Direct electrical stimulation mapping of cognitive functions in the human brain.

Direct electrical stimulation (DES) is a well-established clinical tool for mapping cognitive functions while patients are undergoing awake neurosurgery or invasive long-term monitoring to identify epileptogenic tissue. Despite the proliferation of a range of invasive and noninvasive methods for mapping sensory, motor and cognitive processes in the human brain, DES remains the clinical gold standard for establishing the margins of brain tissue that can be safely removed while avoiding long-term neurological deficits. In parallel, and principally over the last two decades, DES has emerged as a powerful scientific tool for testing hypotheses of brain organization and mechanistic hypotheses of cognitive function. DES can cause transient "lesions" and thus can support causal inferences about the necessity of stimulated brain regions for specific functions, as well as the separability of sensory, motor and cognitive processes. This Special Issue of Cognitive Neuropsychology emphasizes the use of DES as a research tool to advance understanding of normal brain organization and function.

The processing of inflected and derived words in writing.

We report on an English-speaking, aphasic individual (RB) with a spelling deficit more severely affecting orthographically irregular words for which phonologically plausible errors (PPEs) were produced. PPEs were observed for all word forms, with the exception of inflectional suffixes, despite the irregular sound-print mappings of many inflectional suffixes (e.g., walked → /wɔkt/). RB's pattern replicates that reported in Badecker, Rapp, and Caramazza (Badecker, W., Rapp, B., & Caramazza, A. (1996). Lexical Morphology and the Two Orthographic Routes. Cognitive Neuropsychology, 13, 161-176). We extended their investigation by examining RB's spelling of derived words and found a selective deficit for derived words compared to inflected words in writing. This selective deficit did not appear to reflect differences in morphological transparency or suffix frequencies that exist between inflection and derivation. This is the first evidence that distinct neural mechanisms support inflection and derivation in spelling.

How modality-specific is morphology?

Writing has long been considered to be dependent on speaking. However, modality-specific dissociations between written and spoken word production imply that word production is supported by distinct neural mechanisms in writing, which can be impaired or spared regardless of the intactness of spoken word production. Rapp et al. (2015). Modality and morphology: What we write may not be what we say. Psychological Science, 26, 892-902 documented a double dissociation where problems with regular inflections were selectively restricted to writing or speaking. We report on two English-speaking aphasic individuals who exhibit this same modality-specific dissociation of inflectional processing, replicating the original findings. We expand on Rapp et al.'s study by examining whether the dissociations observed with regular inflections extend to other morphological forms, such as derivation and irregular inflection. Results showed that the dissociation holds for derivation; however, both participants were impaired with irregular inflections in both output modalities. Implications of these findings for morphological processing and the independence of the orthographic system are discussed.

Topographical gradients of semantics and phonology revealed by temporal lobe stimulation.

Word retrieval is a fundamental component of oral communication, and it is well established that this function is supported by left temporal cortex. Nevertheless, the specific temporal areas mediating word retrieval and the particular linguistic processes these regions support have not been well delineated. Toward this end, we analyzed over 1000 naming errors induced by left temporal cortical stimulation in epilepsy surgery patients. Errors were primarily semantic (lemon → "pear"), phonological (horn → "corn"), non-responses, and delayed responses (correct responses after a delay), and each error type appeared predominantly in a specific region: semantic errors in mid-middle temporal gyrus (TG), phonological errors and delayed responses in middle and posterior superior TG, and non-responses in anterior inferior TG. To the extent that semantic errors, phonological errors and delayed responses reflect disruptions in different processes, our results imply topographical specialization of semantic and phonological processing. Specifically, results revealed an inferior-to-superior gradient, with more superior regions associated with phonological processing. Further, errors were increasingly semantically related to targets toward posterior temporal cortex. We speculate that detailed semantic input is needed to support phonological retrieval, and thus, the specificity of semantic input increases progressively toward posterior temporal regions implicated in phonological processing. Hum Brain Mapp 38:688-703, 2017. © 2016 Wiley Periodicals, Inc.

A selective morpho-phonological deficit?

We report on an English-speaking, aphasic individual (TB) who showed a striking dissociation in speaking with the different forms (allomorphs) that an inflection can take. Although very accurate in producing the consonantal inflections (-/s/, -/z/, -/d/, -/t/), TB consistently omitted syllabic inflections (-/əz/, -/əd/), therefore correctly saying "dogs" or "walked," but "bench" for benches or "skate" for skated. Results from control tests ruled out that TB's selective difficulties stemmed from problems in selecting the correct inflection for the syntactic context or problems related to phonological or articulatory mechanisms. TB's selective difficulties appeared instead to concern morpho-phonological mechanisms responsible for adapting morphological elements to word phonology. These mechanisms determine whether the plural inflection surfaces in the noun bench as voiced (-/z/), unvoiced (-/s/) or syllabic (-/əz/). Our results have implications for understanding how morphological elements are encoded in the lexicon and the nature of morpho-phonological mechanisms involved in speech production.

Functional differences among stimulation-identified cortical naming sites in the temporal region.

To preserve postoperative language, electrical stimulation mapping is often conducted prior to surgery involving the language-dominant hemisphere. Object naming is the task most widely used to identify language cortex, and sites where stimulation elicits naming difficulty are typically spared from resection. In clinical practice, sites classified as positive undergo no further testing regarding the underlying cause of naming failure. Word production is a complex function involving multiple mechanisms that culminate in the identification of the target word. Two main mechanisms, i.e., semantic and phonological, underlie the retrieval of stored information regarding word meaning and word sounds, and naming can be hampered by disrupting either of these. These two mechanisms are likely mediated by different brain areas, and therefore, stimulation-identified naming sites might not be functionally equivalent. We investigated whether further testing at stimulation-identified naming sites would reveal an anatomical dissociation between these two mechanisms. In 16 patients with refractory temporal lobe epilepsy (TLE) with implanted subdural electrodes, we tested whether, despite inability to produce an item name, patients could reliably access semantic or phonological information regarding objects during cortical stimulation. We found that stimulation at naming sites in superior temporal cortex tended to impair phonological processing yet spared access to semantic information. By contrast, stimulation of inferior temporal naming sites revealed a greater proportion of sites where semantic access was impaired and a dissociation between sites where stimulation spared or disrupted semantic or phonological processing. These functional-anatomical dissociations reveal the more specific contribution to naming provided by these cortical areas and shed light on the often profound, interictal word-finding deficit observed in temporal lobe epilepsy. Additionally, these techniques potentially lay the groundwork for future studies to determine whether particular naming sites that fall within the margins of the desired clinical resection might be resected without significant risk of decline.

Early Parallel Activation of Semantics and Phonology in Picture Naming: Evidence from a Multiple Linear Regression MEG Study.

The time course of brain activation during word production has become an area of increasingly intense investigation in cognitive neuroscience. The predominant view has been that semantic and phonological processes are activated sequentially, at about 150 and 200-400 ms after picture onset. Although evidence from prior studies has been interpreted as supporting this view, these studies were arguably not ideally suited to detect early brain activation of semantic and phonological processes. We here used a multiple linear regression approach to magnetoencephalography (MEG) analysis of picture naming in order to investigate early effects of variables specifically related to visual, semantic, and phonological processing. This was combined with distributed minimum-norm source estimation and region-of-interest analysis. Brain activation associated with visual image complexity appeared in occipital cortex at about 100 ms after picture presentation onset. At about 150 ms, semantic variables became physiologically manifest in left frontotemporal regions. In the same latency range, we found an effect of phonological variables in the left middle temporal gyrus. Our results demonstrate that multiple linear regression analysis is sensitive to early effects of multiple psycholinguistic variables in picture naming. Crucially, our results suggest that access to phonological information might begin in parallel with semantic processing around 150 ms after picture onset.

Why Saturday could be both green and red in synesthesia.

It has long been observed that certain words induce multiple synesthetic colors, a phenomenon that has remained largely unexplored. We report here on the distinct synesthetic colors two synesthetes experienced with closed sets of concepts (digits, weekdays, months). For example, Saturday was associated with green, like other word starting with s; however, Saturday also had its specific color (red). Auditory priming and Visual Color Stroop tasks were used to understand the cognitive mechanisms supporting the distinct synesthetic colors. Results revealed that processing of word segments and whole words was specifically involved in each type of synesthetic colors. However, these mechanisms differed between participants, as they could relate either to orthography (and written words) or phonology (and spoken words). Further differences concerned the word representations, which varied as to whether or not they encoded serial positions. In addition to clarifying the cognitive mechanisms underlying the distinct synesthetic colors, our results offer some clues for understanding the neurocognitive underpinnings of a rather common form of synesthesia.

Modality and morphology: what we write may not be what we say.

Written language is an evolutionarily recent human invention; consequently, its neural substrates cannot be determined by the genetic code. How, then, does the brain incorporate skills of this type? One possibility is that written language is dependent on evolutionarily older skills, such as spoken language; another is that dedicated substrates develop with expertise. If written language does depend on spoken language, then acquired deficits of spoken and written language should necessarily co-occur. Alternatively, if at least some substrates are dedicated to written language, such deficits may doubly dissociate. We report on 5 individuals with aphasia, documenting a double dissociation in which the production of affixes (e.g., the -ing in jumping) is disrupted in writing but not speaking or vice versa. The findings reveal that written- and spoken-language systems are considerably independent from the standpoint of morpho-orthographic operations. Understanding this independence of the orthographic system in adults has implications for the education and rehabilitation of people with written-language deficits.

Foreign and Regional Languages Make You Less Deontological.

Previous studies have shown that foreign languages can change people's responses to moral dilemmas, making them more likely to choose harm (e.g., to kill one individual in order to save a few lives). Regional languages have also been shown to make sacrificial choices more likely. Regional languages are typically acquired early and used routinely among family and acquaintances, thus differing from foreign languages that are typically acquired later and used rather sporadically. Using a process dissociation procedure, we show in the present study that regional and foreign languages weaken the contribution of the deontological view in similar ways. Furthermore, the effects of both languages were modulated by proficiency, so that less proficient bilinguals showed a stronger decrease of the deontological tendency. These similarities suggest that the effects induced by both languages stem from what these languages have in common. Both languages are not experienced in contexts critical in forging moral views (e.g., public institutions, media, schools). We propose that the effects of foreign and regional languages stem from the lack of experience in such contexts.

Independent distractor frequency and age-of-acquisition effects in picture-word interference: fMRI evidence for post-lexical and lexical accounts according to distractor type.

In two fMRI experiments, participants named pictures with superimposed distractors that were high or low in frequency or varied in terms of age of acquisition. Pictures superimposed with low-frequency words were named more slowly than those superimposed with high-frequency words, and late-acquired words interfered with picture naming to a greater extent than early-acquired words. The distractor frequency effect (Experiment 1) was associated with increased activity in left premotor and posterior superior temporal cortices, consistent with the operation of an articulatory response buffer and verbal self-monitoring system. Conversely, the distractor age-of-acquisition effect (Experiment 2) was associated with increased activity in the left middle and posterior middle temporal cortex, consistent with the operation of lexical level processes such as lemma and phonological word form retrieval. The spatially dissociated patterns of activity across the two experiments indicate that distractor effects in picture-word interference may occur at lexical or postlexical levels of processing in speech production.

How the hand has shaped sign languages.

In natural languages, biological constraints push toward cross-linguistic homogeneity while linguistic, cultural, and historical processes promote language diversification. Here, we investigated the effects of these opposing forces on the fingers and thumb configurations (handshapes) used in natural sign languages. We analyzed over 38,000 handshapes from 33 languages. In all languages, the handshape exhibited the same form of adaptation to biological constraints found in tasks for which the hand has naturally evolved (e.g., grasping). These results were not replicated in fingerspelling-another task where the handshape is used-thus revealing a signing-specific adaptation. We also showed that the handshape varies cross-linguistically under the effects of linguistic, cultural, and historical processes. Their effects could thus emerge even without departing from the demands of biological constraints. Handshape's cross-linguistic variability consists in changes in the frequencies with which the most faithful handshapes to biological constraints appear in individual sign languages.

Finding levels of abstraction in speech production: evidence from sound-production impairment.

A widely held view in linguistics and psycholinguistics is that there are distinct levels of processing for context-independent and context-specific representations of sound structure. Recently, this view has been disputed, in part because of the absence of clear evidence that there are abstract mental representations of discrete sound-structure units. Here, we present novel evidence that separate context-independent and context-specific representations of sound structure are supported by distinct brain mechanisms that can be selectively impaired in individuals with acquired brain deficits. Acoustic data from /s/-deletion errors of 2 aphasic speakers suggest both a phonological level of processing at which sound representations (e.g., /p/) do not specify context-specific detail (e.g., aspirated or unaspirated) and a distinct level at which context-specific information is represented. These data help constrain accounts of sound-structure processing in word production and crucially support the claim that context-independent linguistic information affects language production.

Pupillary Stroop effects.

We recorded the pupil diameters of participants performing the words' color-naming Stroop task (i.e., naming the color of a word that names a color). Non-color words were used as baseline to firmly establish the effects of semantic relatedness induced by color word distractors. We replicated the classic Stroop effects of color congruency and color incongruency with pupillary diameter recordings: relative to non-color words, pupil diameters increased for color distractors that differed from color responses, while they reduced for color distractors that were identical to color responses. Analyses of the time courses of pupil responses revealed further differences between color-congruent and color-incongruent distractors, with the latter inducing a steep increase of pupil size and the former a relatively lower increase. Consistent with previous findings that have demonstrated that pupil size increases as task demands rise, the present results indicate that pupillometry is a robust measure of Stroop interference, and it represents a valuable addition to the cognitive scientist's toolbox.

When do combinatorial mechanisms apply in the production of inflected words?

A central question for theories of inflected word processing is to determine under what circumstances compositional procedures apply. Some accounts (e.g., the dual-mechanism model; Clahsen, 1999 ) propose that compositional processes only apply to verbs that take productive affixes. For all other verbs, inflected forms are assumed to be stored in the lexicon in a nondecomposed manner. This account makes clear predictions about the consequences of disruption to the lexical access mechanisms involved in the spoken production of inflected forms. Briefly, it predicts that nonproductive forms (which require lexical access) should be more affected than productive forms (which, depending on the language task, may not). We tested these predictions through the detailed analysis of the spoken production of a German-speaking individual with an acquired lexical impairment resulting from a stroke. Analyses of response accuracy, error types, and frequency effects revealed that combinatorial processes are not restricted to verbs that take productive inflections. On this basis, we propose an alternative account, the stem-based assembly model (SAM), which posits that combinatorial processes may be available to all stems and not only to those that combine with productive affixes.

Language can shape the perception of oriented objects.

Seeing an object is a natural source for learning about the object's configuration. We show that language can also shape our knowledge about visual objects. We investigated sign language that enables deaf individuals to communicate through hand movements with as much expressive power as any other natural language. A few signs represent objects in a specific orientation. Sign-language users (signers) recognized visual objects faster when oriented as in the sign, and this match in orientation elicited specific brain responses in signers, as measured by event-related potentials (ERPs). Further analyses suggested that signers' responsiveness to object orientation derived from changes in the visual object representations induced by the signs. Our results also show that language facilitates discrimination between objects of the same kind (e.g., different cars), an effect never reported before with spoken languages. By focusing on sign language we could better characterize the impact of language (a uniquely human ability) on object visual processing.

Hands show where things are: The close similarity between sign and natural space.

Many of the signs produced across sign languages are iconic, in the sense that they resemble the concepts they represent. We examined whether location, one of basic sign parameters along with handshape and movement, is systematically used for purposes of iconicity. Our findings revealed a mapping of vertical sign space that is exploited in its entirety for encoding typical locations in natural space. In all of the twenty sign languages we analyzed, signs were more likely to have high locations with concepts typically occurring in high vs. low regions of natural space (e.g., cloud vs. root). Furthermore, the height of signs produced to identify a visual object varied depending on object position (e.g., it was higher for basketball in the basket than basketball on the floor). It thus appears that signing space is permeable to semantic and episodic features, and iconicity plays a crucial role in making signing space so adaptable.

Foreign language effect in decision-making: How foreign is it?

It has been shown that decisions and moral judgments differ when made using native languages compared to foreign languages. Cross-linguistic differences appeared in foreign languages that monolinguals typically acquired in school and used neither routinely nor extensively. We replicated these differences with two populations of proficient, native bilinguals (Italian-Venetian; Italian-Bergamasque). Venetian and Bergamasque are spoken in households and informal circles, unlike Italian, which is also used in more formal contexts. The findings reported in foreign languages for the Asian Disease Problem and the Footbridge Dilemma were reproduced in Venetian and Bergamasque. Our results show that language effects on decision-making and moral judgments are not restricted to foreign languages. The explanation proposed for foreign languages of cross-linguistic differences in emotion responses does not apply to our proficient, native bilinguals, who showed emotion responses of equal intensity in their languages. We propose that the contexts in which bilinguals use a language - either native, regional or foreign - could affect decisions.

Knowing where but not what: impaired thematic roles and spatial language.

We describe case J.P. who, following a left inferior frontal lesion, made frequent role confusions in comprehension and production (e.g., saying "The boy kicks the girl" for a picture showing a girl kicking a boy). J.P.'s preserved ability to judge the grammaticality of sentences rules out a syntactic deficit as the primary cause of the role confusions. Thematic role assignment is also required with spatial prepositions such as in or above, and J.P.'s thematic role assignment was also severely impaired with spatial prepositions. We capitalized on prior linguistic analyses and behavioural studies to design accurate tests of the semantics of spatial terms, spatial relations, and critical features of objects. Fine-grain semantic tests revealed that the semantics of spatial terms and objects was intact. We hypothesize that J.P.'s role confusions reflected a failure to integrate objects within semantic representations that define the thematic roles. Our data suggest that properties of objects and thematic roles are specified by distinct semantic processes, which have different brain localizations. J.P.'s lesion further suggests that left inferior frontal regions are critical in thematic role assignment, thus contributing to the understanding of the linguistic functions of these regions.