Perception of frequency, amplitude, and azimuth of a vibratory dipole source by the octavolateralis system of goldfish (Carassius auratus)

Goldfish (Carassius auratus) were conditioned to suppress respiration to a 40-Hz vibratory source and subsequently tested for stimulus generalization to frequency, stimulus amplitude, and position (azimuth). Animals completely failed to generalize to frequencies separated by octave intervals both lesser and greater than the CS. However, they did appear to generalize weakly to an aerial loudspeaker stimulus of the same frequency (40 Hz) after conditioning with an underwater vibratory source. Animals had a gradually decreasing amount of generalization to amplitude changes, suggesting a perceptual dimension of loudness. Animals generalized largely or completely to the same underwater source presented at a range of source azimuths. When these azimuths were presented at a transect of 3 cm, some animals did show decrements in generalization, while others did not. This suggests that although azimuth may be perceived more saliently at distances closer to a dipole source, perception of position is not immediately salient in conditioned vibratory source detection. Differential responding to test stimuli located toward the head or tail suggests the presence of perceptual differences between sources that are rostral or caudal with respect to the position of the animal or perhaps the head.     

Weak stimulus generalization using sinusoidal gratings: A cautionary note in animal psychophysics

Psychophysical techniques involving operant and classical conditioning are used commonly to determine the visual thresholds of nonverbal subjects. Typically, subjects are conditioned to respond to a particular stimulus, and once trained sufficiently, the subject’s sensitivity to this and similar stimuli can be determined by decreasing the stimulus intensity until the animal no longer responds. However, this assumes that the animal’s tendency to respond correlates with its sensory abilities. Stimulus generalization to sinusoidal gratings of different spatial frequencies was examined in goldfish. Subjects were classically conditioned to suppress respiration upon presentation of a sinusoidal grating. Animals’ learning curves to each spatial frequency grating were compared in order to determine whether the animals generalized across stimuli. Results indicate that fish show weak stimulus generalization across spatial frequencies. Also, the shape of the contrast sensitivity function, a common measure of the animal’s visual capabilities, is distorted by insufficient training. We conclude that when the goal is to obtain accurate data on visual capabilities, either subjects should be trained to each stimulus to be tested or their generalization gradients should be measured.     

Stimulus generalization of conditioned taste aversion in rats

Relatively little information is available regarding the intradimensional stimulus generalization of conditioned taste aversion (CTA). Experiment 1 employed a between-groups generalization test to examine the extent to which conditioned flavor aversion to one sucrose solution generalized to other concentrations of sucrose in adult rats. Evidence of a gradient of aversion was obtained. Because generalization gradients in other tasks have been found to flatten over a retention interval, Experiment 2 investigated the effects of delayed testing (2, 7, or 21 days) upon the slope of the generalization gradient. The generalization gradient flattened at the intervals, suggesting that subjects forgot the specific attributes of the conditioning concentration and avoided generalized stimuli as if they were the original CS. Experiment 3 used a long delay between taste and toxicosis to degrade the associative contingency and found no evidence that the generalization gradients found in the first two experiments could be explained in terms of enhanced neophobia due to poisoning. These findings provide further evidence (cf. A. W. Logue, 1979, Psychological Bulletin, 86, 276-296; M. Domjan, 1980, in J. S. Rosenblatt, R. A. Hinde, C. Beer, & M. Busnel (Eds.), Advances in the study of behavior, Vol. 11, New York: Academic Press) that CTA shares a number of similarities with other learning processes. Further, they illustrate that stimulus forgetting can be detected in a paradigm considered relatively immune to retention loss.     

Cardiovascular generalization and differentiation: The relationship between heart rate and blood pressure

The present study was designed to study generalization and differentiation and to check the relation between heart rate and blood pressure in cardiovascular conditioning. Five naive mongrel dogs were presented with 10 sec 1000 cps tone regularly reinforced with 0.5-sec shock to a left foreleg. Four other 10-sec tones (600, 800, 1200, 1400 cps), never reinforced, were administered in a randomized and counterbalanced fashion using a differential reinforcing technique. The response measures were heart rate and three indices of blood pressure,i.e., systolic, diastolic, and pulse pressures.“Initial response” of heart rate showed rapid tachycardia to all tones, with no generalization and differentiation among five tones.“Terminal response” clearly indicated generalization gradients that tachycardia to 800 and 1200 cps tones were higher than those to 600 and 1400 cps tones, respectively. Some differences were noted among systolic, diastolic, and pulse pressures, with pulse pressure showing a trend of generalization and differentiation in the“initial response.” No statistical difference, however, was obtained in blood pressure scores. Beat-by-beat analyses of correlations between heart rate and diastolic blood pressure showed that the correlations successively decreased during the“terminal response.” Overall correlations between heart rate and three indices of blood pressure were obtained, with the highest correlation between heart rate and pulse pressure. Considering the above results in relation to the classical conditioning theory, it was tentatively concluded that“initial response” includes non-conditional factors, while“terminal response” reflects mainly a true conditional response, particularly, at a long CS-US interval. This interpretation leads to the hypothesis that conditional response with a short CS-US interval may attain better conditioning scores in terms of the summation effect between a non-conditional factor and a true conditional factor.     

Stimulus factors in aversive controls: conditioned suppression after equal training to two stimuli

Pigeons trained to peck a key for food were periodically presented with tones ending with electrical shock until tone presentation consistently suppressed ongoing pecks. Shock was then discontinued and gradients of stimulus generalization were assessed by presenting tones with frequencies above, below, and at the frequencies of those used to develop conditioned suppression. When the training tones had frequencies at 670 and 1500 cps, resulting gradients were bi-modal with peak suppression at 670 and 1500 cps. Of the other test tones, 1000 cps produced the most suppression. When the training tones had frequencies at 450 and 2250 cps, bi-modal gradients were again obtained with peak suppression to the 450 and 2250 cps tones. Of the other test tones, 1000 cps produced the least suppression. These results support the hypothesis that generalized response tendencies summate.     

Stimulus generalization of suppression in rats following aversively motivated instrumental or Pavlovian training.

Following 100 or 300 avoidance training trials, instrumental subjects and their yoked Pavlovian counterparts were tested for generalization of lick suppression along the frequency dimension of the avoidance conditioned stimulus. Gradients of stimulus control were evident after 300 instrumental avoidance training trials, and additional intradimensional Pavlovian discrimination training further sharpened the gradients. After 100 trials, the yoked Pavlovian subjects suppressed more than their instrumental counterparts. However, with increased Pavlovian training, flatter gradients with decreased suppression were obtained. Results from a second experiment revealed that, whereas Pavlovian experience decreased suppression to the tone, subjects suppressed drinking in the presence of static, environmental cues. Data from both experiments supported interpretations that stress the role of response control over environmental events.     

Differential effects of omission contingencies on various components of Pavlovian appetitive conditioned responding in rats

Five experiments with rat subjects investigated the effects of omission and partial reinforcement contingencies on five individual behaviors evoked by visual and auditory conditioned stimuli paired with a food unconditioned stimulus. The effects of omission depended on the behavior on which that contingency was placed: One behavior was eliminated, one was unaffected, and three were reduced relative to the performance of yoked controls. Partial reinforcement resulted in lower frequencies of three behaviors and higher frequencies of two behaviors, compared with performance under consistent reinforcement. A partial reinforcement extinction effect was noted with one behavior but not with the others. These results are related to the possible role of instrumental conditioning contingencies in generating conditioned behavior in this appetitive conditioning preparation and to the independence of individual components of a complex conditioned response.     

Periodicity pitch perception and its upper frequency limit in cats

We have established a procedure for behavioral assessment of periodicity pitch perception in the cat. This procedure requires the training of cats to make a differential response to two different pure tones, and then testing for the transfer of learning when complex stimuli known to produce periodicity pitch sensations in humans are substituted for the pure tones. Complex stimuli consisted of three to five harmonically related frequencies of the same amplitude and phase. Eight cats were the subjects of these experiments. Two of them were controls. It is demonstrated here that cats do perceive periodicity pitch. We also used this procedure to measure the upper frequency limit of periodicity pitch perception for two cats. The frequencies tested were 100, 200, 300, 400, 600, 800, 900, and 1,200 Hz. The data indicate that the upper frequency limit of periodicity pitch perception in the cat is on the order of 600 to 800 Hz. There is a trend of decreasing prominence of the perception of periodicity pitch as the component frequencies are increased. These results are parallel to the human psychophysical data.     

Generalization gradients in human predictive learning: Effects of discrimination training and within-subjects testing

Generalization gradients have been investigated widely in animal conditioning experiments, but much less so in human predictive learning tasks. Here, we apply the experimental design of a recent study on conditioned fear generalization in humans (Lissek et al., 2008) to a predictive learning task, and examine the effects of a number of relevant procedural parameters drawn from the generalization literature in animal conditioning. Experiment 1 shows that prior discrimination learning and steady-state testing procedures sharpen the gradient; Experiment 2 shows that within-subjects testing of the range of generalization stimuli also sharpens the gradient. In addition, Experiment 2 shows that, in case of very flat initial generalization, an orderly gradient can reveal itself through differential rates of extinction learning. Finally, Experiment 2 also evidenced an orderly gradient of generalization-of-extinction. These results suggest that discrimination processes have an important effect on the generalization of predictive learning in humans, and highlight behavioral analogies between animal conditioning and human predictive learning.     

CS-specific gamma,theta, and alpha EEG activity detected in stimulus generalization following induction of behavioral memory by stimulation of the nucleus basalis

Tone paired with stimulation of the nucleus basalis (NB) induces behavioral memory that is specific to the frequency of the conditioned stimulus (CS), assessed by cardiac and respiration behavior during post-training stimulus generalization testing. This paper focuses on CS-specific spectral and temporal features of conditioned EEG activation. Adult male Sprague-Dawley rats, chronically implanted with a stimulating electrode in the NB and a recording electrode in the ipsilateral auditory cortex, received either tone (6kHz, 70dB, 2s) paired with co-terminating stimulation of the nucleus basalis (0.2s, 100Hz, 80-105 microA, ITI approximately 45s) or unpaired presentation of the stimuli (approximately 200 trials/day for approximately 14 days). CS-specificity was tested 24h post-training by presenting test tones to obtain generalization gradients for the EEG, heart rate, and respiration. Behavioral memory was evident in cardiac and respiratory responses that were maximal to the CS frequency of 6kHz. FFT analyses of tone-elicited changes of power in the delta, theta, alpha, beta1, beta2, and gamma bands in the paired group revealed that conditioned EEG activation (shift from lower to higher frequencies) was differentially spectrally and temporally specific: theta, and alpha to a lesser extent, decreased selectively to 6kHz during and for several seconds following tone presentation while gamma power increased transiently during and after 6kHz. Delta exhibited no CS-specificity and the beta bands showed transient specificity only after several seconds. The unpaired group exhibited neither CS-specific behavioral nor EEG effects. Thus, stimulus generalization tests reveal that conditioned EEG activation is not unitary but rather reflects CS-specificity, with band-selective markers for specific, associative neural processes in learning and memory.     

Latent inhibition and stimulus generalization of the classically conditioned nictitating membrane response in rabbits (Oryctolagus cuniculus) following dorsal hippocampal ablation.

Rabbits received 0 to 450 exposures of a tone conditioned stimulus (CS) prior to classical defensive conditioning of the nicitating membrane response based on an infraorbital eye shock unconditioned stimulus. Tone preexposure resulted in retarded conditioning in normal rabbits. This latent inhibition effect was not present in animals with bilateral dorsal hippocampectomy produced by aspiration. Control animals with bilateral neocortical and callosal aspiration lesions demonstrated a latent inhibition effect similar to that shown by normal nonoperated animals. The failure of CS preexposure to retard conditioning in hippocampal rabbits was not due to differences in threshold of the conditioned response to the CS or to differences in response mechanisms as determined by tests of habituation and dishabituation of the unconditioned response. A subsequent experiment employed combined-cue summation tests to confirm the fact that preexposure did not endow the tone with conditioned as well as latent inhibitiory properties. Finally, tests of stimulus generalization along the auditory frequency dimension indicated flatter relative gradients for hippocampals than for nonoperated controls, with cortical controls in between. These findings were discussed in terms of Douglas' model of hippocampal function.     

Maintained generalization testing of conditioned suppression

This study concerns the use of a multiple stimulus discrimination procedure for producing data on the generalization of conditioned suppression. Four rats were maintained on a variable interval schedule of milk reinforcement in the presence of five stimuli varying in auditory click rate. When response rates were stable, electric shock was regularly paired with the termination of one of the click stimuli. For two rats the shock was paired with the slowest click rate, and for two rats shock was paired with the fastest click rate. The VI schedule remained in effect. Plots of the relative rates of response to each of the five stimuli yielded concave gradients for both animals suppressed at the slowest click rate, and flat gradients with a sharp drop at the warning stimulus for both animals suppressed at the fastest click rate. When the warning stimuli were reversed for both pairs of subjects, both gradient forms were reproduced. The present procedure was contrasted with procedures used by other investigators.     

Analysis of response rates during stimulus generalization

In the presence of one click frequency, the presses of two hungry rats on one of two levers were reinforced with food on variable-interval schedules; in the presence of a different click frequency, presses on the other lever were reinforced. In stimulus generalization tests, a variety of click frequencies were presented and reinforcement withheld. The test stimuli were found to exert control over which of the two levers the rats pressed, but not over the rate of pressing the selected lever. The results were interpreted as further evidence that intermediate rates in generalization gradients may be the result of the alternation of several distinct behavior patterns.     

The decrement in conditioned fear with increased trials of simultaneous conditioning is not specific to the simultaneous procedure

Three experiments using a conditioned punishment paradigm with rat subjects examined the possibility that the nonmonotonic acquisition function previously found to characterize simultaneous conditioning was due to the noninformative nature of the conditioned stimulus (CS). In Experiment 1 the suppressive effects of a CS previously presented with an unconditioned stimulus (US) in a simultaneous and forward (informative) manner were compared following 20 and an additional 60 conditioning trials. Excitatory conditioning similarly diminished with increased trials for both the simultaneous and forward procedures. Experiment 2 employed a between-groups design. Simultaneous, forward, and trace conditioning procedures were compared following 20 or 100 trials. Each of the three 100-trial groups showed less resistance to extinction than their 20-trial counterparts. Experiment 3 determined that the decrement in excitatory conditioning for the 100-trial groups was not due to the greater number of US presentations, per se, but rather to the number of CS-US pairings. The nonmonotonic acquisition function observed with all three conditioning procedures indicated that informational factors were not responsible for the decrement observed in simultaneous conditioning. The pattern of results suggested that subjects receiving extended conditioning trials were better able to discriminate between training and testing.     

Consolidation and long-term retention of an implanted behavioral memory

Hypothesized circuitry enabling information storage can be tested by attempting to implant memory directly in the brain in the absence of normal experience. Previously, we found that tone paired with activation of the cholinergic nucleus basalis (NB) does induce behavioral memory that shares cardinal features with natural memory; it is associative, highly specific, rapidly formed, consolidates and shows intermediate retention. Here we determine if implanted memory also exhibits long-term consolidation and retention. Adult male rats were first tested for behavioral responses (disruption of ongoing respiration) to tones (1-15 kHz), yielding pre-training behavioral frequency generalization gradients. They next received 3 days of training with a conditioned stimulus (CS) tone (8.0 kHz, 70 dB, 2s) either paired (n=7) or unpaired (n=6) with moderate electrical stimulation of the nucleus basalis (～ 65 μA, 100 Hz, 0.2s, co-terminating with CS offset). Testing for long-term retention was performed by obtaining post-training behavioral frequency generalization gradients 24h and 2 weeks after training. At 24h post-training, the Paired group exhibited specific associative behavioral memory, manifested by larger responses to the CS frequency band than the Unpaired group. This memory was retained 2 weeks post-training. Moreover, 2 weeks later, the specificity and magnitude of memory had become greater, indicating that the implanted memory had undergone consolidation. Overall, the results demonstrate the validity of NB-implanted memory for understanding natural memory and that activation of the cholinergic nucleus basalis is sufficient to form natural associative memory.     

Flicker contrast sensitivity in normal and specifically disabled readers

Temporal contrast sensitivity for counterphase flicker was determined for specifically disabled and normal readers to investigate whether the two groups differ in the functioning of their transient systems. In experiment 1, temporal contrast sensitivity was measured over a range of temporal frequencies with a spatial frequency of 2 cycles deg-1. Disabled readers were less sensitive than the control subjects at all temporal frequencies. In experiment 2, temporal contrast sensitivity was measured at a temporal frequency of 20 Hz over a range of spatial frequencies. Disabled readers were less sensitive than the controls at all spatial frequencies, with the differences between the groups increasing as spatial frequency increased. Both these findings are interpreted as supporting the hypothesis of a transient-system deficit in the visual systems of disabled readers.     

Generalization of contextual fear as a function of familiarity: the role of within- and between-context associations

Three experiments in rats investigated the generalization of conditioned fear from one context (B) to both a preexposed context (A) and a novel context (C). In each experiment, when the conditioning context (B) had been preexposed, there was greater generalization to context A than to context C; but when B was novel at the outset of conditioning this difference between A and C was not observed. The implications of these results for associative treatments of the development of contextual memories are evaluated.     

Vibrotactile frequency discrimination at short durations

This experiment investigated how frequency discrimination of a sinusoidal, mechanical vibration applied to the tip of the right index finger is affected by shortening the duration of the stimuli from 200 ms to 30 ms. Using a standard stimulus of 100 Hz at 30 dB above threshold, seven comparison frequencies (at intervals of 10 Hz) were judged as higher or lower in frequency according to the method of constant differences. Vibrotactile stimuli were matched for subjective intensity across both frequency and duration. Difference limens for vibrotactile frequency were found to decline slightly from 200 ms to 50 ms (attributable to practice) and to increase noticeably at 30 ms. This result is discussed in relation to the seemingly contradictory results for auditory pitch discrimination.     

Effects of peripheral circular contours on dynamic spatial orientation

The rod-and-frame effect (RFE) was investigated with the use of a frame that oscillated about an axis at its center at five different frequencies, ranging from .013 to .213 Hz. The resultant RFE shifted continuously with the roll motion of the frame, and it was significantly larger at the lowest frequency (.013 Hz) than under comparable static conditions. The dynamic RFE was lowest at the higher oscillation frequencies. Oscillatory roll vection--apparent self-motion--was reported by 3 of the 9 subjects when the frame was oscillating at its highest frequency (.213 Hz). The subjects yielded large increases in the RFE during the sessions with reports of vection. Surrounding the kinetic frame with a circular contour eliminated all reports of vection and significantly interacted with frequency to reduce the RFE--but only at low frequencies. The reduction amounted to 21.2% averaged over all 9 subjects at the three lowest frequencies. A surrounding contour, therefore, suppressed low-frequency kinetic visual orientation information that might otherwise have produced larger changes in apparent self-orientation and perceived vertical. Vection-sensitive subjects differed from nonvection subjects by exhibiting (1) a high-frequency fall-off in real-motion gain, (2) a high-frequency enhancement in illusory-motion gain, and (3) only a small and nonsignificant increase in illusory-movement phase lag with increases in frequency.     

CLASSICAL CONDITIONING OF DECREASES IN HUMAN SYSTOLIC BLOOD PRESSURE1

Decrease in human systolic blood pressure of 4.35 mm Hg (range: 0 to 12 mm Hg) were classically conditioned in normal and hypertensive subjects using a delayed conditioning paradigm in which a 30-sec auditory stimulus (the conditioned stimulus) was followed immediately by tilting the subject 15° head-down to elicit small decreases in blood pressure. Conditioning occurred within five trials. A control group demonstrated that sensitization of the reflex by repeated tilting could not account for the blood-pressure decreases associated with the conditioned stimulus in experimental subjects.