A comparison of responses and stimuli as time markers

A rat's behavior, as well as a stimulus, may be a time marker. But do they lead to similar performance? Eight rats were trained on a 20-s DRL procedure in which head-entry responses were time markers, i.e., each head-entry response indicated that food would not be delivered for 20 s. Concurrently, eight rats were trained on a control procedure in which light stimuli, yoked to the responses of a rat in the DRL procedure, were time markers, i.e., each light stimulus indicated that food would not be delivered for 20 s. A comparison of performance between the two groups showed a lower response rate in the DRL procedure than in the yoked control procedure. However, similar response patterns between the two groups were observed, suggesting that rats anticipated the food similarly with a stimulus or a response as the time marker.     

Changing rates of reinforcement perturbs the flow of time

This experiment tested the hypothesis that changes in rate of reinforcement affect the rate of an internal pacemaker as suggested by the Behavioral Theory of Timing (BeT: Killeen and Fetterman, 1988). Pigeons were trained to discriminate durations of 10 s and 20 s, and then exposed to higher or lower rates of freely delivered reinforcers. When returned to the discrimination task, judgments were reliably biased in the predicted directions: those returning from a richer context judged standard durations to be longer than did those returning from a poorer context. These results validate a key assumption of BeT, and provide an explanation of how changing tempos of life bias the perception of time.     

Further tests of the Scalar Expectancy Theory (SET) and the Learning-to-Time (LeT) model in a temporal bisection task

To contrast two models of timing, Scalar Expectancy Theory (SET) and Learning to Time (LeT), pigeons were exposed to a double temporal bisection procedure. On half of the trials, they learned to choose a red key after a 1s signal and a green key after a 4s signal; on the other half of the trials, they learned to choose a blue key after a 4-s signal and a yellow key after a 16-s signal. This was Phase A of an ABA design. On Phase B, the pigeons were divided into two groups and exposed to a new bisection task in which the signals ranged from 1 to 16s and the choice keys were blue and green. One group was reinforced for choosing blue after 1-s signals and green after 16-s signals and the other group was reinforced for the opposite mapping (green after 1-s signals and blue after 16-s signals). Whereas SET predicted no differences between the groups, LeT predicted that the former group would learn the new discrimination faster than the latter group. The results were consistent with LeT. Finally, the pigeons returned to Phase A. Only LeT made specific predictions regarding the reacquisition of the four temporal discriminations. These predictions were only partly consistent with the results.     

Odour representation in honeybee olfactory glomeruli shows slow temporal dynamics: an optical recording study using a voltage-sensitive dye

Stimulation with odours has been shown to elicit characteristic patterns of activated glomeruli in the antennal lobe (AL) of honeybees. In this study we show that these patterns are dynamic in a time window of 2-3 s after stimulus onset. We measured changes in the averaged membrane potential of all cells in the glomerular neuropil by optical imaging of the voltage-sensitive dye RH795 using a slow scan CCD camera (3 frames/s). The four substances 1-hexanol, hexanal, citral and clove-oil as well as the binary mixtures hexanol+hexanal and hexanol+citral were used as stimuli (2 s stimulus duration). We found that: (1) every odour elicited an odour-specific activity pattern, and conversely every glomerulus had a characteristic odour response profile; (2) some glomeruli had a tonic, some a phasic-tonic, and some a slow phasic response pattern; (3) the difference between the glomerular response patterns increased within 2 s of stimulus presentation, which suggests that odour representations became more characteristic over stimulus time; and (4) the responses to odorant mixtures were complex and glomerulus-dependent: some responses correspond to the sum of the compounds' responses, some to the response of one of the components.     

Correlations between activity of pallidal neurons and motor parameters

Abstract The relationships between the activity of 87 pallidal neurons and parameters characterizing motor performance were analysed quantitatively in six unrestrained cats performing a forelimb flexion movement controlled by a reaction-time paradigm. Three motor variables were considered: the static force exerted when the trigger stimulus occurred, the latency of the force change initiating the lever-release movement and the velocity of the force change. For all the cats, the latency of the change in force was the prevalent parameter in determining the movement onset occurrence. The single-unit activity was quantified on a trial-by-trial basis by measuring the mean firing rate in windows of fixed duration. Both normal (total) and partial linear correlation coefficients were used. Partial correlation coefficients were calculated between the mean firing rate during selected windows and each of the motor parameters. A total of 17% and 9% of all partial correlation coefficients tested ( n = 1566) was found to be significantly different from zero, respectively with p 0.05 and p 0.01. The mean percentages of explained variation, measured by the square of the correlation coefficient, were 17% ( p 0.05) and 21% ( p 0.01). The 46 pallidal neurons presenting an increase in discharge after the trigger stimulus could be classified in homogeneous subgroups with respect to the signs of the correlations. Representations of the motor programme underlying reaction-time performance evidenced at the pallidal level further substantiate a specific involvement of the basal ganglia in the latency of motor triggering.     

Coding "what" and "when" in the Archer fish retina

Traditionally, the information content of the neural response is quantified using statistics of the responses relative to stimulus onset time with the assumption that the brain uses onset time to infer stimulus identity. However, stimulus onset time must also be estimated by the brain, making the utility of such an approach questionable. How can stimulus onset be estimated from the neural responses with sufficient accuracy to ensure reliable stimulus identification? We address this question using the framework of colour coding by the archer fish retinal ganglion cell. We found that stimulus identity, “what”, can be estimated from the responses of best single cells with an accuracy comparable to that of the animal''s psychophysical estimation. However, to extract this information, an accurate estimation of stimulus onset is essential. We show that stimulus onset time, “when”, can be estimated using a linear-nonlinear readout mechanism that requires the response of a population of 100 cells. Thus, stimulus onset time can be estimated using a relatively simple readout. However, large nerve cell populations are required to achieve sufficient accuracy.

Authors Summary

In our interaction with the environment we are flooded with a stream of numerous objects and events. Our brain needs to understand the nature of these complex and rich stimuli in order to react. Research has shown ways in which a ‘what’ stimulus was presented can be encoded by the neural responses. However, to understand ‘what was the nature of the stimulus’ the brain needs to know ‘when’ the stimulus was presented. Here, we investigated how the onset of visual stimulus can be signalled by the retina to higher brain regions. We used archer fish as a framework to test the notion that the answer to the question of ‘when’ something has been presented lies within the larger cell population, whereas the answer to the question of ‘what’ has been presented may be found at the single-neuron level. The utility of the archer fish as model animal stems from its remarkable ability to shoot down insects settling on the foliage above the water level, and its ability to distinguish between artificial targets. Thus, the archer fish can provide the fish equivalent of a monkey or a human that can report psychophysical decisions.     

Conditioning and time representation in long short-term memory networks

Dopaminergic models based on the temporal-difference learning algorithm usually do not differentiate trace from delay conditioning. Instead, they use a fixed temporal representation of elapsed time since conditioned stimulus onset. Recently, a new model was proposed in which timing is learned within a long short-term memory (LSTM) artificial neural network representing the cerebral cortex (Rivest et al. in J Comput Neurosci 28(1):107–130, 2010). In this paper, that model’s ability to reproduce and explain relevant data, as well as its ability to make interesting new predictions, are evaluated. The model reveals a strikingly different temporal representation between trace and delay conditioning since trace conditioning requires working memory to remember the past conditioned stimulus while delay conditioning does not. On the other hand, the model predicts no important difference in DA responses between those two conditions when trained on one conditioning paradigm and tested on the other. The model predicts that in trace conditioning, animal timing starts with the conditioned stimulus offset as opposed to its onset. In classical conditioning, it predicts that if the conditioned stimulus does not disappear after the reward, the animal may expect a second reward. Finally, the last simulation reveals that the buildup of activity of some units in the networks can adapt to new delays by adjusting their rate of integration. Most importantly, the paper shows that it is possible, with the proposed architecture, to acquire discharge patterns similar to those observed in dopaminergic neurons and in the cerebral cortex on those tasks simply by minimizing a predictive cost function.     

Sensitization and habituation of the swimming behavior in ascidian larvae to light

Ascidian larvae of Ciona intestinalis change their photic behavior during the course of development. Newly hatched larvae show no response to a light stimulus at any intensity. At 4 hr after hatching, larvae were induced to start to swimming upon the cessation of illumination, and to stop swimming upon the onset of illumination. At a weaker light intensity (5.0 x 10(-3) J/m (2).s), the larvae showed similar responses to either a single stimulus or repeated stimuli of onset and cessation of light until 10 hr after hatching. At a stronger light intensity (3.2 x 10(-1) J/m(2).s), when the stimulus was repeated, they showed sensitization and habituation of the swimming response. At 3 hr after hatching the larvae failed to show any response to an initial stimulus at any intensity of light, but after several repeated stimuli (sensitization) they showed a swimming response at light intensities above 4.0 x 10(-2) J/m (2).s. At 5 hr and with intensity above 1.0 x 10 (-2) J/m(2).s, the larvae showed photoresponses to the first stimulus, but after several repetitions the larvae failed to stop swimming upon the onset of light (habituation). A repeated series of stimuli at stronger intensities of light caused greater habituation; this habituation was retained for about 1 min. Since the larval central nervous system in Ciona is comprised of only about 100 neurons, learning behavior in ascidian larvae should provide insights for a minimal mechanism of memory in vertebrates.     

Visual Responses of Crayfish Ocular Motoneurons: An Information Theoretical Analysis

Motoneuron responses were elicited by global visual motion and stepwise displacements of an illuminated stripe. Stimulus protocols were identical to those used in previous behavioral studies of compensatory eyestalk reflexes. The firing rates and directional selectivity of the motoneuron responses were measured with respect to four stimulus dimensions (spatial frequency, contrast, angular displacement and velocity). The directional selectivity of the motoneuron response was correlated to the previously measured gain of the reflex for each stimulus dimension. The information theoretical analysis is based upon Kullback-Leibler (K-L) distances which measure the dissimilarity of responses to different stimuli. K-L distances for single neurons are strongly influenced by the mean rate difference of the responses to any pair of stimuli. Because of redundancy, the joint K-L distances of pairs of neurons were less than the sum of the K-L distances of the individual neurons. Furthermore, the joint K-L distances were only weakly influenced by correlations among coactivated neurons. For most of the stimulus dimensions, the K-L distances of single motoneurons were not sufficient to account for the stimulus discriminations exhibited by the eyestalk reflex which typically required the summed output of 2 to 5 motoneurons. Thus the behaviorally relevant information is encoded in the motoneuron ensemble. The minimum time required to discriminate the direction of motion (the encoding window) for a single motoneuron is about 380 to 480 ms (including a 175 ms response latency) for stepwise displacements and up to 1.0 s for global motion. During this period a motoneuron fires 2 to 3 impulses.     

Respiratory muscle responses elicited by dorsal periaqueductal gray stimulation in rats

The periaqueductal gray matter is an essential neural substrate for central integration of defense behavior and accompanied autonomic responses. The dorsal half of the periaqueductal gray matter (dPAG) is also involved in mediating emotional responses of anxiety and fear, psychological states that often are associated with changes in ventilation. However, information regarding respiratory modulation elicited from this structure is limited. The present study was undertaken to investigate the relationship between stimulus frequency and magnitude on ventilatory pattern and respiratory muscle activity in urethane-anesthetized, spontaneously breathing rats. Electrical stimulation in the dPAG-recruited abdominal muscle activity increased ventilation and increased respiratory frequency by significantly shortening both inspiratory time and expiratory time. Ventilation increased within the first breath after the onset of stimulation, and the respiratory response increased with increasing stimulus frequency and magnitude. dPAG stimulation also increased baseline EMG activity in the diaphragm and recruited baseline external abdominal oblique EMG activity, normally quiescent during eupneic breathing. Significant changes in cardiorespiratory function were only evoked by stimulus intensities >10 microA and when stimulus frequencies were >10 Hz. Respiratory activity of both the diaphragm and abdominal muscles remained elevated for a minimum of 60 s after cessation of stimulation. These results demonstrate that there is a short-latency respiratory response elicited from the dPAG stimulation, which includes both inspiratory and expiratory muscles. The changes in respiratory timing suggest rapid onset and sustained poststimulus dPAG modulation of the brain stem respiratory network that includes expiratory muscle recruitment.     

Comparison of Scalar Expectancy Theory (SET) and the Learning-to-Time (LeT) model in a successive temporal bisection task

The present research tested the generality of the "context effect" previously reported in experiments using temporal double bisection tasks [e.g., Arantes, J., Machado, A. Context effects in a temporal discrimination task: Further tests of the Scalar Expectancy Theory and Learning-to-Time models. J. Exp. Anal. Behav., in press]. Pigeons learned two temporal discriminations in which all the stimuli appear successively: 1s (red) vs. 4s (green) and 4s (blue) vs. 16s (yellow). Then, two tests were conducted to compare predictions of two timing models, Scalar Expectancy Theory (SET) and the Learning-to-Time (LeT) model. In one test, two psychometric functions were obtained by presenting pigeons with intermediate signal durations (1-4s and 4-16s). Results were mixed. In the critical test, pigeons were exposed to signals ranging from 1 to 16s and followed by the green or the blue key. Whereas SET predicted that the relative response rate to each of these keys should be independent of the signal duration, LeT predicted that the relative response rate to the green key (compared with the blue key) should increase with the signal duration. Results were consistent with LeT's predictions, showing that the context effect is obtained even when subjects do not need to make a choice between two keys presented simultaneously.     

Learning performances in young horses using two different learning tests

To achieve optimal performance in equine sports as well as in leisure not only the physical abilities of the horse should be considered, but also the horse’s personality. Besides temperamental aspects, like emotionality, or the horse’s reactivity towards humans in handling situations, the learning ability of the horse is another relevant personality trait. To study whether differences in learning performance are consistent over time and whether individual learning performance differs between learning tests or is affected by emotionality, 39 young horses (Dutch Warmblood) were tested repeatedly in two learning tests. An aversive stimulus (AS) was used in one learning test (the avoidance learning test) and a reward was used in the other learning test (the reward learning test). During both learning tests behaviour as well as heart rate were measured. Each test was executed four times, twice when horses were 1 year of age, and twice when they were 2 years of age. Half of the horses received additional physical training from 6 months onwards. In both tests horses could be classified as either performers, i.e. completing the daily session, or as non-performers, i.e. returning to the home environment without having completed the daily session. There were some indications that emotionality might have caused non-performing behaviour, but these indications are not convincing enough to exclude other causes. Furthermore, there seem to be no simple relationships between measures of heart rate, behavioural responses putatively related to emotionality and learning performance. Horses revealed consistent individual learning performances within years in both tests, and in the avoidance learning test also between years. There was no significant correlation between learning performances in the avoidance learning test and the learning performances in the reward learning test. It is concluded that individual learning abilities are consistent over a short time interval for an avoidance learning test and a reward learning test and over a longer time for the avoidance learning test. Furthermore, results indicate that some horses perform better when they have to learn to avoid an aversive stimulus while others perform better when they are rewarded after a correct response. It is suggested that these differences may be relevant to design optimal individual training programmes and methods.     

Speech-evoked activity in primary auditory cortex: effects of voice onset time

Neural encoding of temporal speech features is a key component of acoustic and phonetic analyses. We examined the temporal encoding of the syllables /da/ and /ta/, which differ along the temporally based, phonetic parameter of voice onset time (VOT), in primary auditory cortex (A1) of awake monkeys using concurrent multilaminar recordings of auditory evoked potentials (AEP), the derived current source density, and multiunit activity. A general sequence of A1 activation consisting of a lamina-specific profile of parallel and sequential excitatory and inhibitory processes is described. VOT is encoded in the temporal response patterns of phase-locked activity to the periodic speech segments and by “on” responses to stimulus and voicing onset. A transformation occurs between responses in the thalamocortical (TC) fiber input and A1 cells. TC fibers are more likely to encode VOT with “on” responses to stimulus onset followed by phase-locked responses during the voiced segment, whereas A1 responses are more likely to exhibit transient responses both to stimulus and voicing onset. Relevance to subcortical speech processing, the human AEP and speech psychoacoustics are discussed. A mechanism for categorical differentiation of voiced and unvoiced consonants is proposed.     

High Repeatability of Anti-Predator Responses and Resting Metabolic Rate in a Beetle

Measures of repeatability are essential for understanding behavioral consistency and individual differences in behavior, i.e. animal personalities. We studied anti-predator responses of the yellow mealworm beetle (Tenebrio molitor) and performed behavioral tests in plastic containers representing a typical laboratory environment of T. molitor. Behavioral tests were repeated in Eppendorf test tubes where we also measured resting metabolic rate (RMR). Results show that the response latency to a threatening/startling stimulus, and the total time spent in the state of tonic immobility, correlated across the tests. The behavioral responses were repeatable and RMR covaried phenotypically with personality: we found a negative correlation between response latency time and time spent immobile, a positive correlation between response latency and RMR, and a negative correlation between RMR and total time spent immobile. These correlations were also similar across trials performed in the Eppendorf test tubes and the plastic containers.     

Determinants of tonic and phasic reactions in transswitching

Forty-eight college students were assigned randomly to four groups in a 2 X 2 factorial arrangement of phasic conditional stimuli (same vs. different) and tonic conditional stimuli (same vs. different) to receive 2 days of classical conditioning with a transswitching procedure. Tonic stimuli were a 5-minute projected white triangle or circle; phasic stimuli were a 5-second red or green square superimposed over the tonic stimuli. There were six tonic stimulus segments each day, separated by 20-second periods of no stimulus, three containing six trials of the phasic stimulus paired with shock and three containing six trials of the phasic stimulus alone, in the counterbalanced order. Tonic responding at the onset of the tonic stimuli or during brief periods following its onset were recorded, along with phasic responses to the phasic stimuli. Responses included magnitude of skin conductance responses, frequency of unelicited skin conductance responses, and tonic heart rate. Both skin conductance measures of responding to the tonic stimuli differentiated significantly between positive and negative tonic segments during Day 2, but only in the group with two different tonic stimuli and one phasic stimulus ("standard" transswitching). This supported the hypothesis that tonic stimulus differentiation would be absent when two different phasic stimuli were present. The heart rate data did not support this hypothesis, showing tonic differentiation in both groups with two tonic stimuli. Phasic differentiation controlled by the different phasic stimuli was observed on Day 1; on Day 2, phasic differentiation was present only in the group with two tonic and one phasic stimuli and the group with one tonic and two phasic stimuli.(ABSTRACT TRUNCATED AT 250 WORDS)     

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

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

Frequency-dependent excitability of "membrane" slow responses of Rabbit left atrial trabeculae in the presence of Ba2+ and high K+

Small trabeculae of rabbit left atrium immersed in TKBa solution (Tyrode with 10 mM K+ and 1 mM Ba2+) were used to study frequency dependence of "membrane" slow response excitability at long cycle lengths (greater than 1 s). In TKBa, stimuli generate graded, low- amplitude (2-15 mV) subliminal responses of variable long duration (up to 450 ms). A full all-or-none slow response is generated when a subliminal response depolarizes the membrane to about--35 mV. Subliminal response amplitude and rate of rise augment with stimulus intensity-duration product. For a fixed stimulus, the subliminal response is larger and faster at higher frequencies. Sudden changes in stimulus frequency or time course induce changes in subliminal response tha take four to eight cycles to attain steady state. For a fixed stimulus, slow response latency shortens progressively during the first few cycles after a sudden increase in frequency or when a rested preparation is excited (latency adaptation phenomenon, LAP). Slow response threshold stimulus requirements decrease during LAP (excitability hysteresis). The degree of excitability hysteresis is dependent on stimulation frequency and is more pronounced at higher frequencies. Frequency sensitivity of subliminal response (which causes frequency sensitivity of slow response excitability) is explained in terms of a transient state of enhancement set up by each stimulus. The enhanced state decays between stimuli with a half-time of approximately 4 s, thus allowing cumulative effects to become evident at rates above 0.1 Hz.     

Recalibration of the Multisensory Temporal Window of Integration Results from Changing Task Demands

The notion of the temporal window of integration, when applied in a multisensory context, refers to the breadth of the interval across which the brain perceives two stimuli from different sensory modalities as synchronous. It maintains a unitary perception of multisensory events despite physical and biophysical timing differences between the senses. The boundaries of the window can be influenced by attention and past sensory experience. Here we examined whether task demands could also influence the multisensory temporal window of integration. We varied the stimulus onset asynchrony between simple, short-lasting auditory and visual stimuli while participants performed two tasks in separate blocks: a temporal order judgment task that required the discrimination of subtle auditory-visual asynchronies, and a reaction time task to the first incoming stimulus irrespective of its sensory modality. We defined the temporal window of integration as the range of stimulus onset asynchronies where performance was below 75% in the temporal order judgment task, as well as the range of stimulus onset asynchronies where responses showed multisensory facilitation (race model violation) in the reaction time task. In 5 of 11 participants, we observed audio-visual stimulus onset asynchronies where reaction time was significantly accelerated (indicating successful integration in this task) while performance was accurate in the temporal order judgment task (indicating successful segregation in that task). This dissociation suggests that in some participants, the boundaries of the temporal window of integration can adaptively recalibrate in order to optimize performance according to specific task demands.     

gamma Interferon production appears to predict time of recurrence of herpes labialis

Peripheral blood mononuclear cells separated from the blood of 29 volunteers within 3 wk of the onset of recurrent oral herpes labialis spontaneously secreted IFN-gamma (and small amounts of IFN-beta) into the culture medium in varying amounts (mean = 77 U/ml, SE = 17). However, interferon could not be detected in the serum of 10 of these patients tested. In the group as a whole, peak levels of interferon were secreted in vitro at 6 to 20 days after the onset of herpes labialis. Serial studies in eight patients showed peak production in the second or third week, with subsequent decrease to undetectable levels at 6 wk. A strong correlation between peak supernatant interferon level and the time to the next recurrence of herpes labialis in each patient was noted (r = 0.82, p less than 0.0001). Herpes simplex antigen-stimulated mononuclear cell cultures from each patient produced a mixture of IFN-alpha and IFN-gamma. A less marked but still significant correlation was noted between the peak mixed interferon level and the inter-recurrence interval (r = 0.52, p less than 0.005). These results suggest that a recurrent herpes labialis acts as an in vivo stimulus to the induction of IFN-gamma-producing cells that circulate in peripheral blood. The IFN-gamma produced is either a direct determinant of frequency of recurrence of herpes or a quantitative marker for other cellular immune events determining frequency.     

Reduced variability of ongoing and evoked cortical activity leads to improved behavioral performance

Sensory responses of the brain are known to be highly variable, but the origin and functional relevance of this variability have long remained enigmatic. Using the variable foreperiod of a visual discrimination task to assess variability in the primate cerebral cortex, we report that visual evoked response variability is not only tied to variability in ongoing cortical activity, but also predicts mean response time. We used cortical local field potentials, simultaneously recorded from widespread cortical areas, to gauge both ongoing and visually evoked activity. Trial-to-trial variability of sensory evoked responses was strongly modulated by foreperiod duration and correlated both with the cortical variability before stimulus onset as well as with response times. In a separate set of experiments we probed the relation between small saccadic eye movements, foreperiod duration and manual response times. The rate of eye movements was modulated by foreperiod duration and eye position variability was positively correlated with response times. Our results indicate that when the time of a sensory stimulus is predictable, reduction in cortical variability before the stimulus can improve normal behavioral function that depends on the stimulus.