. Six-month Old Babies Prefer to Watch Asl Over Pantomimes. B. Deaf Mothers Sometimes Fail to Produ

Lang Cogn Process. Author manuscript; bachelor in PMC 2012 Jan one.

Published in terminal edited form as:

PMCID: PMC3167215

NIHMSID: NIHMS219015

Sign language and pantomime product differentially engage frontal and parietal cortices

Karen Emmorey

San Diego State University

Stephen McCullough

San Diego State Academy

Sonya Mehta

University of Iowa

Laura L. B. Ponto

University of Iowa

Thomas J. Grabowski

Academy of Iowa

Abstract

We investigated the functional organization of neural systems supporting linguistic communication production when the main language articulators are also used for meaningful, but non-linguistic, expression such as pantomime. Fourteen hearing non-signers and 10 deaf native users of American Sign Language (ASL) participated in an H₂ ¹⁵O-PET study in which they generated activeness pantomimes or ASL verbs in response to pictures of tools and manipulable objects. For pantomime generation, participants were instructed to "show how y'all would utilise the object." For verb generation, signers were asked to "generate a verb related to the object." The objects for this condition were selected to elicit handling verbs that resemble pantomime (e.g., TO-HAMMER (hand configuration and movement mimic the act of hammering) and non-handling verbs that exercise non (e.1000., Cascade-SYRUP, produced with a "Y" handshape). For the baseline task, participants viewed pictures of manipulable objects and an occasional not-manipulable object and decided whether the objects could exist handled, gesturing "yes" (thumbs up) or "no" (hand wave). Relative to baseline, generation of ASL verbs engaged left junior frontal cortex, but when non-signers produced pantomimes for the same objects, no frontal activation was observed. Both groups recruited left parietal cortex during pantomime production. Withal, for deaf signers the activation was more extensive and bilateral, which may reverberate a more complex and integrated neural representation of mitt actions. Nosotros conclude that the production of pantomime versus ASL verbs (even those that resemble pantomime) appoint partially segregated neural systems that support praxic versus linguistic functions.

1.one Introduction

For sign languages, the chief language articulators (the hands) are too used for meaningful, merely non-linguistic, expression, eastward.yard., pantomime product or emblematic gestures such as "thumbs up/down." In dissimilarity, the use of the speech articulators to express meaningful non-linguistic information is relatively limited, e.g., song imitation, a few emblematic facial gestures such as sticking out i's tongue. Furthermore, the sign articulators perform everyday actions that are clearly related to many symbolic gestures (due east.g., object manipulation), whereas everyday actions of the speech communication articulators (e.g., chewing, swallowing) are normally unrelated to symbolic gesture. There is also convincing evidence for a diachronic human relationship between certain symbolic gestures and lexical signs within a signing community. For example, the "departure" gesture used beyond the Mediterranean was incorporated into French Sign Linguistic communication as the verb PARTIR, "to leave" (Janzen & Shaffer, 2002), and the Primal American gesture indicating a small creature has been incorporated into Nicaraguan Sign Language every bit the classifier class pregnant "pocket-sized animal" (Kegl, Senghas, & Coppola,1999). Although signs may take a gestural origin, they differ in systematic means from gesture. For example, signs exhibit duality of patterning and hierarchical phonological structure (east.g., Sandler & Lillo-Martin, 2006), whereas gestures are holistic with no internal structure (McNeill, 1992). Signs vest to grammatical categories, and gestures do not. Sign production involves lemma choice and phonological code retrieval (Emmorey, 2007), whereas pantomimic gesture involves visual-motoric imagery of action production. Here, we investigate the engagement of neural substrates that might differentiate sign product from gesture production.

We focus on verbs in American Sign Language (ASL) that resemble pantomimic actions: handling classifier verbs in which the hand configuration depicts how the human hand holds and manipulates an musical instrument. For case, the sign Castor-HAIR is made with a grasping handshape and a "brushing" motion at the head. Such verbs are almost often referred to equally classifier verbs because the handshape is morphemic and refers to a belongings of the referent object (e.one thousand., the handle of a brush; Supalla, 1986; see papers in Emmorey, 2003, for discussion). The course of handling classifier verbs is quite iconic, depicting the paw configuration used to grasp and dispense an object and the motion that is typically associated with the object's manipulation. Given the pantomime-similar quality of these verbs, we investigated the neural bases for the production of pantomimes, handling classifier verbs (hereafter referred to as handling verbs), and ASL verbs that practise not exhibit such sensory-motoric iconicity (hereafter referred to equally non-handling verbs). Nosotros restricted our investigation to pantomimes and excluded emblems considering both signs and pantomimes can easily be elicited with moving picture stimuli and because pantomimes are similar in form to handling verbs.

1.2 Dissociations betwixt sign and pantomime product

Evidence that the ability to sign can exist dissociated from pantomime ability comes from two case studies of deaf aphasic signers (Corina et al., 1992; Marshall et al., 2004). Corina et al. (1992) described the case of WL who had a big fronto-temporo-parietal lesion in the left hemisphere. WL exhibited poor sign comprehension, and his signing was characterized by phonological and semantic errors with reduced grammatical structure. A relevant example of a phonological error by WL is his production of the sign SCREWDRIVER, which iconically depicts the shaft of a screwdriver (index and middle fingers extended) twisting into the palm of the non-dominant hand. He substituted an A-bar handshape (fist with thumb extended) for the target handshape. In contrast to his sign production, WL was unimpaired in his power to produce pantomime. For example, instead of signing DRINK (a curved "C" hand moves toward the mouth, with wrist rotation – as if drinking), WL cupped his hands together to grade a pocket-sized bowl. WL was able to produce stretches of pantomime and tended to substitute pantomimes for signs, even when pantomime required more than complex movements. Similarly, Marshall et al. (2004) written report some other case of a deaf aphasic signer, "Charles," who likewise had a left temporo-parietal lesion and exhibited sign anomia that was parallel to speech anomia. For example, his sign finding difficulties were sensitive to sign frequency and to cueing, and he produced both semantic and phonological errors. However, his gesture product was intact and superior to his sign production even when the forms of the signs and gestures were very similar. These lesion data signal that the neural systems for sign language and pantomime product are not identical, just it is unlikely that these neural systems are completely distinct and contained. To our knowledge, there are no existing reports of brain-damaged patients with intact signing ability but impaired power to produce pantomimic gestures, which suggests that the neural systems supporting pantomimic communication might too exist involved in sign language processing (MacSweeney, Capek, Campbell, & Woll, 2008).

To place overlapping neural substrates as well as those that might differentiate between sign and gesture production, we elicited signs and pantomimes from neurologically-intact deaf signers and from hearing not-signers. In the written report reported hither, participants viewed pictures of manipulable objects (e.chiliad., a saw, a fork) and were asked to generate either an associated ASL verb (deaf signers only) or a pantomime showing how the object is typically used (run across Figure 1). In a control task, participants viewed manipulable objects and an occasional not-manipulable object (e.g., a house) and were asked to determine whether the objects could be held in the hand, gesturing either "yes" (thumbs upward) or "no" (hand wave). The transmission response controlled for mitt motion, and the determination task controlled for semantic processing of manipulable objects. This relatively high-level control chore immune usa to examine the specific neural regions that are engaged when participants pantomime an activeness or produce an ASL verb associated with an object, equally distinct from recognizing the object and its manipulability.

Analogy of sample object pictures and the verbs or pantomimes generated in response to these stimuli.

1.iii Neural substrates for pantomime production past hearing non-signers vs. sign production by deaf signers

Previous studies of pantomime production in hearing non-signers point to a disquisitional role for left parietal cortices. In a positron emission tomography (PET) written report, Rumiati et al. (2004) reported increased neural activity in a dorsal region of the inferior parietal lobule (IPL) when participants pantomimed the use of a pictured object compared to naming that object. Using fMRI, Imazu et al. (2007) also found increased activity in left IPL when Japanese participants pantomimed picking upward objects with chopsticks, compared to the actual use of chopsticks. Rumiati et al. (2004) hypothesize that left IPL is involved in the retrieval of object-related action schemata during pantomime execution (meet likewise Buxbaum, 2001). In add-on, several studies written report increased neural action in the left superior parietal lobule (SPL) when participants pantomime using tools compared to a motoric baseline task (Choi et al., 2001; Moll et al., 2000). Left SPL may be involved in the on-line arrangement of arm and hand actions in relation to an imagined object in space. Superior parietal cortex has been implicated in the representation of dynamic trunk schemas that monitor the location of the hand and arm during execution of movements (Khan et al., 2005; Wolpert, Goodbody, & Husain, 1998).

Left parietal cortices are besides recruited during sign linguistic communication product (for reviews see Corina & Knapp, 2006; MacSweeney et al., 2008). Greater activation has been reported in left inferior and superior parietal lobules for signing compared to speaking (Braun et al., 2001; Emmorey, Mehta, & Grabowski, 2007). Emmorey et al. (2007) suggested that the left superior parietal lobule may be involved in proprioceptive monitoring during sign linguistic communication production. The inferior parietal lobule, specifically, the supramarginal gyrus (SMG), has been implicated in lexical-phonological processing for sign language. Corina et al. (1999) constitute that direct stimulation of SMG resulted in semantic errors and handshape substitutions in a picture naming task.

In addition, phonological similarity judgments in sign language elicited activation in the superior portion of left SMG, extending into SPL (MacSweeney, Waters, Brammer, Woll, & Goswami, 2008). In add-on to left parietal cortex, sign linguistic communication production depends upon left inferior frontal cortex. Impairment to this region causes sign linguistic communication aphasia characterized by non-fluent signing and production errors (Poizner, Bellugi, & Klima, 1987; Hickok et al., 1996). During overt signing, several studies report meaning neural action in left junior frontal cortex (BA 44, 45, 46, and 47) (Corina et al., 2003; Braun et al. 2001; Emmorey et al., 2004; Petitto et al., 2000). In a conjunction analysis, Emmorey et al. (2007) establish that Broca's area (BA 45) was equivalently engaged for both sign and word product. The lexical-level functions of the left inferior frontal cortex are varied, but include phonological encoding, syntactic computations, and lexical semantic processing.

It is unclear whether the left junior frontal gyrus plays a critical role in pantomime production (Frey, 2008). Neuroimaging results are mixed, with some studies reporting left IFG activation (Hermsdörfer et al., 2007; Peigneux et al., 2004; Rumiati et al., 2004), while others report no left IFG activation during pantomime production (Choi et al., 2001; Imazu et al., 2007; Moll et al., 2000). Activation in left IFG is generally attributed to the retrieval and processing of object semantics associated with tool use pantomime. Nonetheless, an impairment of the semantic knowledge of tools and manipulable objects can occur in the face up of an intact ability to produce tool-use pantomimes (Buxbaum, Schwartz, & Carew, 1997; Rosci, Chiesa, Laiacona, & Capitani, 2003). Impairments in pantomime product are nigh oft observed with damage to left parietal cortex (eastward.g., Buxbaum, Johnson-Frey, & Barlett-Williams, 2005; Rothi, Heilman, & Watson, 1985), although there is some evidence that harm to left IFG is also associated with poor pantomime performance (Goldenberg et al., 2007).

Thus, in that location is testify that both sign and pantomime product engage left fronto-parietal regions. Semantic processes are associated with junior frontal cortex, while parietal regions are associated with manual articulation, object manipulation, and visually guided movements. Past straight comparing sign and pantomime production by deaf signers, we can examine the extent to which distinct frontal and parietal regions are engaged during lexical verb retrieval and production compared to the production of familiar, non-linguistic actions. We predict that both verb generation and pantomime generation volition engage fronto-parietal regions, but each may have a singled-out neural signature. Specifically, we predict that verb generation volition appoint the left inferior frontal cortex to a greater extent than pantomime production because central linguistic processes are associated with this area (e.g., lexical search, phonological encoding, semantic processing) and because tool-utilize pantomime does not always entail semantic processing of the object (e.k., Rosci et al., 2003). Nosotros also predict that verb and pantomime generation volition exhibit distinct patterns or levels of activation within parietal cortex, reflecting differences in the lexical versus praxic functions that underlie these tasks.

To further investigate the neural systems that underlie pantomime versus sign production, nosotros included a group of hearing non-signers in the report. These participants produced pantomimes for two sets of manipulable objects. For 1 fix, the deaf signers were asked to generate linguistic responses (ASL handling verbs related to the object), while the hearing non-signers were asked to produce pantomimes (probable to exist like in grade to the ASL treatment verbs). The between-group comparison for this set of objects provides an additional measure of the stardom betwixt the neural systems that support sign language versus pantomime production. For the second set of objects, both deafened and hearing participants were asked to generate pantomimes related to object utilize. The between-grouping comparison for these objects will reveal whether pantomime production past deaf signers and by hearing non-signers engages similar cortical regions. It is possible that knowledge and employ of ASL may alter the neural systems that are recruited for pantomime production.

Recently, Corina et al. (2007) reported that deaf signers did not engage frontal-parietal cortices when passively viewing transmission actions that were cocky-oriented (e.k., scratch neck, lick lips, rub shoulder) or object-oriented (e.grand., bite an apple, read a book, pop a balloon; the model handled the objects). In contrast, hearing not-signers showed robust activation in frontal-parietal regions when viewing these actions. Corina et al. (2007) hypothesized that life-long experience with a visual linguistic communication shifts neural processing of man deportment to extrastriate association areas, regions that were particularly active for the deaf signers. Corina et al. (2007) suggested that this shift arises because signers must actively filter human actions to chop-chop distinguish communicative from non-chatty actions for further linguistic processing. Such preprocessing of homo action is non required for non-signers. Similarly, Emmorey, Xu, Gannon, Goldin-Meadow, and Braun (2009) establish that passively viewing pantomimes strongly activated frontal-parietal regions in hearing not-signers, whereas activation was primarily observed in bilateral middle temporal regions in deaf signers. Emmorey et al. (2009) hypothesized that deafened signers recognize pantomimes quickly and relatively automatically and that this ease of processing leads to a substantial reduction of neuronal firing within frontal-parietal cortices.

1.4 Handling ("pantomimic") verbs vs. non-handling verbs

Finally, given that ASL handling verbs are nearly identical in form to pantomimic gestures (run into Figure ane), information technology was important to likewise elicit not-treatment ASL verbs in society to appraise whether the pantomimic properties of handling verbs affects the pattern of neural activeness in frontal or parietal cortices. Nosotros did non anticipate differences between handling and non-handling verbs because we previously found very similar results for the product of treatment verbs (e.g., Brush-Hair, ERASE-BLACKBOARD) and non-treatment "general verbs" (eastward.grand., YELL, SLEEP) (Emmorey et al., 2004). Specifically, the production of both verb types in a motion-picture show-naming task engaged left junior frontal cortex and the left superior parietal lobule (the baseline comparison was a face-orientation judgment task), and there were no significant differences in neural activation betwixt the two verb types. However, the contrast between pantomimes and not-handling verbs may reveal differences that are non observed when pantomimes are contrasted with handling verbs. Specifically, considering the phonological course of not-handling verbs does not involve reaching movements or grasping-type hand configurations (different object-use pantomimes or handling verbs), we may detect a greater divergence within parietal cortices between pantomimes and non-handling verbs than between pantomimes and handling verbs.

In sum, nosotros investigated whether deafened signers and hearing non-signers differ with respect to neural regions engaged during the production of pantomimed actions and whether distinct patterns of neural activeness are observed for the production of pantomimed actions, pantomimic-like ASL verbs, and non-pantomimic ASL verbs past deaf signers.

2. Methods

2.1. Participants

Twenty-four correct-handed adults participated in the study. X participants (six males) were native deaf signers, anile xx–27 years (mean age = 23.two years). All had deaf parents and acquired ASL every bit their first language from birth. Viii participants had severe to profound hearing loss, and hearing loss level was unknown for two participants. All deaf participants used ASL as their primary and preferred linguistic communication. The other fourteen participants (seven males) reported normal hearing, had no knowledge of ASL, and were native English speakers (mean historic period = 28.vii years; range 22–41 years). All deafened and hearing participants had 12 or more than years of formal educational activity, and all gave informed consent in accordance with Federal and institutional guidelines.

2.2 Materials

To select objects that would elicit handling verbs as well equally non-handling verbs, 166 photographs of manipulable objects were presented to an contained grouping of 16 deaf signers. Pictures were presented using Psyscope software (Cohen et al., 1993) at the aforementioned rate every bit for the neuroimaging experiment (see below). Participants were asked to "produce a verb that names an action you lot would perform with that object." Instructions were given in ASL by a native signer. Nosotros selected 32 object pictures that consistently elicited treatment classifier verbs and 32 objects that consistently elicited non-handling classifier verbs (i.e., a bulk of participants produced the desired response). In addition, the object pictures that elicited treatment verbs were selected because they also elicited consistent and hands performed pantomimic gestures from hearing not-signers, as described below. For instance, objects in the treatment verb elicitation set included a scrub brush (TO-SCRUB), a sprint (THROW-Sprint), and a mug (TO-DRINK). Objects in the non-handling verb elicitation set included a ruler (TO-MEASURE), a pigment brush (TO-Pigment), and syrup (TO-POUR-SYRUP). Example stimuli and verbs are presented in Figure 1.

To select object pictures that would elicit clear and appropriate pantomimic gestures, we presented the same object photographs to a split grouping of twenty hearing non-signers and xvi deaf signers. Participants were asked "to pantomime how you would utilize that object or what you would do with that object." We selected 32 object pictures that elicited consequent and easily performed pantomimes from both participant groups (i.e., the majority of participants produced appropriate pantomimes with little hesitation). Objects in the pantomime elicitation set included a fork, a drill, a key, and scissors (come across Figure 1 for an example target pantomime). In add-on, these object pictures were selected because they elicited pantomimes from deaf signers that were distinct from ASL verbs associated with the objects. For instance, given a picture of a broom, participants produced a pantomime of sweeping with a broom, which does not resemble the ASL verb TO-SWEEP. Thus, nosotros could determine whether deaf participants in the neuroimaging study were producing the expected pantomimes, rather than generating ASL verbs.

Finally, to assess the iconicity and meaningfulness of the elicited handling verbs, pantomimic gestures, and non-handling ASL verbs, we presented video of the target ASL verbs and pantomimes produced by a deaf native signer to another group of 10 hearing not-signers. Participants were asked to rate each gesture as having no meaning (rated as 0), weak meaning (rated as one), moderate/fairly articulate meaning (rated as ii), or absolute/strong direct pregnant (rated equally iii) and to write down the meaning of each gesture. The non-handling verbs (hateful rating = 0.70; SD = 0.30) were rated as less meaningful than the handling verbs (hateful rating = 1.73; SD = 0.39) and the pantomimic gestures (mean rating = i.60; SD = 0.33). The majority of participants were too able to identify the general pregnant of the target ASL handling verbs and the pantomimic gestures.

two.3 Procedure

Paradigm Conquering

All participants underwent MR scanning in a 3.0T TIM Trio Siemens scanner to obtain a 3D T1-weighted structural scan with isotropic 1 mm resolution using the post-obit protocol: MP-RAGE, TR 2530, TE iii.09, TI 800, FOV 25.6cm, matrix 256 × 256 × 208. The MR scans were used to confirm the absenteeism of structural abnormalities, assistance in anatomical interpretation of results, and facilitate registration of PET information to a Talairach-uniform atlas.

Positron emission tomography (PET) data were acquired with a Siemens/CTI Hr+ PET organization using the following protocol: 3D, 63 prototype planes, 15 cm axial FOV, four.5mm transaxial and four.2mm axial FWHM resolution. Participants performed the experimental tasks during the intravenous bolus injection of fifteen mCi of [^xvO]water. Arterial blood sampling was not performed.

Images of rCBF were computed using the [¹⁵O]water autoradiographic method (Herscovitch et al, 1983, Hichwa et al, 1995) as follows. Dynamic scans were initiated with each injection and continued for 100 seconds, during which twenty five-second frames were acquired. To decide the time course of bolus transit from the cognitive arteries, time-activity curves were generated for regions of interest placed over major vessels at the base of operations of the brain. The 8 frames representing the start forty seconds immediately afterwards transit of the bolus from the arterial puddle were summed to make an integrated 40-second count image. These summed images were reconstructed into 2mm pixels in a 128x128 matrix.

Tasks

While undergoing PET scanning, participants performed the post-obit tasks when presented with object photographs: a) generate an associated ASL verb (deafened simply), b) generate an object-related pantomime, or c) make a handling decision. For the verb generation task, one set of objects elicited handling verbs and another elicited non-handling verbs. For both sets of objects, deaf participants were instructed in ASL past a deaf native signer to produce a verb that named an action performed with the object. They were not specifically told to produce handling or non-handling verbs. For the pantomime generation task, both participant groups were instructed to "show me how you would use the pictured object." Hearing participants were too instructed to generate pantomimes for the same object stimuli that were used to elicit handling verbs from the deaf participants. For the control task, participants were instructed to decide whether the pictured object could be held in the hand and to indicate their response with either a "yes" gesture (thumbs up) or a "no" gesture (palm downwardly, horizontal mitt moving ridge). Three to four objects in each cake of 16 were not manipulable (due east.g., a highway, a radio tower, a dam). 4 practice items preceded each target task.

For all tasks, picture stimuli were presented for two south followed past a 500 ms inter-stimulus-interval. For each task condition, 32 picture stimuli were presented to participants using I-glasses SVGA Pro goggles (I-O Display Systems; Sacramento, CA), in two split up blocks of 16 pictures. For each block, the motion-picture show stimuli were presented from v s subsequently the injection (approximately seven–10 s before the bolus arrived in the brain) until 35 s afterward injection.

Participants' responses were recorded during the PET report by either a native ASL signer or a native English speaker. Responses were also digitally recorded for confirmation and later analysis; yet, due to equipment malfunction, the responses of five deaf and iii hearing participants were not recorded. Participants were immune to produce verbs and/or gestures with both hands (a flexible intravenous cannula was used for the [¹⁵O]water injection, which permitted arm movement after the injection).

Spatial normalization

PET data were spatially normalized to a Talairach-compatible atlas through a series of coregistration steps (run into Damasio et al., 2004; Grabowski et al., 1995, for details). Prior to registration, the MR information were manually traced to remove extracerebral voxels. Talairach space was constructed directly for each participant via user-identification of the inductive and posterior commissures and the midsagittal aeroplane on the 3D MRI data prepare in Brainvox. An automated planar search routine defined the bounding box and piecewise linear transformation was used (Frank et al., 1997), as defined in the Talairach atlas (Talairach and Tournoux, 1988). Later Talairach transformation, the MR data sets were warped (AIR 5^th order nonlinear algorithm) to an atlas infinite constructed by averaging fifty normal Talairach-transformed brains, rewarping each brain to the average, and finally averaging them again, analogous to the procedure described in Woods et al., (1999). Additionally, the MR images were segmented using a validated tissue division algorithm (Grabowski et al., 2000), and the gray thing partition images were smoothed with a 10 mm kernel. These smoothed gray matter images served as the target for registering participants' PET information to their MR images.

For each participant, PET data from each injection were coregistered to each other using Automatic Epitome Registration (AIR 5.25, Roger Woods, UCLA). The coregistered PET information were averaged, and the mean PET paradigm was and so registered to the smoothed grey matter partition using FSL's linear registration tool (Jenkinson & Smith, 2001; Jenkinson et al., 2002). The deformation fields computed for the MR images were then applied to the PET data to bring them into register with the Talairach-compatible atlas. After spatial normalization, the PET data were smoothed with a sixteen mm FWHM Gaussian kernel using complex multiplication in the frequency domain. The final calculated voxel resolution was 17.9 x 17.9 10 18.9 mm. PET data from each injection were normalized to a global hateful of 1000 counts per voxel.

Regression Analyses

PET data were analysed with a pixelwise general linear model (Friston et al. 1995). Regression analyses were performed using tal_regress, a customized software module based on Admirer's to the lowest degree squares routines (Miller, 1991) and cross-validated against SAS (Grabowski et al, 1996). Regression analyses were performed separately for data from the deaf ASL participants and hearing not-signing participants. Models for both regression analyses included participant and job condition effects. For the deaf ASL participants, half-dozen contrasts were tested: 1) pantomime – decision, 2) handling verbs – decision, 3) non-handling verbs – decision, iv) pantomime – handing verbs, 5) pantomime – non-handling verbs, and 6) handling verbs – non-handling verbs. For the hearing not-signing participants, only contrasts 1, 2, and 4 were tested, and contrasts two and four differed for these participants in that they pantomimed when presented the stimuli for the handling verbs.

In addition, two random furnishings analyses were performed to assess differences betwixt ASL and non-signing participant groups in the pantomime and handling verb atmospheric condition. For these analyses, data were averaged across task condition (either the pantomime job or the handling verbs/pantomime task). The deviation epitome betwixt the task and control task was used as the dependent mensurate to examination for differences betwixt deaf and non-signing participants.

Contrasts were tested with t-test and thresholded using random field theory (Worsley 1994), with a Familywise type I error set at alpha less than 0.05. In accord with our hypotheses, an a priori search volume was used that included left frontal lobe, left temporal lobe, and bilateral parietal lobes. The search book was delineated by manual tracing on the averaged MR and measured 258 cm3, calculated to comprise 43 resels. A whole encephalon search was likewise performed to look for areas of activation outside of this restricted search volume.

3.1 Results

For the behavioral results, non-responses were scored equally wrong for both verbs and pantomimes. Wrong verb responses included naming the object (rather than generating a verb) or producing a non-target verb. Incorrect pantomime responses included producing a not-target pantomime or a not-pantomimic gesture, such as a hesitation gesture (e.g., waving the hand as if searching for an idea). Deafened participants were more authentic when generating handling verbs (93.13% correct) than when generating non-treatment verbs (77.12% right), t(nine) = −3.414, p < .01. Incorrect responses for the non-handling verbs were primarily not-target verb responses in which the participant produced a treatment, rather than a not-treatment verb. Response accuracy for handling verbs did not differ significantly from response accurateness for generating pantomimes (91.88%), t(9) = 0.612, p = .55. Hearing participants were equally accurate as deafened participants when generating pantomimes (94.01%), and response accuracy was similar for the "handling verb" object pantomimes (92.89%; deaf participants generated ASL verbs for these objects).

Tables 1 and ​ 2 present the local maxima of areas of increased activity for each target task contrasted with the command task for deaf and hearing participants, respectively. As shown in Effigy 2, when deaf participants generated pantomimes, they exhibited extensive bilateral activation in the superior parietal lobule (SPL) and forth the precentral gyrus; in contrast, when hearing participants generated pantomimes to the same stimuli, activation was left lateralized within SPL. The hearing participants as well exhibited activation inside left middle/inferior temporal cortex (BA 37) when producing pantomimes for the same objects that elicited pantomimes from the deaf signers and when producing pantomimes for the objects that elicited handling verbs from signers ("handling verb" pantomimes). Finally, for both deaf and hearing participants, pantomime generation resulted in significant activation within the right cerebellum. For deaf signers, correct cerebellar activation was as well observed during the generation of non-treatment verbs.

Results of the dissimilarity betwixt pantomime production and baseline task in the deaf participants (row A) and hearing not-signers (row B). Row C shows the results for the betwixt subjects contrast (deaf – hearing non-signers) for the same job comparisons. The colour overlay reflects pregnant increased activity at an uncorrected level, and regions in red reflect increased activeness at a RFT corrected level (p < 0.05). Compared to the baseline job, pantomime production past both deafened and hearing participants resulted in greater activity in left superior parietal lobule. For the deaf participants, the increased activity was more extensive, encompassing the correct parietal lobule, the left and right precentral gyrus, and right supplementary motor area.

Table 1

Summary of PET activation results for deafened participants. Local maxima of areas with increased activation for the target tasks in contrast to the control job and for pantomime production contrasted with ASL verb product. Results are from the a priori search volume (critical t(86) = 4.03) and the whole brain analysis (critical t(86) = four.60; indicated by an asterisk).

Region	Side	X	Y	Z	T
Pantomime
Superior frontal gyrus (BA 6)	L/R	0	0	+66	five.02*
Supplementary Motor Expanse (BA half dozen)	R	+5	−one	+58	5.05*
Precentral gyrus (BA half-dozen)	R	+26	−9	+49	5.13*
Precentral gyrus (BA 4)	L	−29	−29	+48	half-dozen.54*
Superior parietal lobule (BA 7)	Fifty	−9	−66	+58	4.76
	R	+24	−55	+sixty	four.77
Cerebellum	L	−28	−49	−26	5.32*
	R	+24	−48	−28	5.24*
Treatment Verbs
Junior frontal gyrus (BA 46)	L	−43	+37	+11	4.50
	50	−33	+36	+three	4.53
Inferior frontal gyrus (BA 44)	L	−42	+9	+18	iv.34
Cerebellum	R	+22	−52	−28	5.74*
Not-handling verbs
Junior frontal gyrus (BA 46)	L	−52	+33	+xiii	3.36‡
Cerebellum	R	+4	−58	−19	v.l*
	R	+22	−51	−27	v.71*
	R	+10	−69	−28	5.43*
Pantomime > handling verbs
Superior parietal lobule (BA vii)	R	+28	−55	+54	4.62
Pantomime > non-handling verbs
Precentral gyrus (BA six)	R	+25	−7	+49	5.22*
Intraparietal sulcus	L	−41	−38	+47	v.twoscore
Junior parietal lobule (BA twoscore)	R	+45	−33	+41	v.55
Superior parietal lobule (BA vii)	R	+29	−53	+54	5.64

Table two

Summary of PET activation results for hearing participants. Local maxima of areas with increased activation for the target tasks in comparing to the control task. Results are from the a priori search volume (critical t(121) = 3.96) and the whole brain analysis (critical t(121) = 4.51; indicated by an asterisk).

Region	Side	X	Y	Z	T
Pantomime
Superior parietal lobule (BA 7)	50	−30	−43	+60	iv.84
Inferior temporal gyrus (BA 37)	L	−55	−65	+ii	4.05
"Handling verb" pantomime
Superior parietal lobule (BA 7)	Fifty	−26	−51	+63	4.44
Intraparietal sulcus	L	−31	−33	+32	four.94
Junior temporal gyrus (BA 37)	L	−49	−61	+1	iv.17
Cerebellum	R	+26	−52	−29	iv.90*

Unlike pantomime generation, when deafened signers produced ASL treatment verbs, activation was observed at two local maxima inside the left inferior frontal gyrus (IFG) (see Figure iii). Similarly, when deaf participants generated ASL non-handling verbs, activation was observed in left IFG, but this neural activity was weaker, reaching significance only at an uncorrected level. When pantomime product was directly contrasted with ASL treatment or not-treatment verb production, greater neural activity was observed for pantomimes inside the right SPL (see Tabular array 1). The contrast betwixt pantomime production and the non-handling ASL verbs also revealed additional regions of greater neural activity inside the right precentral gyrus and bilateral inferior parietal lobule. There were no neural regions that were significantly more active for handling verb than for pantomime product. For non-handling verbs compared to pantomimes, greater activation was observed in a small region within the posterior right cerebellum (+28, −85, −25; t = 4.68). Finally, a directly contrast between the generation of handling verbs and non-treatment verbs revealed no differences in neural activeness.

Results of the contrast between handling verb generation and baseline task in the deafened participants (row A) and pantomime production (for the same 'treatment verb' stimuli) and baseline task in hearing not-signers (row B). Row C shows the results for the between subjects contrast (deaf – hearing non-signers) for the same task comparisons. The color overlay reflects pregnant increased action at an uncorrected level, and regions in red reverberate increased activity at a RFT corrected level (p < 0.05). Increased action in the inferior frontal gyrus is observed in the deaf signers but non the hearing participants.

We next conducted random event analyses comparison the deaf and hearing groups (encounter Table 3). For pantomime product, greater activation was observed in right SPL and correct precentral gyrus for deafened signers compared to hearing non-signers (see Figure iv); greater activation was observed within the retrosplenial area and hippocampus/parahippocampal gyrus for hearing non-signers compared to deaf signers. When deaf signers generated ASL verbs and hearing not-signers generated pantomimes to the aforementioned objects, we observed greater activation within left IFG and the middle frontal gyrus (BA 10) for deaf signers (meet Effigy 5). At that place were no regions that were significantly more active for hearing not-signers than for deaf signers for this dissimilarity.

Table 3

Summary of PET activation results for the 2 random effects analyses. Local maxima of areas with increased activation for the target tasks in comparing to the control task. Results are from the a priori search book (critical t(22) = 4.88) and the whole brain analysis (critical t(22) = 5.82; indicated by an asterisk).

Region	Side	X	Y	Z	T
Pantomime
Deaf > hearing
Precentral gyrus (BA 6)	R	+31	−12	+48	half-dozen.68*
Superior parietal lobule (BA 7)	R	+31	−46	+49	5.82
Hearing > deaf
Retrosplenium (BA 23)	Fifty	−2	−56	+15	5.35
Hippocampus/parahippocampal gyrus	Fifty	−32	−27	−4	half-dozen.42*
Handling verbs/pantomime
Deaf > hearing
Inferior/middle frontal gyrus (BA 46/10)	Fifty	−26	+40	+v	v.21
Middle frontal gyrus (BA ten)	L	−30	+54	−xi	6.24*
Hearing > deaf No regions were more active for hearing participants

4.1 Discussion

The production of ASL verbs past deaf signers engaged the left junior frontal gyrus (Figure 3A), replicating several previous studies of sign linguistic communication production (e.thou., Braun et al., 2001; Corina et al., 2003; Emmorey et al., 2007; Petitto et al., 2000). Verb generation tasks for spoken linguistic communication also consistently elicit activation in left IFG (due east.g., Buckner, Raichle, & Peterson, 1995; Klein et al., 1999). In contrast, activation in left inferior frontal cortex was not observed when either deaf signers or hearing not-signers generated pantomimic gestures in response to pictures of manipulable objects (Figures 2 and ​ 3B). During verb generation, left IFG activation has been argued to reflect semantic search and lexical selection processes (e.chiliad., Thompson-Schill et al., 1997), and such lexical processes are not required for pantomime generation. Nonetheless, it is possible that some covert verb generation may have occurred for the deafened signers during pantomime product considering the direct contrast betwixt ASL verb and pantomime generation did not result in differential activation in left IFG for ASL verbs. Signers may have occasionally retrieved an ASL verb related to the pictured object, even though their task was to generate a pantomime of the objects' use. Such covert lexical retrieval may have reduced the difference in neural action within left IFG for the direct contrast between ASL verbs and pantomimes.

Further evidence that ASL verb generation engages frontal regions to a greater extent than pantomime is constitute in the random effects analysis comparison deaf participants' product of handling verbs and hearing participants' production of pantomimes in response to the aforementioned objects (Tabular array iii). Deaf signers' product of ASL verbs engaged left anterior frontal cortex (BA 46/x) to a greater extent than hearing participants' production of pantomimes that resembled ASL handling verbs (Effigy 3C).

Equally predicted, in comparison to the baseline task, pantomime product engaged the left superior parietal lobule for both hearing and deafened participants (Figure 2). Activation inside the left inferior parietal lobule may not have been observed because the baseline task involved a determination of whether an object could be held in the hand, and the planning and representation of grasp configuration is associated with inferior parietal cortex (eastward.thou., Grafton et al., 1996; Jeannerod, Arbib, Rizzolatti, & Sakata, 1995). Consistent with this explanation, we observed increased activity in IPL when contrasting pantomime production with non-handling verb generation (but not with treatment verb generation).

Activation within SPL was bilateral and more than all-encompassing for deaf signers than for hearing non-signers, which may be due to the richer and more precisely articulated pantomimes that were produced past the deafened signers. Deaf signers produced crisp and clear manus configurations in their pantomimes (e.thou., depicting a specific type of grip on a paint brush handle), while the hearing participants produced more lax, cryptic handshapes. Deaf signers tended to produce longer pantomimes; for example, lx% of their pantomimes repeated the depicted action, compared to 46% for hearing participants. Signers also produced somewhat more than complex pantomimes. In particular, the signers produced more than 2-handed pantomimes (80% vs. 41% for hearing not-signers), and their pantomimes represented more aspects of an activity. For instance, in response to a picture of a teapot, both groups tended to produce a pouring gesture, but the deafened signers were more likely to simultaneously produce a gesture representing the teacup with their other hand. Deafened signers may be more practiced in pantomime generation than hearing people because ASL narratives and stories often include pantomimic gestures (Emmorey, 1999; Liddell & Metzger, 1998), and they too sometimes need to pantomime when communicating with hearing people. Contempo studies have shown that expertise can lead to an expanded neural representation for a given task or ability (eastward.g., Beilock et al., 2008; Landau & D'Esposito, 2006), and we hypothesize that the more all-encompassing SPL activation observed for the deaf signers may reflect experience-dependent changes in the neural network that supports pantomime product.

When hearing non-signers produced pantomimes, we observed significant activation in left junior temporal cortex (Figures 2 and ​ 3), which was not observed for deaf signers. Activation within the ventral stream may reverberate more extensive visual analysis of the object to exist manipulated past the hearing non-signers as they planned their object pantomimes. The group comparison revealed greater activation in the retrosplenial cortex and the hippocampus/parahippocampal gyrus for the hearing group than the deaf group. These regions are known to exist involved in the recall of episodic data, and we advise that activation in these regions arises because hearing participants may be more likely to generate pantomimes by retrieving a specific memory of how they used the object in the by. In contrast, deaf signers may be more likely to generate a pantomimed action on the wing without attempting to remember a particular instance of using the object.

The direct dissimilarity between pantomime production and the generation of both handling and non-handling verbs revealed activation in the correct superior parietal lobule specific to pantomimes (Table 1). Kroliczak et al. (2007) reported that pantomimed grasping of an object recruited right SPL in comparison to pantomimed reaching toward the object, while the comparison between real grasping vs. real reaching did not show a deviation in right SPL activation. Kroliczak et al. (2007) hypothesized that correct parietal cortex may be involved in less familiar pantomimes that crave additional spatial transformations and/or reprogramming of move kinematics that depend upon right-hemisphere processing. Chaminade, Meltzoff, & Decety (2005) likewise contend that correct superior parietal cortex is involved in the visuospatial assay of gestures, especially when spatial features are necessary to place an action to exist imitated. Such visuospatial analysis of an object to be grasped or of the movements needed to pantomime how an object is used are non required for the generation of ASL verbs. During verb generation, hand configuration and movement are retrieved as function of a phonological representation of a lexical sign – the form of the sign is not direct computed from an assay of the visuospatial characteristics of the stimulus object.

The contrast between pantomime production and the generation of not-handling verbs revealed boosted activation in the left inferior parietal lobule for pantomimes (Tabular array 1). Left IPL activation is likely due to the grasping hand configurations and movements that were used for the pantomimes. By definition, the not-handling verbs did not involve mitt configurations or motions that resembled how objects are grasped. The contrast betwixt handling verb and pantomime generation revealed no differential activation within left IPL. This pattern of results suggests that left IPL may play a role in the production of both pantomimes and ASL signs that involve similar grasping paw configurations and movements and/or that convey semantic information about object manipulation.

These results are relevant to the hypothesis that motor systems are involved in the semantic processing and representation of action words (e.g., Boulenger, Hauk, & Pulvermüller, 2009; Hauk, Johnsrude, & Pulvermüller, 2004; Rizzolatti & Craighero, 2004). The fronto-parietal regions engaged during the generation of ASL handling verbs and pantomimic gestures were not identical, despite very similar motoric patterns and action semantics. The production of "pantomimic" ASL verbs does non appear to involve cerebral embodiment processes that are involved in demonstrating object manipulation or apply. The findings are consistent, however, with the hypothesis that motor areas play a role in the conceptual representation of action verbs, every bit evidenced by the blueprint of contrasts within left IPL described above. In improver, Emmorey et al. (2004) found very like activation peaks in left premotor and left inferior parietal cortices for both ASL signers and English speakers when naming pictures of tool-based deportment (e.g. stir, hammer, scrub). Emmorey et al. (2004) proposed that activation within these regions reflects retrieval of noesis near the sensory- and motor-based attributes that partially constitute conceptual representations for these types of actions.

Compared to baseline, the generation of both treatment and not-handling verbs activated left inferior frontal cortex; however, activation was weaker for the non-treatment verbs. It is not clear why left frontal activation was reduced for the non-treatment verbs. Weaker left IFG activation during verb generation has been associated with lower demands on lexical selection and with less lexical interference (due east.grand., Nelson et al., 2009; Thompson-Schill et al., 1997). Even so, more lexical competition may have occurred for non-treatment verbs considering participants sometimes produced non-target handling verbs, which suggests more possible responses existed for the non-handling verbs. However, it is possible that retrieval of these verbs was less enervating because they are morphologically less circuitous than handling classifier verbs. Specifically, non-handling verbs are monomorphemic, whereas the handshape of classifier verbs represents a distinct morpheme that specifies either an instrument or treatment agent (Benedicto & Brentari, 2004; see likewise papers in Emmorey, 2003).

In sum, the production of pantomime and ASL verbs relies on partially distinct neural systems. Pantomime production engaged superior parietal cortex bilaterally for deaf signers, while verb production engaged left inferior frontal cortex. Pantomime production by hearing non-signers also did non engage left inferior frontal cortex, indicating the left frontal cortex is non critical to the execution of pantomimes. Interestingly, the neural networks for pantomime generation were non identical for the deaf and hearing groups. Deaf signers recruited more than extensive regions within superior parietal cortex, perhaps reflecting more than practice with pantomime generation; whereas, hearing non-signers recruited neural regions associated with episodic memory retrieval, maybe reflecting a recall-based strategy for pantomime generation. In line with previous research, nosotros suggest that left inferior frontal cortex subserves lexical retrieval functions associated with verb generation (even for verbs that mimic hand actions), whereas superior parietal cortex subserves the execution of praxic motor movements involved in the pantomimed utilize of an object.

Acknowledgments

This research was supported by a grant from the National Institute on Deafness and other Communication Disorders (R01 DC006708). We give thanks Jocelyn Cole, Jarret Frank, Franco Korpics, and Heather Larrabee for their help with the study.

Correspondent Data

Karen Emmorey, San Diego State University.

Stephen McCullough, San Diego State University.

Sonya Mehta, University of Iowa.

Laura L. B. Ponto, University of Iowa.

Thomas J. Grabowski, Academy of Iowa.

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