Reinforcement-related neurons in the primate basal forebrain respond to the learned significance of task events rather than to the hedonic attributes of reward

The objective of this study was to determine if the responses of basal forebrain neurons are related to the cognitive processes necessary for the performance of behavioural tasks, or to the hedonic attributes of the reinforcers delivered to the monkey as

Increasing top-down suppression from prefrontal cortex facilitates tactile working memory

Navigated transcranial magnetic stimulation (TMS) combined with diffusion-weighted magnetic resonance imaging (DW-MRI) and tractography allows investigating functional anatomy of the human brain with high precision. Here we demonstrate that working memory (WM) processing of tactile temporal information is facilitated by delivering a single TMS pulse to the middle frontal gyrus (MFG) during memory maintenance. Facilitation was obtained only with a TMS pulse applied to a location of the MFG with anatomical connectivity to the primary somatosensory cortex (S1). TMS improved tactile WM also when distractive tactile stimuli interfered with memory maintenance. Moreover, TMS to the same MFG site attenuated somatosensory evoked responses (SEPs). The results suggest that the TMS-induced memory improvement is explained by increased top-down suppression of interfering sensory processing in S1 via the MFG-S1 link. These results demonstrate an anatomical and functional network that is involved in maintenance of tactile temporal WM. (C) 2009 Elsevier Inc. All rights reserved.

左侧梭状回内文字的字形加工：中文正常阅读者，中文文盲、非中文母语被试的对比研究

Reading is an important human-specific skill obtained through extensive learning experience and is reliance on the ability to rapidly recognize single words. According to the behavioral studies, the most important stage of reading is the representation of “visual word form”, which is independent on surface visual features of the reading materials. The prelexical visual word form representation is characterized by the abstractive and highly effective and precise processing. Neuroimaging and neuropsychological studies have investigated the neural basis underlying the visual word form processing. On the basis of summary of the existing literature, the current thesis aimed to address three fundamental questions involving neural basis of word recognition. First, is there a dedicated neural network that is specialized for word recognition? Second, is the orthographic information represented in the putative word/character selective region (VWFA)? Third, what is the role of reading experience in the genesis of the VWFA, is experience a main driver to shape VWFA instead of evolutionary selectivity? Nineteen Chinese literate volunteers, 5 Chinese illiterates and 4 native English speakers participated in this study, and performed perceptual tasks during fMRI scanning. To address the first question, we compared the differential responses to three categories of visual objects, i.e., faces, line drawings of objects and Chinese characters, and defined the region of interesting (ROI) for the next experiment. To address the second question, Chinese character orthography was manipulated to reveal possible differential responses to real characters, false characters, radical combinations, and stroke combinations in the regions defined by the first experiment. To examine the role of reading experience in genesis of specialization for character, the responses for unfamiliar Chinese characters in Chinese illiterates and native English speakers were compared with that in the Chinese literates, and tracked the change in cortical activation after a short-term reading training in the illiterates. Data were analyzed in two dimensions. Both BOLD signal amplitude and spatial distribution pattern among multi-voxels were used to systematically investigate the responsiveness of the left fusiform gyrus to Chinese characters. Our results provide strong and clear evidence for the existence of functionally specialized regions in the human ventral occipital-temporal cortex. In the skilled readers a region specialized for written words could be consistently found in the lateral part of the left fusiform gyrus, line drawings in the median part and faces in the middle. Our results further show that spatial distribution analysis, a method that was not commonly used in neuroimaging of reading, appears to be a more effective measurement for category specialization for visual objects processing. Although we failed to provide evidence that VWFA processes orthographic information in terms of signal intensitiy, we do show that response pattern of real characters and radical collections in this area is different from that of false characters and random stroke combinations. Our last set of experiments suggests that the selective bias to reading material is clearly experience dependent. The response to unknown characters in both English speakers/readers and Chinese illiterates is fundamentally different from that of the skilled Chinese readers. The response pattern for unknown characters is more similar to that for line drawings rather as a weak version of character in skilled Chinese readers. Short-term training is not sufficient to produce VWFA bias even when tested with learned characters, rather the learned characters generated a overall upward shift of the activation of the left fusiform region. Formation of a dedicated region specialized for visual word/character might depend on long-term extensive reading experience, or there might be a critical period for reading acquisition.

左中部梭状回枕颞沟内皮质对汉字加工的必要性、选择性和作用性质

A number of functional neuroimaging studies with skilled readers consistently showed activation to visual words in the left mid-fusiform cortex in occipitotemporal sulcus (LMFC-OTS). Neuropsychological studies also showed that lesions at left ventral occipitotemporal areas result in impairment in visual word processing. Based on these empirical observations and some theoretical speculations, a few researchers postulated that the LMFC-OTS is responsible for instant parallel and holistic extraction of the abstract representation of letter strings, and labeled this piece of cortex as “visual word form area” (VWFA). Nonetheless, functional neuroimaging studies alone is basically a correlative rather than causal approach, and lesions in the previous studies were typically not constrained within LMFC-OTS but also involving other brain regions beyond this area. Given these limitations, it remains unanswered for three fundamental questions: is LMFC-OTS necessary for visual word processing? is this functionally selective for visual word processing while unnecessary for processing of non-visual word stimuli? what are its function properties in visual word processing? This thesis aimed to address these questions through a series of neuropsychological, anatomical and functional MRI experiments in four patients with different degrees of impairments in the left fusiform gyrus. Necessity: Detailed analysis of anatomical brain images revealed that the four patients had differential foci of brain infarction. Specifically, the LMFC-OTS was damaged in one patient, while it remained intact in the other three. Neuropsychological experiments showed that the patient with lesions in the LMFC-OTS had severe impairments in reading aloud and recognizing Chinese characters, i.e., pure alexia. The patient with intact LMFC-OTS but information from the left visual field (LVF) was blocked due to lesions in the splenium of corpus callosum, showed impairment in Chinese characters recognition when the stimuli were presented in the LVF but not in the RVF, i.e. left hemialexia. In contrast, the other two patients with intact LMFC-OTS had normal function in processing Chinese characters. The fMRI experiments demonstrated that there was no significant activation to Chinese characters in the LMFC-OTS of the pure alexic patient and of the patient with left hemialexia when the stimuli were presented in the LVF. On the other hand, this patient, when Chinese characters were presented in right visual field, and the other two with intact LMFC-OTS had activation in the LMFC-OTS. These results together point to the necessity of the LMFC-OTS for Chinese character processing. Selectivity: We tested selectivity of the LMFC-OTS for visual word processing through systematically examining the patients’ ability for processing visual vs. auditory words, and word vs. non-word visual stimuli, such as faces, objects and colors. Results showed that the pure alexic patients could normally process auditory words (expression, understanding and repetition of orally presented words) and non-word visual stimuli (faces, objects, colors and numbers). Although the patient showed some impairments in naming faces, objects and colors, his performance scores were only slightly lower or not significantly different relative to those of the patients with intact LMFC-OTS. These data provide compelling evidence that the LMFC-OTS is not requisite for processing non-visual word stimuli, thus has selectivity for visual word processing. Functional properties: With tasks involving multiple levels and aspects of word processing, including Chinese character reading, phonological judgment, semantic judgment, identity judgment of abstract visual word representation, lexical decision, perceptual judgment of visual word appearance, and dictation, copying, voluntary writing, etc., we attempted to reveal the most critical dysfunction caused by damage in the LMFC-OTS, thus to clarify the most essential function of this region. Results showed that in addition to dysfunctions in Chinese character reading, phonological and semantic judgment, the patient with lesions at LMFC-OTS failed to judge correctly whether two characters (including compound and simple characters) with different surface features (e.g., different fonts, printed vs. handwritten vs. calligraphy styles, simplified characters vs. traditional characters, different orientations of strokes or whole characters) had the same abstract representation. The patient initially showed severe impairments in processing both simple characters and compound characters. He could only copy a compound character in a stroke-by-stroke manner, but not by character-by-character or even by radical-by-radical manners. During the recovery process, namely five months later, the patient could complete the abstract representation tasks of simple characters, but showed no improvement for compound characters. However, he then could copy compound characters in a radical-by-radical manner. Furthermore, it seems that the recovery of copying paralleled to that of judgment of abstract representation. These observations indicate that lesions of the LMFC-OTS in the pure alexic patients caused several damage in the ability of extracting the abstract representation from lower level units to higher level units, and the patient had especial difficulty to extract the abstract representation of whole character from its secondary units (e.g., radicals or single characters) and this ability was resistant to recover from impairment. Therefore, the LMFC-OTS appears to be responsible for the multilevel (particularly higher levels) abstract representations of visual word form. Successful extraction seems independent on access to phonological and semantic information, given the alexic patient showed severe impairments in reading aloud and semantic processing on simple characters while maintenance of intact judgment on their abstract representation. However, it is also possible that the interaction between the abstract representation and its related information e.g. phonological and semantic information was damaged as well in this patient. Taken together, we conclude that: 1) the LMFC-OTS is necessary for Chinese character processing, 2) it is selective for Chinese character processing, and 3) its critical function is to extract multiple levels of abstract representation of visual word and possibly to transmit it to phonological and semantic systems.

听觉想象的大脑表征

As a species of internal representation, how is mental imagery organized in the brain? There are two issues related to this question: the time course and the nature of mental imagery. On the nature of mental imagery, today's imagery debate is influenced by two opposing theories: (1) Pylyshyn’s propositional theory and (2) Kosslyn’s depictive representation theory. Behavioural studies indicated that imagery encodes properties of the physical world, such as the spacial and size information of the visual world. Neuroimaging and neuropsychological data indicated that sensory cortex; especially the primary sensory cortex, is involved in imagery. In visual modality, neuroimaging data further indicated that during visual imagery, spatial information is mapped in the primary visual, providing strong evidences for depictive theory. In the auditory modality, behavioural studies also indicated that auditory imagery represents loudness and pitch of sound; this kind of neuroimaging evidence, however, is absent. The aim of the present study was to investigate the time course of auditory imagery processing, and to provide the neuroimaging evidence that imaginal auditory representations encode loudness and pitch information, using the ERP method and a cue-imagery (S1)-S2 paradigm. The results revealed that imagery effects started with an enhancement of the P2, probably indexing the top-down allocation of attention to the imagery task; and continued into a more positive-going late positive potentials (LPC), probably reflecting the formation of auditory imagery. The amplitude of this LPC was inversely related to the pitch of the imagined sound, but directly related to the loudness of the imagined sound, which were consistent with auditory perception related N1 component, providing evidences that auditory imagery encodes pitch and loudness information. When the S2 showed difference in pitch of loudness from the previously imagined S1, the behavioral performance were significantly worse and accordingly a conflict related N2 was elicited; and the high conflict elicited greater N2 amplitude than low conflict condition, providing further evidences that imagery is analog of perception and can encode pitch and loudness information. The present study suggests that imagery starts with an mechanism of top-down allocation of attention to the imagery task; and continuing into the step of imagery formation during which the physical features of the imagined stimulus can be encoded, providing supports to Kosslyn’s depictive representation theory.