**9. Acknowledgements**

The authors are funded by grant PSI2010-19619 from Ministerio de Ciencia e Innovacion, Spain (MICINN).

### **10. References**

86 Neuroimaging – Cognitive and Clinical Neuroscience

depiction seems that as we move from more posterior/dorsal regions to more anterior/ventral in the left IFG, a gradient of activations can be found to be specifically involved (in this order) in phonology/articulation, syntax, and semantics. A similar gradient could be found in the STG when moving from the primary auditory association areas in or around Hesch's gyrus, spreading widely to both anterior and posterior regions in the STG, probably covering also parts of the inferior parietal cortex. In the latter case, the gradient seems to cover, following this order, phonology/articulation and syntax. If we want to expand these functions to semantic processes, then STS and at least several

From there, the system spreads to notably many other brain regions, comprising, posteriorly, large portions of the whole temporal lobes, including the temporal poles and part of the basal regions, as well as significant portions of the parietal cortex. Frontally, the system spreads to more anterior regions, including large extensions of the prefrontal cortex; among them, an area showing the most substantial increase in size in humans when compared to other primates and importantly involved in general intelligence. Significant medial regions, both in the prefrontal cortex and in the parietal cortex, are also included in this system. This *extended language network* (using an expression coined by Ferstl et al., 2008) largely overlaps with the *human default system*, a bilateral network in the human brain active when we are involved in "internal" mental tasks. If the linguistic message implies the visualization or representation of a given situation, then the corresponding primary or

The system can therefore be viewed as a continuous flux of information spreading from perisylvian areas toward multiple, distant areas. In turn, it also seems that the limits between linguistic and non-linguistic processes within this system appear blurred. An overall rule seems to be that the closer we move toward the sylvian fissure, the more specifically linguistic the process is. But even in this case (as we have seen), these regions are

Finally, that the flux of information spreads from perisylvian areas toward extensive regions of the cerebral cortex (actually, nearly all portions of the cortex appear susceptible of being involved) does not necessarily mean that this spreading strictly follows a temporal (sequential) order. Actually, brain networks continuously fire at different frequencies (e.g., Buzsáki, 2006), and it is plausible that information fluxes continuously in a reciprocal way and almost simultaneously between perysilvian and more distant areas. This would be a possible underlying mechanism explaining the large number of mutual influences from one structural layer of language (phonology, syntax, and semantics) to each other, as reported in the literature (e.g., Pulvermüller et al., 2009b). Indeed, considering that there are about 10.000 connections per neuron in the cerebral cortex, firing up to 1.000 times per second and therefore performing a comparable number of calculations (Previc, 2009), a parallel or at least cascade mode of operation of the whole (extended) language network emerges as a very plausible picture. On the other hand, the centrality of auditory/verbal (i.e., phonological/articulatory) information in human language would be consistent with the position of the two main hubs involved in language processing and the direction of the information flux spreading from them as primary receptors of language information to widespread areas, even if the overall

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**5** 

*Japan* 

Satoru Yokoyama *Tohoku University* 

**Neuro-Anatomical Overlap Between Language** 

In the nineteenth century, two studies in aphasiology comprise a turning point for research of brain-language relationships: Broca, 1861 and Wernicke, 1874. Based on these two studies, it was claimed that Broca's area (i.e., the pars triangularis and pars opercularis of the left inferior frontal gyrus) and Wernicke's area (i.e., the posterior part of the left superior/middle temporal gyrus, but in some situations including a part of the inferior parietal lobule) were involved in language production and comprehension, respectively (Geschwind, 1970). Recently, due to the development of functional brain imaging techniques (e.g., PET and fMRI), normal brains have been measured to examine the neuro-cognitive architecture of language processing. In particular, both Broca's and Wernicke's areas have been shown to be responsible for several language functions, such as single word processing

However, these two important regions are also activated for working memory-related processes, at least, including executive functions and short term memory processes of linguistic information, and the processes of storage and access to long term memory of linguistic information. This memory system could be assumed essential for language comprehension. For example, in order to comprehend a word, we have to first identify a series of sounds or letters as a certain word and to access its semantic information from long term memory. For sentence comprehension, we have to tentatively memorize several words comprising the sentence to compute the syntactic and semantic structure of the sentence. For example, it is clear that if we do not tentatively memorize words comprising the sentence, we cannot comprehend the sentence, since we have to compute the syntactic/semantic information of the sentence by using these words. Hence, in order to understand a language expression, we need the involvement of both the short and long term memory systems. In previous studies, there were essentially two types of standpoints regarding the involvement of the memory system in language comprehension. The first is that of the "specialist", who assumes that the syntactic processing system of the language processing system exists in our brain and is independent from other congnitive functions. The second is that of the "generalist", who assumes that the syntactic processing system has neural substrates in

common with other cognitive functions, mainly the working memory system.

In this chapter, recent neuroimaging studies of the neuro-cognitive architecture of single word and sentence processing will be briefly reviewed and the relationships between language and memory in the human brain will be discussed in the context of functional

**1. Introduction** 

and sentence processing (Fig. 1).

neuroimaging evidence.

**and Memory Functions in the Human Brain** 

