Role of left posterior superior temporal gyrus in phonological processing for speech perception and production

Models of both speech perception and speech production typically postulate a processing level that involves some form of phonological processing. There is disagreement, however, on the question of whether there are separate phonological systems for speech input versus speech output. We review a range of neuroscientific data that indicate that input and output phonological systems partially overlap. An important anatomical site of overlap appears to be the left posterior superior temporal gyrus. We then present the results of a new event-related functional magnetic resonance imaging (fMRI) experiment in which participants were asked to listen to and then (covertly) produce speech. In each participant, we found two regions in the left posterior superior temporal gyrus that responded both to the perception and production components of the task, suggesting that there is overlap in the neural systems that participate in phonological aspects of speech perception and speech production. The implications for neural models of verbal working memory are also discussed in connection with our findings.     

Towards a functional neuroanatomy of speech perception

The functional neuroanatomy of speech perception has been difficult to characterize. Part of the difficulty, we suggest, stems from the fact that the neural systems supporting 'speech perception' vary as a function of the task. Specifically, the set of cognitive and neural systems involved in performing traditional laboratory speech perception tasks, such as syllable discrimination or identification, only partially overlap those involved in speech perception as it occurs during natural language comprehension. In this review, we argue that cortical fields in the posterior-superior temporal lobe, bilaterally, constitute the primary substrate for constructing sound-based representations of speech, and that these sound-based representations interface with different supramodal systems in a task-dependent manner. Tasks that require access to the mental lexicon (i.e. accessing meaning-based representations) rely on auditory-to-meaning interface systems in the cortex in the vicinity of the left temporal-parietal-occipital junction. Tasks that require explicit access to speech segments rely on auditory-motor interface systems in the left frontal and parietal lobes. This auditory-motor interface system also appears to be recruited in phonological working memory.     

Left posterior auditory-related cortices participate both in speech perception and speech production: Neural overlap revealed by fMRI

Recent neuroimaging studies and neuropsychological data suggest that there are regions in posterior auditory cortex that participate both in speech perception and speech production. An outstanding question is whether the same neural regions support both perception and production or whether there exist discrete cortical fields subserving these functions. Previous neurophysiological studies suggest that there is indeed regional overlap between these systems, but those studies used a rehearsal task to assess production. The present study addressed this question in an event-related fMRI experiment in which subjects listened to speech and in separate trials, performed a covert object naming task. Single subject analysis revealed regions of coactivation for speech perception and production in the left posterior superior temporal sulcus (pSTS), left area Spt (a region in the Sylvian fissure at the parietal-temporal boundary), and left inferior frontal gyrus. These results are consistent with lesion data and previous physiological data indicating that posterior auditory cortex plays a role in both reception and expression of speech. We discuss these findings within the context of a neuroanatomical framework that proposes these neural sites are a part of an auditory-motor integration system.     

Distributed processing and cortical specialization for speech and environmental sounds in human temporal cortex

Using functional MRI, we investigated whether auditory processing of both speech and meaningful non-linguistic environmental sounds in superior and middle temporal cortex relies on a complex and spatially distributed neural system. We found that evidence for spatially distributed processing of speech and environmental sounds in a substantial extent of temporal cortices. Most importantly, regions previously reported as selective for speech over environmental sounds also contained distributed information. The results indicate that temporal cortices supporting complex auditory processing, including regions previously described as speech-selective, are in fact highly heterogeneous.     

Parametrically Dissociating Speech and Nonspeech Perception in the Brain Using fMRI

Candidate brain regions constituting a neural network for preattentive phonetic perception were identified with fMRI and multivariate multiple regression of imaging data. Stimuli contrasted along speech/nonspeech, acoustic, or phonetic complexity (three levels each) and natural/synthetic dimensions. Seven distributed brain regions' activity correlated with speech and speech complexity dimensions, including five left-sided foci [posterior superior temporal gyrus (STG), angular gyrus, ventral occipitotemporal cortex, inferior/posterior supramarginal gyrus, and middle frontal gyrus (MFG)] and two right-sided foci (posterior STG and anterior insula). Only the left MFG discriminated natural and synthetic speech. The data also supported a parallel rather than serial model of auditory speech and nonspeech perception.     

The effects of simulated stuttering and prolonged speech on the neural activation patterns of stuttering and nonstuttering adults

Functional magnetic resonance imaging was used to investigate the neural correlates of passive listening, habitual speech and two modified speech patterns (simulated stuttering and prolonged speech) in stuttering and nonstuttering adults. Within-group comparisons revealed increased right hemisphere biased activation of speech-related regions during the simulated stuttered and prolonged speech tasks, relative to the habitual speech task, in the stuttering group. No significant activation differences were observed within the nonstuttering participants during these speech conditions. Between-group comparisons revealed less left superior temporal gyrus activation in stutterers during habitual speech and increased right inferior frontal gyrus activation during simulated stuttering relative to nonstutterers. Stutterers were also found to have increased activation in the left middle and superior temporal gyri and right insula, primary motor cortex and supplementary motor cortex during the passive listening condition relative to nonstutterers. The results provide further evidence for the presence of functional deficiencies underlying auditory processing, motor planning and execution in people who stutter, with these differences being affected by speech manner.     

Modeling the categorical perception of speech sounds: A step toward biological plausibility

Our native language has a lifelong effect on how we perceive speech sounds. Behaviorally, this is manifested as categorical perception, but the neural mechanisms underlying this phenomenon are still unknown. Here, we constructed a computational model of categorical perception, following principles consistent with infant speech learning. A self-organizing network was exposed to a statistical distribution of speech input presented as neural activity patterns of the auditory periphery, resembling the way sound arrives to the human brain. In the resulting neural map, categorical perception emerges from most single neurons of the model being maximally activated by prototypical speech sounds, while the largest variability in activity is produced at category boundaries. Consequently, regions in the vicinity of prototypes become perceptually compressed, and regions at category boundaries become expanded. Thus, the present study offers a unifying framework for explaining the neural basis of the warping of perceptual space associated with categorical perception.     

Pure word deafness and the bilateral processing of the speech code

The analysis of pure word deafness (PWD) suggests that speech perception, construed as the integration of acoustic information to yield representations that enter into the linguistic computational system, (i) is separable in a modular sense from other aspects of auditory cognition and (ii) is mediated by the posterior superior temporal cortex in both hemispheres. PWD data are consistent with neuropsychological and neuroimaging evidence in a manner that suggests that the speech code is analyzed bilaterally. The typical lateralization associated with language processing is a property of the computational system that acts beyond the analysis of the input signal. The hypothesis of the bilateral mediation of the speech code does not imply that both sides execute the same computation. It is proposed that the speech signal is asymmetrically analyzed in the time domain, with left‐hemisphere mechanisms preferentially extracting information over shorter (25–50 ms) temporal integration windows and right mechanisms over longer (150–250 ms) windows.     

Behavioral and fMRI evidence that cognitive ability modulates the effect of semantic context on speech intelligibility

Text cues facilitate the perception of spoken sentences to which they are semantically related (Zekveld, Rudner, et al., 2011). In this study, semantically related and unrelated cues preceding sentences evoked more activation in middle temporal gyrus (MTG) and inferior frontal gyrus (IFG) than nonword cues, regardless of acoustic quality (speech in noise or speech in quiet). Larger verbal working memory (WM) capacity (reading span) was associated with greater intelligibility benefit obtained from related cues, with less speech-related activation in the left superior temporal gyrus and left anterior IFG, and with more activation in right medial frontal cortex for related versus unrelated cues. Better ability to comprehend masked text was associated with greater ability to disregard unrelated cues, and with more activation in left angular gyrus (AG). We conclude that individual differences in cognitive abilities are related to activation in a speech-sensitive network including left MTG, IFG and AG during cued speech perception.     

An fMRI investigation of covertly and overtly produced mono- and multisyllabic words

Studies suggest that the left insula may play an important role in speech motor programming. We used functional magnetic resonance imaging to investigate the role of the left insula in the production of monosyllabic or multisyllabic words during overt and covert speech conditions. The left insula did not show a BOLD response for multisyllabic words (which should require more speech motor programming) that was different from that for monosyllabic words. Left parietal lobe regions showed a greater response for multisyllabic as compared to monosyllabic words. This is consistent with clinical studies showing that left parietal lobe lesions can produce a deficit in speech programming. Despite similarities, covert and overt speech did not demonstrate the same patterns of BOLD response. The BOLD response was greater during overt speech in areas that have been shown to play an important role in speech production including left premotor cortex/BA6, left primary motor cortex, left insula, and left superior temporal gyrus.     

Do temporal processes underlie left hemisphere dominance in speech perception?

It is not unusual to find it stated as a fact that the left hemisphere is specialized for the processing of rapid, or temporal aspects of sound, and that the dominance of the left hemisphere in the perception of speech can be a consequence of this specialization. In this review we explore the history of this claim and assess the weight of this assumption. We will demonstrate that instead of a supposed sensitivity of the left temporal lobe for the acoustic properties of speech, it is the right temporal lobe which shows a marked preference for certain properties of sounds, for example longer durations, or variations in pitch. We finish by outlining some alternative factors that contribute to the left lateralization of speech perception.     

新生儿语音感知的神经基础:元分析

语言习得能力是人类在进化中获得的重要能力之一。语言认知的核心功能是语音加工, 因此, 语音加工的脑机制是认知心理学研究的重要课题。我们采用元分析方法, 对使用近红外技术考察新生儿语音感知的结构检测、偏差检测和母语感知的研究进行系统的定量分析, 探究新生儿的典型语音感知脑机制以及这三类语音加工相关脑区的异同。结果显示, 左侧额下回是新生儿检测语音结构的关键脑区; 双侧颞叶在新生儿语音偏差刺激检测中发挥关键作用; 新生儿的母语语音加工存在左侧化优势。     

Articulatory mediation of speech perception: A causal analysis of multi-modal imaging data

The inherent confound between the organization of articulation and the acoustic-phonetic structure of the speech signal makes it exceptionally difficult to evaluate the competing claims of motor and acoustic-phonetic accounts of how listeners recognize coarticulated speech. Here we use Granger causation analyzes of high spatiotemporal resolution neural activation data derived from the integration of magnetic resonance imaging, magnetoencephalography and electroencephalography, to examine the role of lexical and articulatory mediation in listeners’ ability to use phonetic context to compensate for place assimilation. Listeners heard two-word phrases such as pen pad and then saw two pictures, from which they had to select the one that depicted the phrase. Assimilation, lexical competitor environment and the phonological validity of assimilation context were all manipulated. Behavioral data showed an effect of context on the interpretation of assimilated segments. Analysis of 40 Hz gamma phase locking patterns identified a large distributed neural network including 16 distinct regions of interest (ROIs) spanning portions of both hemispheres in the first 200 ms of post-assimilation context. Granger analyzes of individual conditions showed differing patterns of causal interaction between ROIs during this interval, with hypothesized lexical and articulatory structures and pathways driving phonetic activation in the posterior superior temporal gyrus in assimilation conditions, but not in phonetically unambiguous conditions. These results lend strong support for the motor theory of speech perception, and clarify the role of lexical mediation in the phonetic processing of assimilated speech.     

言语想象的神经机制

言语想象不仅在大脑预处理机制方面起到重要的作用,还是目前脑机接口领域研究的热点。与正常言语产生过程相比,言语想象的理论模型、激活脑区、神经传导路径等均与其有较多相似之处。而言语障碍群体的言语想象、想象有意义的词语和句子时的脑神经机制与正常言语产生存在差异。鉴于人类言语系统的复杂性,言语想象的神经机制研究还面临一系列挑战,未来研究可在言语想象质量评价工具及神经解码范式、脑控制回路、激活通路、言语障碍群体的言语想象机制、词语和句子想象的脑神经信号等方面进一步探索,为有效提高脑机接口的识别率提供依据,为言语障碍群体的沟通提供便利。     

Sensorimotor foundations of speech perception in infancy

The perceptual system for speech is highly organized from early infancy. This organization bootstraps young human learners’ ability to acquire their native speech and language from speech input. Here, we review behavioral and neuroimaging evidence that perceptual systems beyond the auditory modality are also specialized for speech in infancy, and that motor and sensorimotor systems can influence speech perception even in infants too young to produce speech-like vocalizations. These investigations complement existing literature on infant vocal development and on the interplay between speech perception and production systems in adults. We conclude that a multimodal speech and language network is present before speech-like vocalizations emerge.     

Dorsal and ventral streams: a framework for understanding aspects of the functional anatomy of language

Despite intensive work on language-brain relations, and a fairly impressive accumulation of knowledge over the last several decades, there has been little progress in developing large-scale models of the functional anatomy of language that integrate neuropsychological, neuroimaging, and psycholinguistic data. Drawing on relatively recent developments in the cortical organization of vision, and on data from a variety of sources, we propose a new framework for understanding aspects of the functional anatomy of language which moves towards remedying this situation. The framework posits that early cortical stages of speech perception involve auditory fields in the superior temporal gyrus bilaterally (although asymmetrically). This cortical processing system then diverges into two broad processing streams, a ventral stream, which is involved in mapping sound onto meaning, and a dorsal stream, which is involved in mapping sound onto articulatory-based representations. The ventral stream projects ventro-laterally toward inferior posterior temporal cortex (posterior middle temporal gyrus) which serves as an interface between sound-based representations of speech in the superior temporal gyrus (again bilaterally) and widely distributed conceptual representations. The dorsal stream projects dorso-posteriorly involving a region in the posterior Sylvian fissure at the parietal-temporal boundary (area Spt), and ultimately projecting to frontal regions. This network provides a mechanism for the development and maintenance of "parity" between auditory and motor representations of speech. Although the proposed dorsal stream represents a very tight connection between processes involved in speech perception and speech production, it does not appear to be a critical component of the speech perception process under normal (ecologically natural) listening conditions, that is, when speech input is mapped onto a conceptual representation. We also propose some degree of bi-directionality in both the dorsal and ventral pathways. We discuss some recent empirical tests of this framework that utilize a range of methods. We also show how damage to different components of this framework can account for the major symptom clusters of the fluent aphasias, and discuss some recent evidence concerning how sentence-level processing might be integrated into the framework.     

Cerebral specialization for speech production in persons with Down syndrome.

The study of cerebral specialization in persons with Down syndrome (DS) has revealed an anomalous pattern of organization. Specifically, dichotic listening studies (e.g., Elliott & Weeks, 1993) have suggested a left ear/right hemisphere dominance for speech perception for persons with DS. In the current investigation, the cerebral dominance for speech production was examined using the mouth asymmetry technique. In right-handed, nonhandicapped subjects, mouth asymmetry methodology has shown that during speech, the right side of the mouth opens sooner and to a larger degree then the left side (Graves, Goodglass, & Landis, 1982). The phenomenon of right mouth asymmetry (RMA) is believed to reflect the direct access that the musculature on the right side of the face has to the left hemisphere's speech production systems. This direct access may facilitate the transfer of innervatory patterns to the muscles on the right side of the face. In the present study, the lateralization for speech production was investigated in 10 right-handed participants with DS and 10 nonhandicapped subjects. A RMA at the initiation and end of speech production occurred for subjects in both groups. Surprisingly, the degree of asymmetry between groups did not differ, suggesting that the lateralization of speech production is similar for persons with and persons without DS. These results support the biological dissociation model (Elliott, Weeks, & Elliott, 1987), which holds that persons with DS display a unique dissociation between speech perception (right hemisphere) and speech production (left hemisphere).     

动词论元结构复杂性加工的认知神经机制

动词论元结构复杂性表现在论元数量、论元范畴选择模式、题元角色指派模式和映射方式四个方面。大部分实证研究表明, 更多的论元数量、选择性论元范畴、选择性题元角色指派以及非典型映射, 使动词论元结构加工的认知神经机制更复杂。多论元加工功能脑区主要涉及左侧额下回和外侧裂周后部; 选择性论元范畴加工功能脑区主要涉及左侧额下回、额叶中后部、颞上回和颞叶中后部; 选择性题元角色指派加工功能脑区主要涉及外侧裂周后部、左侧额叶中后部和额下回; 非典型映射加工功能脑区主要涉及左侧额下回、颞上回、颞中回和颞叶后部。左侧额下回可能涉及初始句法加工、动词次范畴确定、句法移位和非宾格动词语义加工, 左侧额叶中后部可能涉及初始句法加工和动词次范畴确定, 左侧颞上回和颞叶中后部可能涉及表层句法加工和表层论元句法-语义整合, 外侧裂周后部可能涉及论元语义表征。动词论元结构加工过程和动词词汇特征表明, 复杂性某些方面存在交互作用。动词论元结构复杂性与加工难易的对应关系、复杂性加工难度层级和交互作用的认知神经机制以及汉语动词论元结构复杂性加工认知神经机制等议题, 有待进一步探讨。     

The effects of attention on speech perception: an fMRI study

Focusing of attention to a specific speech source plays an important role in everyday speech perception. However, little is known of the neuronal substrates of focused attention in speech perception. Thus, the present study investigated the effects on neuronal activation of directed attention to auditory stimuli that differed in semantic content. Using an event-related fMRI protocol, single vowels, three-phoneme pseudowords and three- and four-phoneme real nouns and words were randomly presented to the subjects during four different instructional conditions. One condition was passive listening without any specific instructions of focusing of attention. The other conditions were attention focused on either the vowels, the pseudowords or the words. Thus, the acoustic stimulation was constant across conditions. The subjects were 13 healthy adults. Functional MRI was performed with a 1.5 T scanner, using an event-related design. During passive listening, there were significant activations bilaterally in the superior temporal gyrus. Instruction to attend to the pseudowords caused activation in middle temporal lobe areas, extending more anterior compared to the activations seen during passive listening. Instruction to attend to the vowel sounds caused an increase in activation in the superior/medial temporal lobe, with a leftward asymmetry. Instruction to attend to the words caused a leftward asymmetry, particularly in the middle and superior temporal gyri. It is concluded that attention plays a modulatory role in neuronal activation to speech sounds, producing specific activations to specific stimulus categories that may act to facilitate speech perception.     

Neuroanatomically separable effects of imageability and grammatical class during single-word comprehension

The present study characterizes the neural correlates of noun and verb imageability and addresses the question of whether components of the neural network supporting word recognition can be separately modified by variations in grammatical class and imageability. We examined the effect of imageability on BOLD signal during single-word comprehension of nouns and verbs. Subjects made semantic similarity judgments while undergoing functional magnetic resonance imaging (fMRI). Nouns and verbs were matched on imageability, and imageability varied continuously within a grammatical category. We observed three anatomically separable effects: a main effect of grammatical class, a main effect of imageability, and an imageability by grammatical class cross-over interaction. The left superior parietal lobule and a region in the left fusiform responded similarly to increases in noun and verb imageability; the left superior temporal gyrus showed greater activity for verbs than nouns after imageability was matched across grammatical class; and, in both the left middle temporal gyrus and the left inferior frontal lobe, a decrease in noun but not verb imageability resulted in higher BOLD signal. The presence of reliable and anatomically separable main effects of both imageability and grammatical class renders unlikely the hypothesis that previously reported dissociations between nouns and verbs can be dismissed as imageability effects. However, some regions previously thought to respond to grammatical class or imageability instead respond to the interaction of these variables.