Differences between simultaneous and blocked training detected by a transfer test

Secondary data analysis was used to compare responding early on a transfer test from rats previously trained simultaneously or successively on multiple temporal discriminations for the same number of trials [Guilhardi, P., Church, R.M., 2005 a. Dynamics of temporal discrimination. Learn. Behav., 33, 399-416]. Three fixed intervals (30, 60, and 120 s) were signaled by three stimuli (light, noise, and clicker). Twelve rats were trained with the three stimulus-interval pairs intermixed on each experimental session (simultaneous condition); 12 other rats were trained in successive blocks of 10 sessions on each pair (blocked condition). Then, all rats had a transfer test in which all three stimulus-interval pairs were presented intermixed on each session. Rats in the simultaneous and blocked condition responded similarly during training, but differently during early stages of the transfer test. One possibility is that rats in the blocked condition were controlled by the previous interval, not by the current stimulus. These results challenge the usual assumptions from models of timing and conditioning that both simultaneous and blocked training produce learning of the associations between stimulus and interval in a multiple interval training task.     

Shifting attention within memory representations involves early visual areas

Prior studies have shown that spatial attention modulates early visual cortex retinotopically, resulting in enhanced processing of external perceptual representations. However, it is not clear whether the same visual areas are modulated when attention is focused on, and shifted within a working memory representation. In the current fMRI study participants were asked to memorize an array containing four stimuli. After a delay, participants were presented with a verbal cue instructing them to actively maintain the location of one of the stimuli in working memory. Additionally, on a number of trials a second verbal cue instructed participants to switch attention to the location of another stimulus within the memorized representation. Results of the study showed that changes in the BOLD pattern closely followed the locus of attention within the working memory representation. A decrease in BOLD-activity (V1-V3) was observed at ROIs coding a memory location when participants switched away from this location, whereas an increase was observed when participants switched towards this location. Continuous increased activity was obtained at the memorized location when participants did not switch. This study shows that shifting attention within memory representations activates the earliest parts of visual cortex (including V1) in a retinotopic fashion. We conclude that even in the absence of visual stimulation, early visual areas support shifting of attention within memorized representations, similar to when attention is shifted in the outside world. The relationship between visual working memory and visual mental imagery is discussed in light of the current findings.     

Timescale dependence in a conditional temporal discrimination procedure

A conditional temporal discrimination procedure was used to test the scope and generality of the principle of timescale invariance (TSI). Rats were trained to make different temporal discriminations following four different auditory stimuli. After shorter stimuli, rats were reinforced for pressing the left lever, and after longer stimuli, rats were reinforced for pressing the right lever. The four auditory stimuli (a pure tone, pulsed tone, click, and white noise) were each presented for different pairs of durations (ranging from 2 vs. 8s to 32 vs. 128 s) within a single session, but the ratio between the shorter and longer duration was maintained constant at 1:4. With the shortest pair of durations (2 vs. 8s), rats showed a clear violation of TSI in both overall and relative response rates. Rats started responding later and persisted relatively longer on the left lever when the shorter duration was only 2s. We interpret these results with regards to a framing hypothesis, whereby TSI does not apply to relatively short durations when animals are simultaneously trained with wide ranges of intervals.     

Interstimulus interval as a discriminative stimulus: Evidence of the generality of a novel asymmetry in temporal discrimination learning

Three appetitive conditioning experiments with rats examined temporal discrimination learning within Pavlovian conditioning trials. In all experiments, the duration of a feature white noise stimulus signaled whether or not a subsequent 10-s target tone would be reinforced. In Experiment 1, the feature durations were 4 and 1 min. For one group of rats (Group 4+/1−), 4 min of noise signaled that the tone would be reinforced and 1 min of noise signaled that the tone would not be reinforced. A second group (Group 1+/4−) was trained with the reverse contingency. The results showed a clear asymmetry in temporal discrimination learning: rats trained with 4+/1− (Long+/Short−) learned the discrimination readily (responding more in the tone on reinforced than on nonreinforced trials), whereas rats trained with 1+/4− (Short+/Long) did not. In Experiment 2, the feature durations were shortened to 60 and 15 s. Due to strong excitatory conditioning of the 15-s feature, the reverse asymmetry was observed, with the Short+/Long− discrimination learned more readily than the Long+/Short− discrimination. However, Experiment 3 demonstrated that the original Long+/Short− advantage could be recovered while using 60− and 15-s feature durations if the excitatory conditioning of the feature was reduced by including nonreinforced feature trials. The results support previous research involving the timing of intertrial intervals and are consistent with the temporal elements hypothesis which holds that the passage of time is encoded as a series of hypothetical stimulus elements.     

Occasion setting in Pavlovian ambiguous target discriminations

Rats were trained in a Pavlovian serial ambiguous target discrimination, in which a target cue was reinforced if it was preceded by one stimulus (P-->T+) but was not reinforced if it was preceded by another stimulus (N-->T-). Test performance indicated that stimulus control by these features was weaker than that acquired by features trained within separate serial feature positive (P-->T+, T-) and serial feature negative (N-->W-, W+) discriminations. The form of conditioned responding and the patterns of transfer observed suggested that the serial ambiguous target discrimination was solved by occasion setting. The data are discussed in terms of the use of retrospective coding strategies when solving Pavlovian serial conditional discriminations, and the acquisition of special properties by both feature and target stimuli.     

Neuronal responses related to long-term recognition memory processes in prefrontal cortex

Much evidence indicates that prefrontal cortex plays an important role in long-term recognition memory processes. Here, we report primate prefrontal neuronal responses carrying information necessary for long-term visual recognition memory. The responses of many neurons signaled stimulus familiarity even when the period over which stimuli had to be remembered extended to 24 hr. Such responses occurred frequently in ventromedial, orbitofrontal, and anterior cingulate but not dorsolateral prefrontal cortex. Prefrontal information processing, as indicated by the response latencies, started after that in inferior temporal cortex and might be related to retrieval processes, as responses were typically larger for familiar than for novel stimuli.     

Effects of delays on human performance on a temporal discrimination procedure: evidence of a choose-short effect

Studies of temporal discrimination in non-human subjects have reliably shown a choose-short effect: higher matching accuracy on short-duration-sample trials than on long-duration-sample trials. This effect occurs as a function of increasing the delay between the onset of sample and comparison stimuli in a delayed matching-to-sample procedure. The present experiment investigated whether the choose-short effect could be demonstrated in human subjects under conditions which paralleled those used with non-human subjects. Subjects responded under a discrete-trial procedure in which they were required to push one of two buttons depending on the duration of a sample stimulus (a blue square on a computer monitor). Delays (0, 8, 16, and 32s) separated sample and comparison stimuli (two white boxes) and were tested both within and across several sessions. Intermediate durations (probe stimuli between 2 and 4s) were also presented. The addition of a delay between the sample and comparison stimuli produced a bias to judge intervals as short when the 8 and 32-s delays were tested across sessions and when the 0, 16, and 32-s delays were tested within the same session. Thus, the choose-short effect was produced in human subjects using the interval bisection procedure regardless of delay length.     

Memory strength and repetition suppression: multimodal imaging of medial temporal cortical contributions to recognition

Declarative memory permits an organism to recognize stimuli that have been previously encountered, discriminating them from those that are novel. One basis for recognition is item memory strength, which may support the perception of stimulus familiarity. Though the medial temporal lobes are known to be critical for declarative memory, at present the neural mechanisms supporting perceived differences in memory strength remain poorly specified. Here, functional MRI (fMRI) and anatomically constrained magnetoencephalography (MEG) indexed correlates of graded memory strength in the human brain, focusing on medial temporal cortex. fMRI revealed a decrease in medial temporal cortical activation that tracked parametric levels of perceived memory strength. Anatomically constrained MEG current estimates revealed that strength-dependent signal reductions onset within 150-300 ms. Memory strength appears to be rapidly signaled by medial temporal cortex through repetition suppression (activation reductions), providing a basis for the subjective perception of stimulus familiarity or novelty.     

Modality and intermittency effects on time estimation

The modality of a stimulus and its intermittency affect time estimation. The present experiment explores the effect of a combination of modality and intermittency, and its implications for internal clock explanations. Twenty-four participants were tested on a temporal bisection task with durations of 200-800 ms. Durations were signaled by visual steady stimuli, auditory steady stimuli, visual flickering stimuli, and auditory clicks. Psychophysical functions and bisection points indicated that the durations of visual steady stimuli were classified as shorter and more variable than the durations signaled by the auditory stimuli (steady and clicks), and that the durations of the visual flickering stimuli were classified as longer than the durations signaled by the auditory stimuli (steady and clicks). An interpretation of the results is that there are different speeds for the internal clock, which are mediated by the perceptual features of the stimuli timed, such as differences in time of processing.     

Response strategies during acquisition of serial learning in pigeons (Columba livia)

Pigeons were exposed on fixed duration sessions to four chromatic stimuli and trained to perform a 2-item sequence. A detailed analysis of incorrect responses performed on each trial prior to completion of the sequence led to distinguish three learning stages. Already on initial sessions animals responded significantly more frequently to relevant stimuli than to irrelevant ones indicating stimulus control by salience. During an intermediary stage, responding to item 2 increased significantly and animals started an increasing number of trials with item 2, suggesting stimulus control by recency. On the final session the correct sequence outnumbered significantly the reverse sequence, suggesting control by order, while recency and salience effects were present to a lesser extent. Repeated pecks, i.e. consecutive pecks on the same cue, occurred on the four cues on the first session and decreased significantly on the last session for irrelevant colours. In addition, when key pecks were followed by a brief sound signal, repeats on relevant stimuli decreased while efficient colour pecks (i.e. when repeats were ignored) on these stimuli increased. These findings are discussed in the light of those obtained in traditional chaining paradigms.     

Differential effects of empty and filled intervals on duration estimation by pigeons: tests of an attention-sharing explanation

Pigeons were trained in a within-subjects design to discriminate durations of an empty interval and a filled interval. Even when different stimuli were used to mark empty intervals and to signal filled intervals, pigeons judged empty intervals to be longer than equal-length filled intervals. This timing difference was not a result of pigeons timing marker duration on empty interval trials. Increasing marker duration did not produce an overestimation of the empty time intervals. It was suggested that this timing difference could be due to a reduction in attention to temporal processing on filled interval trials when visual stimuli are used. Consistent with this hypothesis, it was found that empty intervals were judged longer than filled intervals when testing occurred in a darkened test room, but not when the test room was illuminated. In addition, no timing difference was observed when different auditory stimuli were used as markers for empty intervals and as signals for filled intervals.     

Factors that influence negative summation in a spatial-search task with pigeons

A variant of the standard conditioned inhibition procedure was used to evaluate landmark-based spatial search in a touchscreen preparation. Pigeons were given compound trials with one landmark (A) positioned in a consistent spatial relationship to a hidden goal and another landmark (B) positioned randomly with respect to A and the hidden goal (AB+). On half of the non-reinforced inhibitory trials, A was paired with landmark X (AX-) and on the remaining trials B was paired with Y (BY-). All subjects were also given reinforced trials with a transfer excitor (T+). During conditioned inhibition training, subjects showed no change in overall responding during AX- trials but did show a decrease in the number of pecks to the goal location signaled by A. During non-reinforced summation tests with landmark T, X had a greater suppressive effect than did Y on overall responding but the percentage of pecks at the goal did not differ unless X was positioned near the expected goal signaled by T. These data demonstrate that the effectiveness of a stimulus trained as an inhibitor is dependent on the strength of the association between its training excitor (A) and the US, as well as, the spatial arrangement of stimuli during testing.     

Effects of training history on resurgence in humans

This experiment assessed the effects of training history on resurgence in three college students. Four-choice arbitrary-matching-to-sample trials occurred in two components of a multiple schedule. An A1 or A2 sample stimulus and four (B) comparison stimuli occurred on AB trials, and a C1 or C2 sample stimulus and four (D) comparison stimuli occurred on CD trials. By the end of training, accuracy and latency measures were comparable across separate discriminations, selecting B2 in the presence of A2 and selecting D2 in the presence of C2, despite a lengthier training correlated with the former discrimination. Next, in the presence of A2 and C2, respectively, responses to B2 and D2 were extinguished and responses to B3 and D3 were reinforced. These responses to B3 and D3 then were extinguished in a final condition. In this final condition, resurgence to B2 occurred for each participant, whereas resurgence to D2 occurred for only one participant. Thus, there was greater resurgence of the discrimination with the lengthier training history, despite the discriminations being similar in terms of accuracy and latency before extinction. This result, therefore, can be classified as a latent, or remote, behavioral history effect.     

Stimulus coding rules for perceptual learning

Perceptual learning of visual features occurs when multiple stimuli are presented in a fixed sequence (temporal patterning), but not when they are presented in random order (roving). This points to the need for proper stimulus coding in order for learning of multiple stimuli to occur. We examined the stimulus coding rules for learning with multiple stimuli. Our results demonstrate that: (1) stimulus rhythm is necessary for temporal patterning to take effect during practice; (2) learning consolidation is subject to disruption by roving up to 4 h after each practice session; (3) importantly, after completion of temporal-patterned learning, performance is undisrupted by extended roving training; (4) roving is ineffective if each stimulus is presented for five or more consecutive trials; and (5) roving is also ineffective if each stimulus has a distinct identity. We propose that for multi-stimulus learning to occur, the brain needs to conceptually “tag” each stimulus, in order to switch attention to the appropriate perceptual template. Stimulus temporal patterning assists in tagging stimuli and switching attention through its rhythmic stimulus sequence.     

Prospective timing under dual-task paradigms: attentional and contextual-change mechanisms

The effect of a concurrent memory task on prospective time estimates by human participants was investigated in two experiments. The objective was to isolate task effects from those of participant timing strategy (self-paced counting) and number of contextual changes during the temporal stimulus. Accordingly, self-paced counting was suppressed by requiring participants to perform a word-reading task during the temporal stimuli, while number of stimulus changes presented during temporal stimuli was controlled. Presence versus absence of the concurrent memory task was manipulated in Experiment 1, and instruction to focus on timing or to focus on memory was manipulated in Experiment 2. There was no significant effect of presence versus absence of the concurrent memory task on time estimates; however, time estimates were shorter when participants were instructed to focus on memory versus timing. In both experiments, time estimates were positively correlated with participants' estimates of the number of words presented during the interval, even though number of words presented was invariant. These findings were generally consistent with resource-allocation attentional accounts of concurrent task effects; however, support for a contextual-change model of timing was also obtained.     

Cognitive tactics of Bengalese finch (<Emphasis Type="Italic">Lonchura striata </Emphasis>var. <Emphasis Type="Italic">domestica</Emphasis>) for song discrimination in a go/no-go operant task

Twelve Bengalese finches (Lonchura striata var. domestica) were trained to discriminate between a conspecific and a heterospecific song in a go/no-go operant task. Training the birds to go for the conspecifics song or to go for the heterospecific song required the same number of training sessions. Nine possible cognitive tactics could be used to solve this task, but probe tests revealed that the birds used only four. Six birds memorized only the “no-go” stimulus and responded to the rest of the stimuli (no-go memory), two birds classified songs according to the species category (open-ended categorization), one bird memorized both of the training stimuli but responded by chance to the probe stimuli (rote categorization), and two birds combined open-ended and rote categorization tactics (combined categorization). These tactics were related to the number of sessions needed to reach the species-discrimination criterion. Our results suggest that investigators should consider individual cognitive tactics and the pitfalls of go/no-go discrimination when interpreting the results of operant discrimination tasks.     

Neural correlates of knowledge: stable representation of stimulus associations across variations in behavioral performance

Behavioral responses to a sensory stimulus are often guided by associative memories. These associations remain intact even when other factors determine behavior. The substrates of associative memory should therefore be identifiable by neuronal responses that are independent of behavioral choices. We tested this hypothesis using a paired-associates task in which monkeys learned arbitrary associations between pairs of visual stimuli. We examined the activity of neurons in inferior temporal cortex as the animals prepared to choose a remembered stimulus from a visual display. The activity of some neurons (22%) depended on the monkey's behavioral choice; but for a novel class of neurons (54%), activity reflected the stimulus that the monkey was instructed to choose, regardless of the behavioral response. These neurons appear to represent memorized stimulus associations that are stable across variations in behavioral performance. In addition, many neurons (74%) were modulated by the spatial arrangement of the stimuli in the display.     

The effect of study-test modalities on the remembrance of subjective duration from long-term memory

It was examined whether stimulus modality (auditory vs. visual) affects the retrieval of subjective duration from memory. In two experiments the temporal generalization paradigm was used. Participants had to decide whether the previously learned standard duration (400 ms) occurred in the context of comparison stimuli. Two major results were found. (1) Discrimination was more accurate if the training and testing stimuli were of the same modality than if they were of opposite modalities. (2) If both modality of learning and modality of testing were different, subjects systematically underestimated the test durations, i.e. temporal generalization gradients (the proportion of identifications of a stimulus as the standard, plotted against stimulus duration) shifted to the right. The observed shift is interpreted as a result of a delayed timing process.     

Reaction times in discriminations of varying difficulty: decision modulated by arousal

Pigeons discriminated the hue of a spot of light that appeared in discrete trials. A green spot always signaled food ("green S+") and so did a red spot of constant hue ("red S+"), but on most trials a different red hue appeared and no food was given for pecks ("red S-"). The hue of red S- stayed the same during blocks of up to twelve sessions, but it changed from block to block. During a final group of sessions red S- was omitted and the percent reinforcement to the two S+ stimuli was varied. Major findings were that (1) percent response (Pct(R)) to S- varied with S+/S- similarity, describing a typical ogival psychometric function; (2) Reaction times (RTs) to both red and green S+s were minimal when the red discrimination was impossible (that is, when red S+ and S- were identical); (3) RTs to red S+ were greatest during discriminations of intermediate difficulty; (4) as Pct(R) declined during the learning of a difficult discrimination, RTs increased to red S+ as well as red S-. Most aspects of the data were reproduced by simulations with a quantitative model that incorporated reinforcement-based decision and arousal processes.     

Rats exhibit asymmetrical retention functions for hedonic samples

Rats were trained in a symbolic delayed matching-to-sample task to discriminate hedonic sample stimuli that consisted of food or no food. Retention functions decreased more rapidly on trials initiated by a food sample than on trials initiated by a no-food sample when retention intervals were manipulated within session. The asymmetrical functions could not be explained in terms of mediation of choice responding by magazine head-entry behavior during the retention interval. Unlike within-session changes in retention interval, between-session changes did not result in steeper forgetting functions for food samples. These results in rats are consistent with previous findings reported for the presence and absence of visual samples in pigeons (Wixted, 1993).