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.     

Conditioning in human opiate addicts

Eight volunteers maintained on daily methadone participated in a classical conditioning procedure to determine which if any of the elements of narcotic withdrawal could be conditioned. The unconditioned stimulus was the injection of a small dose of naloxone. The unconditioned response was a brief precipitated withdrawal syndrome. The conditioning stimulus was a tone, odor, and injection of saline. Conditioning was successful in the pilot study in 5 of 8 subjects. The conditioned response consisted of tearing, yawning, lacrimation, systolic blood pressure increase, respiratory irregularities and subjective feelings of narcotic withdrawal sickness (nausea, muscle aches, chills). A second group of 8 subjects showed, in addition to the above, evidence of conditioning of heart rate, respiratory rate and skin temperature decrease. These laboratory findings support the clinical reports of a conditioned withdrawal syndrome and suggest ways to improve treatment results by detecting and extinguishing or modifying conditioned responses.     

Visual encoding of patterns is subject to dual-task interference

Two stimuli were presented at varying stimulus onset asynchronies (SOAs), with each stimulus associated with a distinct task. The first stimulus was a tone at one of either two or three frequencies. In two conditions, the task associated with a tone was either a speeded two-alternative discrimination (2AD), or a speeded three-alternative discrimination (3AD) based on the pitch of the tone. In a third condition, subjects were told to ignore the tone. The second stimulus was a briefly exposed study matrix of red and black squares followed by a mask. After a fixed delay, the mask was replaced by a test matrix that was either the same or different from the study matrix. The task associated with the matrices was to indicate, with no speed pressure, whether the study matrix and the test matrix were the same or different. Results from each speeded AD condition showed that subject's accuracy in the matrix task decreased as the SOA between the tone and the study matrix decreased. This effect was larger for the 3AD tone task than with a 2AD tone task. In addition, within each speeded AD condition, longer RTs in the tone task were associated with lower accuracy in the matrix task. None of these effects was evident when the subjects were told to ignore the tone. These results suggest that encoding visual information can be subject to significant capacity limitations imposed by cross-modal multitasking.     

Signal recognition as influenced by presentation schedules

Performance in a recognition task involving two amplitudes of the same tone was investigaled over a wide range of presentation schedules. The task was arranged so that there was no trial-to-trial feedback or other information regarding the relative frequencies of the two tones. The hit and false alarm rates (the proportion of “loud” responses to loud and soft stimuli, respectively) on any given trial were strongly influenced by the stimulus and response on the preceding trial. In general, Ss tended to repeat the last response and were more accurate after a stimulus alternation than after a stimulus repetition. In addition, hit and false alarm rates were inversely related to the presentation probability of the loud tone, in contrast to the direct relation typically found in signal detection experiments and in recognition experiments with trial-to-trial feedback. A mathematical model incorporating three processes (memory, comparison, and decision) was shown to give a good account of these data.     

Potentiation and overshadowing in odor-aversion learning: Role of method of odor presentation,the distal-proximal cue distinction,and the conditionability of odor

Four experiments with rat subjects examined interactions between odors and tastes in compound aversion conditioning. In Experiments 1 and 2, a saccharin taste potentiated the conditioning of an almond odorant presented on a cup near the location of the drinking spout, but overshadowed the same odorant when it was mixed in drinking water. This difference depended on the type of odor administered during conditioning rather than testing (Experiment 2). Experiments 3 and 4 examined implications of two other differences between the cup and drink odors: (1) while the cup odor was a distal cue that suppressed approach to the spout, the drink odor was primarily a proximal cue that suppressed consumption, and (2) the cup odor was less conditionable than the drink odor when the stimuli were conditioned alone. In Experiment 3, unlike the proximal saccharin taste, the proximal drink odor failed to potentiate the conditioning of a different distal odor. In Experiment 4, substantial dilution of the proximal drink's concentration resulted in both weaker conditioning of the stimulus alone, and potentiation of its conditioning by saccharin. The results suggest that the difference in the conditionability of the cup and drink odors, and not their status as distal and proximal cues, may be critical in the production of potentiation. Weakly conditionable target stimuli are uniquely potentiated by saccharin.     

Generalization gradients of inhibition following auditory discrimination learning

A more direct method than the usual ones for obtaining inhibitory gradients requires that the dimension of the nonreinforced stimulus selected for testing be orthogonal to the dimensions of the reinforced stimulus. In that case, the test points along the inhibitory gradient are equally distant from the reinforced stimulus. An attempt was made to realize this condition by obtaining inhibitory gradients along the frequency dimension of a pure tone after discrimination training in which the nonreinforced stimulus was a pure tone (or tones), and the reinforced stimulus was either white noise or the absence of a tone. The results showed that some degree of specific inhibitory control was exerted by the frequency of the tone, although the gradients were broad and shallow in slope.

A further experiment was conducted to see whether the modification of an excitatory gradient resulting from training to discriminate neighboring tones could arise from a simple interaction of inhibitory and excitatory gradients. The results indicated that it could not, since discrimination training produced a concentration of responding in the vicinity of the reinforced stimulus which cannot be derived from any plausible gradient of inhibition.

     

Recovery of an overshadowed association achieved by extinction of the overshadowing stimulus

Three experiments are reported in which conditioned lick suppression by water-deprived rats was used as an index of associative strength. In Experiment 1, overshadowing of a light by a tone was observed when the light-tone compound stimulus was paired with foot shock. After initial compound pairings, the tone-shock association was extinguished in one group of subjects. Subsequently, these animals demonstrated significantly higher levels of suppression to the light relative to a control group in which the tone had not been extinguished. Experiment 2 replicated this effect while failing to find evidence to support the possibilities that extinction presentations of the overshadowing tone act as retrieval cues for the light-shock association, or that, via second-order conditioning, the light-shock association is actually formed during extinction of the tone. Experiment 3 determined that the recovery from overshadowing observed in Experiments 1 and 2 was specific to the extinction of the overshadowing stimulus rather than the extinction of any excitatory cue. Collectively, these results suggest that the debilitated response to an overshadowed stimulus does not represent an acquisition failure, but rather the failure of an acquired association to be manifest in behavior.     

Forward and backward second-order Pavlovian conditioning in honeybees

Second-order conditioning (SOC) is the association of a neutral stimulus with another stimulus that had previously been combined with an unconditioned stimulus (US). We used classical conditioning of the proboscis extension response (PER) in honeybees (Apis mellifera) with odors (CS) and sugar (US). Previous SOC experiments in bees were inconclusive, and, therefore, we attempted to demonstrate SOC in the following three experiments: (Experiment 1) After differential conditioning (pairing odor A with US and presenting odor B without US), the bees experienced two pairs of partially overlapping odors, either a new odor C followed by a previously reinforced odor A (C-A) or a new odor C followed by a previously nonreinforced odor B (C-B). (Experiment 2) After differential conditioning, bees were presented with C-A or A-C. (Experiment 3) Bees were first presented with C-A or A-C before differential conditioning and were tested with odor C. We observed: (Experiment 1) 40% of the bees showed PER to the C-A presentation, but only 20% showed PER to the C-B presentation. (Experiment 2) 40% of the bees showed PER to the C-A presentation, while only 20% showed PER to the reversed sequence A-C. Experiments 1 and 2 showed that a previously reinforced odor can be a secondary reinforcer for excitatory SOC only with forward-pairing. (Experiment 3) PER toward C was lower (15%) in bees presented with A-C than with C-A (25%). This showed that backward SOC is not as effective as forward SOC. These results help to delineate different conditions that are critical for the phenomenon of SOC.     

Rhythmic context modulates foreperiod effects

Two experiments examined hypotheses about the roles of probabilistic uncertainty and rhythmic context on attentional preparation as reflected by choice response times (RTs) to the final tone of auditory sequences. Nonisochronous sequences with tone timings either arranged metrically or scrambled were linked with one of three different sequence-final time intervals, or foreperiods (FPs), which varied randomly from trial to trial. Two primary results emerged. First, RTs were faster to target tones ending metrical rhythms than to targets ending scrambled rhythms. Second, metrical contexts elicited RTs that increased with FP duration, whereas scrambled contexts elicited RTs that often decreased with FP duration, despite equivalent variability of time intervals in metrical and scrambled contexts. The results suggest that time relations implied by metrical rhythms systematically modulate preparatory responses to sequence-final FPs.     

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.     

Simultaneous conditioning in honeybees (Apis mellifera).

Honeybees (Apis mellifera) were classically conditioned with odor as conditioned stimulus (CS), sucrose as unconditioned stimulus (US), and proboscis extension as response. The purpose of Experiment 1 (Ns = 26 and 27) was to look for facilitation of forward conditioning by CS-US overlap, but rapid conditioning without overlap left little room for improvement. In 2 further experiments, CS and US were simultaneous, and response to odor alone was measured in subsequent tests. In Experiment 2, a Simultaneous group (N = 25) responded more to the training odor than did an Unpaired control group (N = 25). In Experiment 3, a differentially conditioned Simultaneous group (N = 29) responded more to an odor paired with sucrose in training (S+) than to an odor presented alone (S-). The implications of the results for the problem of the role of amount of reward in honeybee learning are considered.     

Naturalistic psychophysics: thresholds of ducklings (Anas platyrynchos) and chicks (Gallus gallus) to tones that resemble mallard calls

Neonatal ducklings and chickens were tested for responsiveness to a pulsing pure tone that was as similar as possible to the mallard maternal alarm call. It is known that ducklings momentarily cease vocalizing when they hear the alarm call and that chicks do the same when they hear pure tones. The duration of peep suppression can thus be used as a measure of whether subjects of either species heard the stimulus. Chicks might not be as sensitive as ducklings to a mallard alarm call because the signal is less significant to them. An adaptive or staircase procedure was used to estimate absolute thresholds, and group psychometric functions were reconstructed for each species from the trial-by-trial data. Ducklings had lower thresholds than chickens as well as steeper psychometric functions to this stimulus. The results suggest that more sensitive and consistent behavioral responses can be elicited by naturalistic sounds than by more arbitrary acoustic stimuli.     

Context-dependent olfactory learning monitored by activities of salivary neurons in cockroaches

Context-dependent discrimination learning, a sophisticated form of nonelemental associative learning, has been found in many animals, including insects. The major purpose of this research is to establish a method for monitoring this form of nonelemental learning in rigidly restrained insects for investigation of underlying neural mechanisms. We report context-dependent olfactory learning (occasion-setting problem solving) of salivation, which can be monitored as activity changes of salivary neurons in immobilized cockroaches, Periplaneta americana. A group of cockroaches was trained to associate peppermint odor (conditioned stimulus, CS) with sucrose solution reward (unconditioned stimulus, US) while vanilla odor was presented alone without pairing with the US under a flickering light condition (1.0 Hz) and also trained to associate vanilla odor with sucrose reward while peppermint odor was presented alone under a steady light condition. After training, the responses of salivary neurons to the rewarded peppermint odor were significantly greater than those to the unrewarded vanilla odor under steady illumination and those to the rewarded vanilla odor was significantly greater than those to the unrewarded peppermint odor in the presence of flickering light. Similar context-dependent responses were observed in another group of cockroaches trained with the opposite stimulus arrangement. This study demonstrates context-dependent olfactory learning of salivation for the first time in any vertebrate and invertebrate species, which can be monitored by activity changes of salivary neurons in restrained cockroaches.     

Stimulus information vs processing time in auditory pattern recognition

Previous studies have shown that pitch perception improves with increases in tone duration. An increase in tone duration increases the information available in the stimulus presentation. Other recent experiments have shown that sufficient perceptual processing time is also necessary for accurate pitch perception. The present study determines the relative contribution of stimulus information and processing time in an absolute pitch indentification task. The results indicate that processing time is more critical than stimulus information. The results are described accurately by a model that describes the perceptual process in terms of the information in the stimulus and the time the information is available for perceptual processing.     

Rats can make relative perceptual judgments about sequential stimuli

In their natural environment, animals often make decisions based on abstract relationships among multiple stimulus representations. Humans and other primates can determine not only whether a sensory stimulus differs from a remembered sensory representation, but also how they differ along a particular dimension. However, much remains unknown about how such relative comparisons are made, and which species share this capacity, in part because most studies of sensory-guided decision making have utilized instrumental tasks in which choices are based on very simple stimulus-response associations. Here, we used a two-stimulus-interval discrimination task to test whether rats could determine how two sequentially presented stimuli were related along the dimension of odor quality (i.e., what the stimulus smells like). At a central port, rats sampled and compared two odor mixtures that consisted of spearmint and caraway in different ratios, separated by a 2-4-s interval, and then entered the left or right reward port. Water was delivered at the left if the first mixture consisted of more spearmint than the second did, and at the right otherwise. We found that the difference in mixture ratio predicted choice accuracy. Control experiments suggest that rats were indeed basing their choices on a comparison of odor quality across mixtures and were not using associative strategies. This study is the first demonstration of the use of a sequential "more than versus less than" rule in rats and provides a well-controlled paradigm for studying abstract comparisons in a rodent model system.     

An inexpensive and automated method for presenting olfactory or tactile stimuli to rats in a two-choice discrimination task

An inexpensive and automated method for presentation of olfactory or tactile stimuli in a two-choice task for rats was implemented with the use of a computer-controlled bidirectional motor. The motor rotated a disk that presented two stimuli of different texture for tactile discrimination, or different odor for olfactory discrimination. Because the solid olfactory stimuli were placed outside the chamber in metal pods with a mesh at front for odor sampling, "washout" of odors between trials was not necessary. To avoid differential auditory cues from motor rotation, the stimuli were arranged such that on each trial the motor always rotated exactly one quarter revolution (in 1 s), left or right, to present the next stimulus at trial start. To illustrate the use of the equipment, 2 rats were trained on tactile discrimination and 2 rats on olfactory discrimination. The rats sampled the stimulus on the disk through a port on the back wall by sniffing at it (olfactory) or touching it (tactile). The task was a go-left/go-right discrimination with the stimulus on the disk being discriminative for which lever provided reinforcement. The rats reached a stable level above 90% correct after 21 and 32 training sessions for tactile and olfactory discrimination, respectively. The article outlines how the equipment was constructed from low-cost components. Inputs from and outputs to the equipment were implemented through the parallel port of a personal computer without the use of a commercial interface board. The method of automated and low-cost presentation of olfactory or tactile stimuli should be of use for a variety of experimental situations such as matching-to-sample and cross-modal discrimination.     

Referential coding contributes to the horizontal SMARC effect

The present study tested whether coding of tone pitch relative to a referent contributes to the correspondence effect between the pitch height of an auditory stimulus and the location of a lateralized response. When left-right responses are mapped to high or low pitch tones, performance is better with the high-right/low-left mapping than with the opposite mapping, a phenomenon called the horizontal SMARC effect. However, when pitch height is task irrelevant, the horizontal SMARC effect occurs only for musicians. In Experiment 1, nonmusicians performed a pitch discrimination task, and the SMARC effect was evident regardless of whether a referent tone was presented. However, in Experiment 2, for a timbre-judgment task, nonmusicians showed a SMARC effect only when a referent tone was presented, whereas musicians showed a SMARC effect that did not interact with presence/absence of the referent. Dependence of the SMARC effect for nonmusicians on a reference tone was replicated in Experiment 3, in which judgments of the color of a visual stimulus were made in the presence of a concurrent high- or low-pitched pure tone. These results suggest that referential coding of pitch height is a key determinant for the horizontal SMARC effect when pitch height is irrelevant to the task.     

Bilateral Ablation of Auditory Cortex in Mongolian Gerbil Affects Discrimination of Frequency Modulated Tones but not of Pure Tones

This study examines the role of auditory cortex in the Mongolian gerbil in differential conditioning to pure tones and to linearly frequency-modulated (FM) tones by analyzing the effects of bilateral auditory cortex ablation. Learning behavior and performance were studied in a GO/NO-GO task aiming at avoidance of a mild foot shock by crossing a hurdle in a two-way shuttle box. Hurdle crossing as the conditioned response to the reinforced stimulus (CR+), as false alarm in response to the unreinforced stimulus (CR−), intertrial activity, and reaction times were monitored. The analysis revealed no effects of lesion on pure tone discrimination but impairment of FM tone discrimination. In the latter case lesion effects were dependent on timing of lesion relative to FM tone discrimination training. Lesions before training in naive animals led to a reduced CR+ rate and had no effect on CR− rate. Lesions in pretrained animals led to an increased CR− rate without effects on the CR+ rate. The results suggest that auditory cortex plays a more critical role in discrimination of FM tones than in discrimination of pure tones. The different lesion effects on FM tone discrimination before and after training are compatible with both the hypothesis of a purely sensory deficit in FM tone processing and the hypothesis of a differential involvement of auditory cortex in acquisition and retention, respectively.     

Control of preference in children by conditioned positive reinforcement

A preference measure was employed with children to evaluate the conditioned positive reinforcing function of a stimulus that preceded reinforcement. A match-to-sample procedure was arranged in which subjects could respond to either the form or color dimension of a compound sample stimulus. Intermittent token reinforcement was provided equally for color and form matches. Two stimuli were employed (Stimulus A and Stimulus B), each consisting of a distinctive tone and colored light. One of these stimuli (the paired stimulus) preceded each token delivery, and the other did not (nonpaired stimulus). The paired stimulus was dependent upon each response to one match dimension, and the nonpaired stimulus followed each response to the other dimension. Three of the five subjects responded primarily to the dimension that was followed by the paired stimulus. This effect was obtained regardless of which stimulus (A or B) was paired and on which match dimension (color or form) the paired stimulus was dependent. These results were unaltered by discontinuing the nonpaired stimulus. The other two subjects demonstrated consistent preferences for the form dimension and Stimulus A, respectively.     

Effects of the periodic administration of odor or vibration on a 3-hr. vigilance task

The present study examined performance during 3 hr. of simulated sentry duty with and without the intermittent administration of low-level sensory stimuli (odor or vibration). For odor and control conditions, target-detection latency increased steadily over the course of the 3-hr. session. Administration of a tactile stimulus reduced the increase in detection latency compared to that found in odor and control conditions. For all conditions, there were no significant differences in target-detection frequency, shot accuracy, or friend-foe discrimination. Across all conditions, restlessness (motor activity) increased significantly the first hour and remained elevated for the rest of the session. Subjective measures of workload (NASA-TLX) indicated that the 3-hr. task rated high on physical demand, mental demand, frustration, and overall workload. These findings suggest that the intermittent delivery of a clearly detectable tactile stimulus can reduce reaction-time decrements that occur as time on task increases.