Individual differences in skilled adult readers reveal dissociable patterns of neural activity associated with component processes of reading

We used fMRI to examine patterns of brain activity associated with component processes of visual word recognition and their relationships to individual differences in reading skill. We manipulated both the judgments adults made on written stimuli and the characteristics of the stimuli. Phonological processing led to activation in left inferior frontal and temporal regions whereas semantic processing was associated with bilateral middle frontal activation. Individual differences in reading subskills were reflected in differences in the degree to which cortical regions were engaged during reading. Variation in sight word reading efficiency was associated with degree of activation in visual cortex. Increased phonological decoding skill was associated with greater activation in left temporo-parietal cortex. Greater reading comprehension ability was associated with decreased activation in anterior cingulate and temporal regions. Notably, associations between reading ability and neural activation indicate that brain/behavior relationships among skilled readers differ from patterns associated with dyslexia and reading development.     

Transfer of visual information after unilateral input to the brain

Visual callosal connections are more numerous and widespread in the association areas than in the primary visual cortex and adjoining visual areas. In keeping with this, the amount of "wrong" ipsilateral visual field that is represented in the various cortical areas of primates and cats increases as one goes from primary visual cortex to extraoccipital areas. Therefore it can be argued that transfer of unilaterally presented visual stimuli occurs mainly at the temporal and parietal cortical level.     

Precuneus-prefrontal activity during awareness of visual verbal stimuli.

Awareness is a personal experience, which is only accessible to the rest of world through interpretation. We set out to identify a neural correlate of visual awareness, using brief subliminal and supraliminal verbal stimuli while measuring cerebral blood flow distribution with H(2)(15)O PET. Awareness of visual verbal stimuli differentially activated medial parietal association cortex (precuneus), which is a polymodal sensory cortex, and dorsolateral prefrontal cortex, which is thought to be primarily executive. Our results suggest participation of these higher order perceptual and executive cortical structures in visual verbal awareness.     

On the neural correlates of object recognition awareness: relationship to computational activities and activities mediating perceptual awareness

Based on theoretical considerations of Aurell (1979) and Block (1995), we argue that object recognition awareness is distinct from purely sensory awareness and that the former is mediated by neuronal activities in areas that are separate and distinct from cortical sensory areas. We propose that two of the principal functions of neuronal activities in sensory cortex, which are to provide sensory awareness and to effect the computations that are necessary for object recognition, are dissociated. We provide examples of how this dissociation might be achieved and argue that the components of the neuronal activities which carry the computations do not directly enter the awareness of the subject. The results of these computations are sparse representations (i.e., vector or distributed codes) which are activated by the presentation of particular sensory objects and are essentially engrams for the recognition of objects. These final representations occur in the highest order areas of sensory cortex; in the visual analyzer, the areas include the anterior part of the inferior temporal cortex and the perirhinal cortex. We propose, based on lesion and connectional data, that the two areas in which activities provide recognition awareness are the temporopolar cortex and the medial orbitofrontal cortex. Activities in the temporopolar cortex provide the recognition awareness of objects learned in the remote past (consolidated object recognition), and those in the medial orbitofrontal cortex provide the recognition awareness of objects learned in the recent past. The activation of the sparse representation for a particular sensory object in turn activates neurons in one or both of these regions of cortex, and it is the activities of these neurons that provide the awareness of recognition of the object in question. The neural circuitry involved in the activation of these representations is discussed.     

Familiarization, attention, and recognition memory in infancy: an event-related potential and cortical source localization study

This study investigated the effects of familiarization and attention on event-related potential (ERP) correlates of recognition memory in infants. Infants 4.5, 6, or 7.5 months of age were either familiarized with 2 stimuli that were used during later testing or presented 2 stimuli that were not used later. Then, infants were presented with a recording of Sesame Street to elicit attention or inattention and presented with familiar and novel stimuli. A negative ERP component over the frontal and central electrodes (Nc) was larger in the preexposure familiarization group for novel- than for familiar-stimulus presentations, whereas the Nc did not differ for the group not receiving a familiarization exposure. Spatial independent components analysis of the electroencephelogram and "equivalent current dipole" analysis were used to examine putative cortical sources of the ERP components. The cortical source of Nc was located in areas of prefrontal cortex and anterior cingulate cortex.     

Executive control of gaze by the frontal lobes

Executive control requires controlling the initiation of movements, judging the consequences of actions, and adjusting performance accordingly. We have investigated the role of different areas in the frontal lobe in executive control expressed by macaque monkeys performing a saccade stop signal task. Certain neurons in the frontal eye field respond to visual stimuli, and others control the production of saccadic eye movements. Neurons in the supplementary eye field do not control directly the initiation of saccades but, instead, signal the production of errors, the anticipation and delivery of reinforcement, and the presence of response conflict. Neurons in the anterior cingulate cortex signal the production of errors and the anticipation and delivery of reinforcement, but not the presence of response conflict. Intracranial local field potentials in the anterior cingulate cortex of monkeys indicate that these medial frontal signals can contribute to event-related potentials related to performance monitoring. Electrical stimulation of the supplementary eye field improves performance in the task by elevating saccade latency. An interactive race model shows how interacting units produce behavior that can be described as the outcome of a race between independent processes and how conflict between gaze-holding and gaze-shifting neurons can be used to adjust performance.     

Examining the role of feedback in TMS-induced visual suppression: A cautionary tale

Visual suppression by single-pulse transcranial magnetic stimulation (sTMS) has been attributed to interruptions of either feedforward or feedback activity in the visual stream. The relative timing of the C1 event related potential (ERP) and of the TMS suppression, taken from separate studies, supports an interruption of feedback. Here we probe the validity of such cross-study comparisons, both by conducting a literature survey and by measuring each time window within participants for the same stimuli. Cortical transmission time was estimated using the C1. We then suppressed the same stimuli that elicited the C1 using sTMS of variable post-stimulus lags. Results do not conclusively discriminate between interruption of feedback or feedforward mechanisms as the source of the visual suppression. We suggest that more evidence is needed to distinguish between feedback and feedforward interference in TMS suppression effects and we advise caution in making inferences derived from separate literatures, using different stimuli.     

Multimodal action representation in human left ventral premotor cortex

We used functional magnetic resonance imaging (fMRI) to investigate the neural systems responding to the sight and to the sound of an action. Subjects saw a video of paper tearing in silence (V), heard the sound of paper tearing (A), and saw and heard the action simultaneously (A + V). Compared to a non-action control stimulus, we found that hearing action sounds (A) activated the anterior inferior frontal gyrus and middle frontal gyrus in addition to primary auditory cortex. The anterior inferior frontal gyrus, which is known to be activated by environmental sounds, also seems to be involved in recognizing actions by sound. Consistent with previous research, seeing an action video (V) compared with seeing a non-action video activated the premotor cortex, intraparietal cortex, and the pars opercularis of the inferior frontal gyrus. An A + V facilitation effect was found in the ventral premotor cortex on the border of areas 44, 6, 3a, and 3b for the action stimuli but not for the control stimuli. This region may be involved in integrating multimodal information about actions. These data provide evidence that the ventral premotor cortex may provide an action representation that abstracts across both agency (self and other) and sensory modality (hearing and seeing). This function may be an important precursor of language functions.     

Cortical integration of audio–visual speech and non-speech stimuli

Using fMRI we investigated the neural basis of audio–visual processing of speech and non-speech stimuli using physically similar auditory stimuli (speech and sinusoidal tones) and visual stimuli (animated circles and ellipses). Relative to uni-modal stimuli, the different multi-modal stimuli showed increased activation in largely non-overlapping areas. Ellipse-Speech, which most resembles naturalistic audio–visual speech, showed higher activation in the right inferior frontal gyrus, fusiform gyri, left posterior superior temporal sulcus, and lateral occipital cortex. Circle-Tone, an arbitrary audio–visual pairing with no speech association, activated middle temporal gyri and lateral occipital cortex. Circle-Speech showed activation in lateral occipital cortex, and Ellipse-Tone did not show increased activation relative to uni-modal stimuli. Further analysis revealed that middle temporal regions, although identified as multi-modal only in the Circle-Tone condition, were more strongly active to Ellipse-Speech or Circle-Speech, but regions that were identified as multi-modal for Ellipse-Speech were always strongest for Ellipse-Speech. Our results suggest that combinations of auditory and visual stimuli may together be processed by different cortical networks, depending on the extent to which multi-modal speech or non-speech percepts are evoked.     

大脑视知觉调控的神经机制

大脑的知觉加工并非单纯由外部刺激驱动,而是存在自上而下的知觉调控。尽管这一现象被大量实验研究证实,但其神经机制仍然是认知神经科学研究的重要问题。本研究系统介绍了知觉调控的神经基础、实现形式、研究范式,及其理论模型,分析指出了当前研究面临的主要问题,并对未来的研究进行了展望,以期促进该问题研究的进一步开展。     

The role of metabotropic glutamate receptors and cortical adaptation in habituation of odor-guided behavior

Decreases in behavioral investigation of novel stimuli over time may be mediated by a variety of factors including changes in attention, internal state, and motivation. Sensory cortical adaptation, a decrease in sensory cortical responsiveness over prolonged stimulation, may also play a role. In olfaction, metabotropic glutamate receptors on cortical afferent pre-synaptic terminals have been shown to underlie both cortical sensory adaptation and habituation of odor-evoked reflexes. The present experiment examined whether blockade of sensory cortical adaptation through bilateral infusion of the group III metabotropic glutamate receptor antagonist cyclopropyl-4-phosphonophenylglycine (CPPG) into the anterior piriform cortex could reduce habituation of a more complex odor-driven behavior such as investigation of a scented object or a conspecific. The results demonstrate that time spent investigating a scented jar, or a conspecific, decreases over the course of a continuous 10 minute trial. Acute infusion of CPPG bilaterally into the anterior piriform cortex significantly enhanced the time spent investigating the scented jar compared to investigation time in control rats, without affecting overall behavioral activity levels. Infusions into the brain outside of the piriform cortex were without effect. CPPG infusion into the piriform cortex also produced an enhancement of time spent investigating a conspecific, although this effect was not significant.     

Multisensory cortical processing of object shape and its relation to mental imagery

Here, we used functional magnetic resonance imaging to investigate the multisensory processing of object shape in the human cerebral cortex and explored the role of mental imagery in such processing. Regions active bilaterally during both visual and haptic shape perception, relative to texture perception in the respective modality, included parts of the superior parietal gyrus, the anterior intraparietal sulcus, and the lateral occipital complex. Of these bimodal regions, the lateral occipital complexes preferred visual over haptic stimuli, whereas the parietal areas preferred haptic over visual stimuli. Whereas most subjects reported little haptic imagery during visual shape perception, experiences of visual imagery during haptic shape perception were common. Across subjects, ratings of the vividness of visual imagery strongly predicted the amount of haptic shape-selective activity in the right, but not in the left, lateral occipital complex. Thus, visual imagery appears to contribute to activation of some, but not all, visual cortical areas during haptic perception.     

Long-distance feedback projections to area V1: Implications for multisensory integration,spatial awareness,and visual consciousness

It is generally agreed that information flow through the cortex is constrained by a hierarchical architecture. Recent experimental evidence suggests that projections descending the hierarchy and targeting the primary visual cortex (area V1) may play an essential role in perceptual processes. We have, therefore, reexamined feedback projections to area V1, using retrograde tracer injections in this area. In addition to well-known areas, quantification of labeling in higher cortical areas reveals a number of hitherto unknown long-distance feedback connections originating from auditory (A1), multisensory (STP) cortices, but also from a perirhinal area (36). These feedback projections from advanced cortical stations, a global feature shared by areas that belong to the ventral visual stream, could play an important role in early multisensory integration and spatial awareness and could provide the physical substrate for the involvement of area V1 in visual consciousness.     

经颅直流电刺激技术在增强健康个体认知功能中的应用及其影响因素

经颅直流电刺激作为一种无创脑刺激技术,已在临床治疗及康复领域有广泛应用。随着研究的深入和人类对于自身认知需求的提高,近年来也有研究者开始尝试使用该技术增强健康个体的认知功能。本文从感知觉、注意、记忆、学习和复杂任务五个方面对目前经颅直流电刺激技术在健康个体认知增强领域的研究现状进行梳理和总结,讨论了机制和影响因素,以及未来面临的问题和挑战。     

Coordination of voluntary and stimulus-driven attentional control in human cortex

Visual attention may be voluntarily directed to particular locations or features (voluntary control), or it may be captured by salient stimuli, such as the abrupt appearance of a new perceptual object (stimulus-driven control). Most often, however, the deployment of attention is the result of a dynamic interplay between voluntary attentional control settings (e.g., based on prior knowledge about a target's location or color) and the degree to which stimuli in the visual scene match these voluntary control settings. Consequently, nontarget items in the scene that share a defining feature with the target of visual search can capture attention, a phenomenon termed contingent attentional capture. We used functional magnetic resonance imaging to show that attentional capture by target-colored distractors is accompanied by increased cortical activity in corresponding regions of retinotopically organized visual cortex. Concurrent activation in the temporoparietal junction and ventral frontal cortex suggests that these regions coordinate voluntary and stimulus-driven attentional control settings to determine which stimuli effectively compete for attention.     

Vicarious responses to pain in anterior cingulate cortex: Is empathy a multisensory issue?

Results obtained with functional magnetic resonance imaging show that both feeling a moderately painful pinprick stimulus to the fingertips and witnessing another person’s hand undergo similar stimulation are associated with common activity in a pain-related area in the right dorsal anterior cingulate cortex (ACC). Common activity in response to noxious tactile and visual stimulation was restricted to the right inferior Brodmann’s area 24b. These results suggest a shared neural substrate for felt and seen pain for aversive ecological events happening to strangers and in the absence of overt symbolic cues. In contrast to ACC 24b, the primary somatosensory cortex showed significant activations in response to both noxious and innocuous tactile, but not visual, stimuli. The different response patterns in the two areas are consistent with the ACC’s role in coding the motivational-affective dimension of pain, which is associated with the preparation of behavioral responses to aversive events.     

Brain stimulation and conscious experience

Libet discovered that a substantial duration (> 0.5-1.0 s) of direct electrical stimulation of the surface of the somatosensory cortex at threshold currents is required before human subjects can report that a conscious somatosensory experience had occurred. Using a reaction time method we confirm that a similarly long stimulation duration at threshold currents is required for activation of elementary visual experiences (phosphenes) in human subjects following stimulation of the surface of the striate cortex. However, the reaction times for the subject to respond to the cessation of the visual experience after the end of electrical stimulation could be as brief as 225-242 ms. We also carried out extensive studies in cats under a variety of anesthetic conditions using the same electrodes and parameters of stimulation employed in the human studies to study the patterns of neuronal activity beneath the stimulating surface electrode. Whereas sufficiently strong currents can activate neurons within milliseconds, stimulating currents close to threshold activate sustained neural activity only after at least 350-500 ms. When currents are close to threshold, some neurons are inhibited for several hundreds of millisecond before the balance between inhibition and excitation shifts towards excitation. These results suggest that the prolonged latencies, i.e., latencies beyond 200-250 ms, for the emergence of conscious experience following direct cortical stimulation result from a delay in the sustained activation of underlying cortical neurons at threshold currents rather than being due to any unusually long duration in central processing time. Intracellular records from cortical neurological cells during repetitive electrical stimulation of the surface of the feline striate cortex demonstrate that such stimulation induces a profound depolarizing shift in membrane potential that may persist after each stimulus train. Such a depolarization is evidence that extracellular K+ concentrations have increased during electrical stimulation. Such an increase in extracellular K+ progressively increases cortical excitability until the threshold for sustained activation of cortical neurons is reached and then exceeded. Consequently, the long latency for threshold activation of cortical neurons depends upon a dynamically increasing cortical facilatory process that begins hundreds of milliseconds before there is sustained activation of such neurons. In some cases, this facilatory process must overcome an initial stimulus-induced inhibition before neuronal firing commences.     

Abnormal retinotopic representations in human visual cortex revealed by fMRI

The representation of the visual field in early visual areas is retinotopic. The point-to-point relationship on the retina is therefore maintained on the convoluted cortical surface. Functional magnetic resonance imaging (fMRI) has been able to demonstrate the retinotopic representation of the visual field in occipital cortex of normal subjects. Furthermore, visual areas that are retinotopic can be identified on computationally flattened cortical maps on the basis of positions of the vertical and horizontal meridians. Here, we investigate abnormal retinotopic representations in human visual cortex with fMRI. We present three case studies in which patients with visual disorders are investigated. We have tested a subject who only possesses operating rod photoreceptors. We find in this case that the cortex undergoes a remapping whereby regions that would normally represent central field locations now map more peripheral positions in the visual field: In a human albino we also find abnormal visual cortical activity. Monocular stimulation of each hemifield resulted in activations in the hemisphere contralateral to the stimulated eye. This is consistent with abnormal decussation at the optic chiasm in albinism. Finally, we report a case where a lesion to white matter has resulted in a lack of measurable activity in occipital cortex. The activity was absent for a small region of the visual field, which was found to correspond to the subject's field defect. The cases selected have been chosen to demonstrate the power of fMRI in identifying abnormalities in the cortical representations of the visual field in patients with visual dysfunction. Furthermore, the experiments are able to show how the cortex is capable of modifying the visual field representation in response to abnormal input.     

Age-related deficits in generation and manipulation of mental images: II. The role of dorsolateral prefrontal cortex.

The authors investigated neural substrates of age-related declines in mental imagery. Healthy adult participants (ages 19 to 77) performed a series of visual-spatial mental imagery tasks that varied in apparent difficulty and involved stimuli of varying graphic complexity. The volumes of the dorsolateral frontal cortex (DLPFC) and posterior visual processing areas were estimated from magnetic resonance imaging scans. The volume of the DLPFC and the fusiform cortex, working-memory capacity, and performance on the tasks involving image generation and manipulation were significantly reduced with age. Further analyses suggested that age-related deficits in performance on mental imagery tasks may stem in part from age-related shrinkage of the prefrontal cortex and age-related declines in working memory but not from age-related slowing of sensorimotor reaction time. The volume of cortical regions associated with modality-specific visual information processing did not show a consistent relationship with specific mental imagery processes.