Does Perceptual Learning Suffer from Retrograde Interference?

In motor learning, training a task B can disrupt improvements of performance of a previously learned task A, indicating that learning needs consolidation. An influential study suggested that this is the case also for visual perceptual learning. Using the same paradigm, we failed to reproduce these results. Further experiments with bisection stimuli also showed no retrograde disruption from task B on task A. Hence, for the tasks tested here, perceptual learning does not suffer from retrograde interference.     

Acquisition versus Consolidation of Auditory Perceptual Learning Using Mixed-Training Regimens

Learning is considered to consist of two distinct phases–acquisition and consolidation. Acquisition can be disrupted when short periods of training on more than one task are interleaved, whereas consolidation can be disrupted when a second task is trained after the first has been initiated. Here we investigated the conditions governing the disruption to acquisition and consolidation during mixed-training regimens in which primary and secondary amplitude modulation tasks were either interleaved or presented consecutively. The secondary task differed from the primary task in either task-irrelevant (carrier frequency) or task-relevant (modulation rate) stimulus features while requiring the same perceptual judgment (amplitude modulation depth discrimination), or shared both irrelevant and relevant features but required a different judgment (amplitude modulation rate discrimination). Based on previous literature we predicted that acquisition would be disrupted by varying the task-relevant stimulus feature during training (stimulus interference), and that consolidation would be disrupted by varying the perceptual judgment required (task interference). We found that varying the task-relevant or -irrelevant stimulus features failed to disrupt acquisition but did disrupt consolidation, whereas mixing two tasks requiring a different perceptual judgment but sharing the same stimulus features disrupted both acquisition and consolidation. Thus, a distinction between acquisition and consolidation phases of perceptual learning cannot simply be attributed to (task-relevant) stimulus versus task interference. We propose instead that disruption occurs during acquisition when mixing two tasks requiring a perceptual judgment based on different cues, whereas consolidation is always disrupted regardless of whether different stimulus features or tasks are mixed. The current study not only provides a novel insight into the underlying mechanisms of perceptual learning, but also has practical implications for the optimal design and delivery of training programs that aim to remediate perceptual difficulties.     

Sound facilitates visual learning

Numerous studies show that practice can result in performance improvements on low-level visual perceptual tasks [1-5]. However, such learning is characteristically difficult and slow, requiring many days of training [6-8]. Here, we show that a multisensory audiovisual training procedure facilitates visual learning and results in significantly faster learning than unisensory visual training. We trained one group of subjects with an audiovisual motion-detection task and a second group with a visual motion-detection task, and compared performance on trials containing only visual signals across ten days of training. Whereas observers in both groups showed improvements of visual sensitivity with training, subjects trained with multisensory stimuli showed significantly more learning both within and across training sessions. These benefits of multisensory training are particularly surprising given that the learning of visual motion stimuli is generally thought to be mediated by low-level visual brain areas [6, 9, 10]. Although crossmodal interactions are ubiquitous in human perceptual processing [11-13], the contribution of crossmodal information to perceptual learning has not been studied previously. Our results show that multisensory interactions can be exploited to yield more efficient learning of sensory information and suggest that multisensory training programs would be most effective for the acquisition of new skills.     

Cross-Limb Interference during Motor Learning

It is well known that following skill learning, improvements in motor performance may transfer to the untrained contralateral limb. It is also well known that retention of a newly learned task A can be degraded when learning a competing task B that takes place directly after learning A. Here we investigate if this interference effect can also be observed in the limb contralateral to the trained one. Therefore, five different groups practiced a ballistic finger flexion task followed by an interfering visuomotor accuracy task with the same limb. Performance in the ballistic task was tested before the training, after the training and in an immediate retention test after the practice of the interference task for both the trained and the untrained hand. After training, subjects showed not only significant learning and interference effects for the trained limb but also for the contralateral untrained limb. Importantly, the interference effect in the untrained limb was dependent on the level of skill acquisition in the interfering motor task. These behavioural results of the untrained limb were accompanied by training specific changes in corticospinal excitability, which increased for the hemisphere ipsilateral to the trained hand following ballistic training and decreased during accuracy training of the ipsilateral hand. The results demonstrate that contralateral interference effects may occur, and that interference depends on the level of skill acquisition in the interfering motor task. This finding might be particularly relevant for rehabilitation.     

The human basal forebrain integrates the old and the new

Acquisition of new learning is challenged by the phenomenon of proactive interference (PI), which occurs when previous learning disrupts later learning. Whereas human neuroimaging studies have focused on the cortical contributions to interference resolution, animal studies demonstrate that efficient resolution of PI depends on cholinergic modulation from basal forebrain (BF). Whether the BF promotes PI resolution in humans is unknown. Here, we adapted a PI paradigm from animal studies for use in a functional MRI experiment. During PI resolution, neurologically intact subjects recruited a BF network that included afferent anterior and posterior cortical sites associated with efficient memory acquisition and perceptual processing. Despite normal performance, nonamnesic patients with alcoholism, which is known to disrupt BF function, did not activate a BF network but instead invoked anterior cortical sites traditionally associated with executive function. These results provide evidence for parallel neural systems, each with the potential to resolve interference in the face of competing information.     

Perceptual learning reduces interneuronal correlations in macaque visual cortex

Responses of neurons in early visual cortex change little with training and appear insufficient to account for perceptual learning. Behavioral performance, however, relies on population activity, and the accuracy of a population code is constrained by correlated noise among neurons. We tested whether training changes interneuronal correlations in the dorsal medial superior temporal area, which is involved in multisensory heading perception. Pairs of single units were recorded simultaneously in two groups of subjects: animals trained extensively in a heading discrimination task, and "naive" animals that performed a passive fixation task. Correlated noise was significantly weaker in trained versus naive animals, which might be expected to improve coding efficiency. However, we show that the observed uniform reduction in noise correlations leads to little change in population coding efficiency when all neurons are decoded. Thus, global changes in correlated noise among sensory neurons may be insufficient to account for perceptual learning.     

Stimulus Roving and Flankers Affect Perceptual Learning of Contrast Discrimination in Macaca mulatta

‘Stimulus roving’ refers to a paradigm in which the properties of the stimuli to be discriminated vary from trial to trial, rather than being kept constant throughout a block of trials. Rhesus monkeys have previously been shown to improve their contrast discrimination performance on a non-roving task, in which they had to report the contrast of a test stimulus relative to that of a fixed-contrast sample stimulus. Human psychophysics studies indicate that roving stimuli yield little or no perceptual learning. Here, we investigate how stimulus roving influences perceptual learning in macaque monkeys and how the addition of flankers alters performance under roving conditions. Animals were initially trained on a contrast discrimination task under non-roving conditions until their performance levels stabilized. The introduction of roving contrast conditions resulted in a pronounced drop in performance, which suggested that subjects initially failed to heed the sample contrast and performed the task using an internal memory reference. With training, significant improvements occurred, demonstrating that learning is possible under roving conditions. To investigate the notion of flanker-induced perceptual learning, flanker stimuli (30% fixed-contrast iso-oriented collinear gratings) were presented jointly with central (roving) stimuli. Presentation of flanker stimuli yielded substantial performance improvements in one subject, but deteriorations in the other. Finally, after the removal of flankers, performance levels returned to their pre-flanker state in both subjects, indicating that the flanker-induced changes were contingent upon the continued presentation of flankers.     

Frequent Video Game Players Resist Perceptual Interference

Playing certain types of video games for a long time can improve a wide range of mental processes, from visual acuity to cognitive control. Frequent gamers have also displayed generalized improvements in perceptual learning. In the Texture Discrimination Task (TDT), a widely used perceptual learning paradigm, participants report the orientation of a target embedded in a field of lines and demonstrate robust over-night improvement. However, changing the orientation of the background lines midway through TDT training interferes with overnight improvements in overall performance on TDT. Interestingly, prior research has suggested that this effect will not occur if a one-hour break is allowed in between the changes. These results have suggested that after training is over, it may take some time for learning to become stabilized and resilient against interference. Here, we tested whether frequent gamers have faster stabilization of perceptual learning compared to non-gamers and examined the effect of daily video game playing on interference of training of TDT with one background orientation on perceptual learning of TDT with a different background orientation. As a result, we found that non-gamers showed overnight performance improvement only on one background orientation, replicating previous results with the interference in TDT. In contrast, frequent gamers demonstrated overnight improvements in performance with both background orientations, suggesting that they are better able to overcome interference in perceptual learning. This resistance to interference suggests that video game playing not only enhances the amplitude and speed of perceptual learning but also leads to faster and/or more robust stabilization of perceptual learning.     

Interference in ballistic motor learning: specificity and role of sensory error signals

Humans are capable of learning numerous motor skills, but newly acquired skills may be abolished by subsequent learning. Here we ask what factors determine whether interference occurs in motor learning. We speculated that interference requires competing processes of synaptic plasticity in overlapping circuits and predicted specificity. To test this, subjects learned a ballistic motor task. Interference was observed following subsequent learning of an accuracy-tracking task, but only if the competing task involved the same muscles and movement direction. Interference was not observed from a non-learning task suggesting that interference requires competing learning. Subsequent learning of the competing task 4 h after initial learning did not cause interference suggesting disruption of early motor memory consolidation as one possible mechanism underlying interference. Repeated transcranial magnetic stimulation (rTMS) of corticospinal motor output at intensities below movement threshold did not cause interference, whereas suprathreshold rTMS evoking motor responses and (re)afferent activation did. Finally, the experiments revealed that suprathreshold repetitive electrical stimulation of the agonist (but not antagonist) peripheral nerve caused interference. The present study is, to our knowledge, the first to demonstrate that peripheral nerve stimulation may cause interference. The finding underscores the importance of sensory feedback as error signals in motor learning. We conclude that interference requires competing plasticity in overlapping circuits. Interference is remarkably specific for circuits involved in a specific movement and it may relate to sensory error signals.     

Effects of cold narcosis on memory acquisition,consolidation and retrieval in honeybees(Apis mellifera)

In learning and memory studies on honeybees(Apis mellifera),cold-induced narcosis has been widely used to temporarily immobilize honeybees.In this study,we investigated the effects of cold narcosis on the associative memories in honeybees by using the proboscis extension response(PER)paradigm.Severe impairments in memory acquisition was found when cold narcosis was performed 30 min,instead of 1 h before training.Locomotor activities were reduced when honeybees were tested 15 min,instead of30 min after cold narcosis.These results indicate that cold narcosis impairs locomotor activities,as well as memory acquisition in a time-dependent manner,but by comparison no such effects on memory retrieval have yet been observed.[0]     

Effect of daytime nap on consolidation of declarative memory in humans

We studied effects of a daytime nap (1 hour) with including only NREM sleep on performance of declarative memory task (60 semantically unrelated word pairs) and general functional state. During training, procedure of learning of 30 word pairs was presented once, and that of the other 30 pairs was repeated twice. Strength of the task acquisition was tested. Subjects participated in two experiments: basic and control one. After learning participants either took a nap (basic experiment) or kept awake looking movies (control experiment). In 4.5 hours after the training session all the subjects were retested. As compared to the subjects who stayed awake during the training-retesting interval, subjects who had a NREM nap demonstrated enhanced performance. Concerning the strength of task acquisition, sleep-dependent performance was observed only for the word pairs learned once. Naps did not affect the functional state assessed by the reaction time dynamics and psychological testing.     

Estimation of mental effort in learning visual search by measuring pupil response

Perceptual learning refers to the improvement of perceptual sensitivity and performance with training. In this study, we examined whether learning is accompanied by a release from mental effort on the task, leading to automatization of the learned task. For this purpose, we had subjects conduct a visual search for a target, defined by a combination of orientation and spatial frequency, while we monitored their pupil size. It is well known that pupil size reflects the strength of mental effort invested in a task. We found that pupil size increased rapidly as the learning proceeded in the early phase of training and decreased at the later phase to a level half of its maximum value. This result does not support the simple automatization hypothesis. Instead, it suggests that the mental effort and behavioral performance reflect different aspects of perceptual learning. Further, mental effort would be continued to be invested to maintain good performance at a later stage of training.     

Prolonged maturation of auditory perception and learning in gerbils

In humans, auditory perception reaches maturity over a broad age range, extending through adolescence. Despite this slow maturation, children are considered to be outstanding learners, suggesting that immature perceptual skills might actually be advantageous to improvement on an acoustic task as a result of training (perceptual learning). Previous non‐human studies have not employed an identical task when comparing perceptual performance of young and mature subjects, making it difficult to assess learning. Here, we used an identical procedure on juvenile and adult gerbils to examine the perception of amplitude modulation (AM), a stimulus feature that is an important component of most natural sounds. On average, Adult animals could detect smaller fluctuations in amplitude (i.e., smaller modulation depths) than Juveniles, indicating immature perceptual skills in Juveniles. However, the population variance was much greater for Juveniles, a few animals displaying adult‐like AM detection. To determine whether immature perceptual skills facilitated learning, we compared naïve performance on the AM detection task with the amount of improvement following additional training. The amount of improvement in Adults correlated with naïve performance: those with the poorest naïve performance improved the most. In contrast, the naïve performance of Juveniles did not predict the amount of learning. Those Juveniles with immature AM detection thresholds did not display greater learning than Adults. Furthermore, for several of the Juveniles with adult‐like thresholds, AM detection deteriorated with repeated testing. Thus, immature perceptual skills in young animals were not associated with greater learning. © 2010 Wiley Periodicals, Inc. Develop Neurobiol 70: 636–648, 2010     

Perceptual Biases in Relation to Paranormal and Conspiracy Beliefs

Previous studies have shown that one’s prior beliefs have a strong effect on perceptual decision-making and attentional processing. The present study extends these findings by investigating how individual differences in paranormal and conspiracy beliefs are related to perceptual and attentional biases. Two field studies were conducted in which visitors of a paranormal conducted a perceptual decision making task (i.e. the face / house categorization task; Experiment 1) or a visual attention task (i.e. the global / local processing task; Experiment 2). In the first experiment it was found that skeptics compared to believers more often incorrectly categorized ambiguous face stimuli as representing a house, indicating that disbelief rather than belief in the paranormal is driving the bias observed for the categorization of ambiguous stimuli. In the second experiment, it was found that skeptics showed a classical ‘global-to-local’ interference effect, whereas believers in conspiracy theories were characterized by a stronger ‘local-to-global interference effect’. The present study shows that individual differences in paranormal and conspiracy beliefs are associated with perceptual and attentional biases, thereby extending the growing body of work in this field indicating effects of cultural learning on basic perceptual processes.     

Neural aromatization accelerates the acquisition of spatial memory via an influence on the songbird hippocampus

Circulating estrogens affect the neural circuits that underlie learning and memory in several vertebrates via an influence on the hippocampus. In the songbird hippocampus local estrogen synthesis due to the abundant expression of aromatase may modulate hippocampal function including spatial memory performance. Here, we examined the effect of estradiol, testosterone, and dihydrotestosterone on the structure and function of the songbird hippocampus. Adult male zebra finches were castrated, implanted with one of these steroids or a blank implant, and trained on a spatial memory task. The rate of acquisition and overall performance on this task was recorded by direct observation. The size and density of cells in the hippocampus and its volume were measured. Estradiol-treated birds learned the task more rapidly than any other group. Although testosterone- and blank-implanted birds did learn the task, we found no evidence of learning in dihydrotestosterone-implanted subjects. Cells in the rostral hippocampus were larger in estradiol- and testosterone-treated birds relative to other groups. A corresponding decrease in the density of cells was apparent in estradiol-implanted subjects relative to all other groups. These data suggest that estradiol may accelerate the acquisition of a spatial memory task and increase the size of neurons in the rostral hippocampus. Since testosterone-mediated changes in acquisition and cell size were similar to those of estradiol, but not dihydrotestosterone, we conclude that neural aromatization of testosterone to estrogen is responsible for effects on the structure and function of the songbird hippocampus.     

(−)-Linalool, a naturally occurring monoterpene compound, impairs memory acquisition in the object recognition task, inhibitory avoidance test and habituation to a novel environment in rats

It is known that (−)-linalool is a competitive antagonist of NMDA receptors, which play a key role in the learning and memory processes; however, only a few studies have reported a possible interference of (−)-linalool in memory. The purpose of this study was to investigate the (−)-linalool effects on acquisition of short- and long-term memories through the objects recognition task, inhibitory avoidance test and habituation to a novel environment. Furthermore, the open field test was used to investigate the interference of (−)-linalool in motivation, locomotion and exploration by animals. Wistar male adult rats received an intraperitoneal injection (i.p.) of saline (NaCl 0.9%), tween 5% or (−)-linalool (50 or 100 mg/kg) before training in the tasks; MK-801 (0.1 mg/kg), a glutamate antagonist, was used as positive control. Short-term (STM) and long-term (LTM) memories were tested 1.5 and 24 h after training, respectively, in the inhibitory avoidance and recognition objects. The results suggested that (−)-linalool (as 50- and 100-mg/kg doses) impaired LTM acquisition, but not STM acquisition, in the object recognition task. In the inhibitory avoidance test, animals receiving linalool (both doses) showed impairment in acquisition of both memories measured. In the open field test, the animals that received (−)-linalool showed no significant difference in the crossings and latency to start the locomotion in any of the doses tested, although (−)-linalool 100 mg/kg reduced rearing behavior. When re-exposed to open field 24 h after training, the rats that received (−)-linalool 100 mg/kg showed no habituation. Taken together, these data suggested that (−)-linalool was able to impair the acquisition of memory in rats, which can be associated to (−)-linalool antagonist capacity as regards NMDA glutamatergic receptors, since other glutamate antagonists also seem to affect memory.     

Practicing coarse orientation discrimination improves orientation signals in macaque cortical area v4

Practice improves the performance in visual tasks, but mechanisms underlying this adult brain plasticity are unclear. Single-cell studies reported no [1], weak [2], or moderate [3, 4] perceptual learning-related changes in macaque visual areas V1 and V4, whereas none were found in middle temporal (MT) [5]. These conflicting results and modeling of human (e.g., [6, 7]) and monkey data [8] suggested that changes in the readout of visual cortical signals underlie perceptual learning, rather than changes in these signals. In the V4 learning studies, monkeys discriminated small differences in orientation, whereas in the MT study, the animals discriminated opponent motion directions. Analogous to the latter study, we trained monkeys to discriminate static orthogonal orientations masked by noise. V4 neurons showed robust increases in their capacity to discriminate the trained orientations during the course of the training. This effect was observed during discrimination and passive fixation but specifically for the trained orientations. The improvement in neural discrimination was due to decreased response variability and an increase of the difference between the mean responses for the two trained orientations. These findings demonstrate that perceptual learning in a coarse discrimination task indeed can change the response properties of a cortical sensory area.     

Expectation Suppression in Early Visual Cortex Depends on Task Set

Stimulus expectation can modulate neural responses in early sensory cortical regions, with expected stimuli often leading to a reduced neural response. However, it is unclear whether this expectation suppression is an automatic phenomenon or is instead dependent on the type of task a subject is engaged in. To investigate this, human subjects were presented with visual grating stimuli in the periphery that were either predictable or non-predictable while they performed three tasks that differently engaged cognitive resources. In two of the tasks, the predictable stimulus was task-irrelevant and spatial attention was engaged at fixation, with a high load on either perceptual or working memory resources. In the third task, the predictable stimulus was task-relevant, and therefore spatially attended. We observed that expectation suppression is dependent on the cognitive resources engaged by a subjects’ current task. When the grating was task-irrelevant, expectation suppression for predictable items was visible in retinotopically specific areas of early visual cortex (V1-V3) during the perceptual task, but it was abolished when working memory was loaded. When the grating was task-relevant and spatially attended, there was no significant effect of expectation in early visual cortex. These results suggest that expectation suppression is not an automatic phenomenon, but dependent on attentional state and type of available cognitive resources.     

Susceptibility to Declarative Memory Interference is Pronounced in Primary Insomnia

Sleep has been shown to stabilize memory traces and to protect against competing interference in both the procedural and declarative memory domain. Here, we focused on an interference learning paradigm by testing patients with primary insomnia (N = 27) and healthy control subjects (N = 21). In two separate experimental nights with full polysomnography it was revealed that after morning interference procedural memory performance (using a finger tapping task) was not impaired in insomnia patients while declarative memory (word pair association) was decreased following interference. More specifically, we demonstrate robust associations of central sleep spindles (in N3) with motor memory susceptibility to interference as well as (cortically more widespread) fast spindle associations with declarative memory susceptibility. In general the results suggest that insufficient sleep quality does not necessarily show up in worse overnight consolidation in insomnia but may only become evident (in the declarative memory domain) when interference is imposed.     

Memory for semantically related and unrelated declarative information: the benefit of sleep, the cost of wake

Numerous studies have examined sleep's influence on a range of hippocampus-dependent declarative memory tasks, from text learning to spatial navigation. In this study, we examined the impact of sleep, wake, and time-of-day influences on the processing of declarative information with strong semantic links (semantically related word pairs) and information requiring the formation of novel associations (unrelated word pairs). Participants encoded a set of related or unrelated word pairs at either 9 am or 9 pm, and were then tested after an interval of 30 min, 12 hr, or 24 hr. The time of day at which subjects were trained had no effect on training performance or initial memory of either word pair type. At 12 hr retest, memory overall was superior following a night of sleep compared to a day of wakefulness. However, this performance difference was a result of a pronounced deterioration in memory for unrelated word pairs across wake; there was no sleep-wake difference for related word pairs. At 24 hr retest, with all subjects having received both a full night of sleep and a full day of wakefulness, we found that memory was superior when sleep occurred shortly after learning rather than following a full day of wakefulness. Lastly, we present evidence that the rate of deterioration across wakefulness was significantly diminished when a night of sleep preceded the wake period compared to when no sleep preceded wake, suggesting that sleep served to stabilize the memories against the deleterious effects of subsequent wakefulness. Overall, our results demonstrate that 1) the impact of 12 hr of waking interference on memory retention is strongly determined by word-pair type, 2) sleep is most beneficial to memory 24 hr later if it occurs shortly after learning, and 3) sleep does in fact stabilize declarative memories, diminishing the negative impact of subsequent wakefulness.