comparison of the present study with previous experiments reporting additive summation of positively reinforced responding indicates that similar variables—rate and aversive differences between training stimulus conditions—should be considered in accounting for response distributions during stimulus compounding when responding is controlled by either positive or negative contingencies.     

Differential autoshaping to common and distinctive elements of positive and negative discriminative stimuli

The learning by hungry pigeons of a discrimination between two successively presented compound visual stimuli was investigated using a two-key autoshaping procedure. Common and distinctive stimulus elements were simultaneously presented on separate keys and either followed by food delivery, S+, or not, S−. The subjects acquired both between-trial and within-trial discriminations. On S+ trials, pigeons pecked the distinctive stimulus more than the common stimulus; before responding ceased on S− trials, they pecked the common stimulus more than the distinctive one. Mastery of the within-display discrimination during S+ trials preceded mastery of the between-trials discrimination. These findings extend the Jenkins-Sainsbury analysis of discriminations based upon a single distinguishing feature to discriminations in which common and distinctive elements are associated with both the positive and negative discriminative stimuli. The similarity of these findings to other effects found in autoshaping—approach to signals that forecast reinforcement and withdrawal from signals that forecast nonreinforcement—is also discussed.     

The effect of a flashing visual stimulus on the auditory continuity illusion

The effect of a visual stimulus on the auditory continuity illusion was examined. Observers judged whether a tone that was repeatedly alternated with a band-pass noise was continuous or discontinuous. In most observers, a transient visual stimulus that was synchronized with the onset of the noise increased the limit of illusory continuity in terms of maximum noise duration and maximum tone level. The smaller the asynchrony between the noise onset and the visual stimulus onset, the larger the visual effect on this illusion. On the other hand, detection of a tone added to the noise was not enhanced by the visual stimulus. These results cannot be fully explained by the conventional theory that illusory continuity is created by the decomposition of peripheral excitation produced by the occluding sound.     

Human sensory dominance

Human Ss matched an auditory and a visual stimulus for subjective magnitude. Then each stimulus was used as a cue in a reaction time task. On occasions when both stimuli were presented simultaneously, Ss’ responding was seen to be dominated by the visual stimulus. Of further interest was the finding that on some occasions of simultaneous light-tone presentation Ss were unaware that the tone had been presented. This apparent prepotency of the visual over the auditory stimulus was seen to persist across a variety of experimental conditions, which included giving Ss verbal instructions to respond to the tone when both stimuli were presented simultaneously.     

Conditioned inhibition from incomplete reductions in the probability of reinforcement

Five experiments, all using appetitive, discrete trial operant conditioning, studied the properties of a stimulus that signalled a reduction in the probability of reward. Discriminations were trained in which reinforcement was available on 100% of trials signalled by a tone, but only on some occasions when the tone was presented simultaneously with a light. The properties of the light were assessed in summation tests with a clicker. The first two experiments established that if the tone-light compound signalled reinforcement on only 25%, 33% or 50% of trials the light acted as a discriminative inhibitor, suppressing responding maintained by the clicker. In these experiments reinforcement had been available on 86.7% of clicker trials during initial training. Experiments 3, 4 and 5 examined further the properties of the light after animals had been trained on a discrimination where reinforcement was available on 50% of trials signalled by the tone-light compound. The light was evaluated in a summation test with a clicker, which had signalled only a 25% or 15% probability of a reinforcement. In this instance, the light did not suppress the responding maintained by the clicker. Instead, the light acted as an excitatory discriminative stimulus to enhance that responding. This dependence of the properties of the light on the reinforcement history of the clicker poses problems for most contemporary models of inhibitory performance.