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2.1.1.! Mid-cingulate! cortex On! the! dorsomedial! wall! of! the! frontal! lobe,! the! MCC! lies! ventral! to! the! pre-SMA! and! SMA! and! dorsal! to! the! anterior! part! of the! corpus! callosum! just! behind! the! ACC! region,! which! extends anteriorly! and! ventrally! below! the! genu! (Fig.! 2b,! d).! The! MCC! comprises! cytoarchitectonic! area! 24,! the! dorsalmost! subdivision! of which! (24c')! and! area! 32 are! partly! buried! in! the! cingulate! sulcus;! and! the! immediately! ventral! perigenual! areas! 24a'! and! 24b'! on the! cingulate! gyrus!, !, 2009. ,
! Relative! to! the! more! caudal! cingulate! motor! areas, the! CMAr! has! dense! connections! with! the! lateral! prefrontal! cortex and! the! pre-SMA,! but! weak! connections! with! the! primary! motor cortex,! spinal! cord! and, !, 1991. ,
, !, 1994.
! Consistent! with! this! pattern! of! anatomical connectivity,! electrical! stimulation! of! the! CMAr! evokes! movements less! clearly! in! comparison! with! the! caudal! CMAs! (Dum! and! Strick, 1991. ,
! As! such,! the! CMAr! appears to! contribute! more! to! the! higher-order! initiation! and! preparation of! motor! responses,! while! the! more! caudal! areas! appear! to! be! more directly! involved! in! motor! execution.! In! contrast! to! the! MCC,! the ACC,! which! comprises! pregenual! and! subgenual! areas! 24c! and! 32, as! well! as! area! 25! on! the! subcallosal! gyrus, ! However,! CMAr! neurons! exhibit! a! stronger association! with! movement! initiation! and! respond! earlier! before movement! onsets!, 1991. ,
! further! demonstrated! increased! serotoninergic! and! GABAergic! receptor! densities in! the! ACC! in! comparison! with! the! MCC.! These! findings! suggest! an increased! involvement! of! the! ACC! in! emotional! and! arousal! related functions! (Paus,! 2001).! Of! direct! relevance! to! the! current! review! is the! cingulate! vocalization! area!, !, 1970. ,
! 1996)! and! the! PAG! region! (An! et! al.,! 1998)! which provide! direct! access! to! affective! facial! movements! and! affective vocal! patterns,! respectively.! Together! with! direct! connections! to the! amygdala,! spinothalamic! system! and! ventral! striatum,! the! anterior! MCC! appears! to! play! an! important! role! in! primate! emotional expression! via! facial! and! vocal! movements, ! This! region,! at! the! rostral! end! of! CMAr,! has! direct! projections! to! the! facial! motor! nuclei, 1993. ,
! innate! affective! calls! can! be! directly! evoked! by ACC/MCC! stimulation! (Jürgens! and! Ploog, !, 1967. ,
, Lesions! in! this! region! of! the! cingulate! gyrus! reduce! spontaneous calling! in! socially! isolated! settings! (MacLean! and! Newman,! 1988) and! severely! impair! the! ability! to! produce! calls! in! response! to! discriminant! stimuli!, !, 1981.
! demonstrating its! critical! role! in! the! voluntary! production! of! spontaneous! and! conditioned! calls.! In! humans,! electrical! stimulation! of! the! MCC! evokes orofacial! and! tongue! movements! but! less! consistently! produces affective! vocalizations! in! comparison! with! similar! stimulation! in NHPs! (Talairach! et! al.,! 1973).! However,! more! recent! investigations demonstrated! that! ACC/MCC! stimulation! triggered! laughter,! along with! positive! emotions, !, 1974. ,
! These! patients! typically! display! a! reduction! in! spontaneous! speech! and! increased hesitation! when! responding! to! direct! questions,! suggesting! that the! human! MCC! may! be! involved! in! the! motivational! initiation of! speech.! However,! the! MCC! is! not! critical! for! volitional! speech production! in! humans:! in! a! comprehensive! review! of! cingulectomy! cases,! Brotis! et! al.! (2009)! reported! that! patients! with! bilateral ACC/MCC! resections! could! generally! recover! their! verbal! abilities and! social! functioning! with! time. 2.1.2.! SMA! and! pre-SMA The! SMA! and! pre-SMA! represent! architectonically! different! brain! regions! that! are! associated! with! distinct! connectivity patterns.! In! the! human! brain,! the! SMA! has! been! related! to! cytoarchitectonic! area! 6a! and! the! pre-SMA! to! cytoarchitectonic! area! 6a as! defined! by! Vogt! and! Vogt! (1919).! In! the! macaque! monkey,! the SMA! has! also! been! referred! to! as! area! F3! and! the! pre-SMA! as! area F6!, ! Patients! with! ACC/MCC! lesions! experience! akinetic! mutism,! which! is! associated! with! a! lack! of! motivation! to! speak, 1988. ,
! Further! support! for! SMA's! direct! involvement NHPs! and! monkeys!, !, 1987. ,
! the! leg! representation! is! situated! in the! most! caudal! part! of! the! SMA,! followed! by! the! hand,! and! finally the! orofacial! representation! around! the! pre-SMA/anterior! SMA! border.! Concerning! the! pre-SMA,! movements! could! also! be! evoked! by electrical! stimulations,! but! these! were! less! consistent! and! required higher! currents, ! these! somatotopic! maps, 2008. ,
! stimulations! in! the! pre-SMA/anterior! SMA! region! produces! orofacial! movements! but! not! vocalizations! ,
! led! to! increased latencies! of! spontaneous! and! conditioned! calls.! Notably,! the! areas that! Kirzinger! and! Jürgens! (1982)! referred! to! as! the! "anterior"! and "posterior"! SMAs! corresponded! to! the! Vogt! and! Vogt! (1919)! areas 6a! and! 6a,! respectively,! and! therefore,! are! considered! pre-SMA and! SMA! in! this! article.! In! addition,! monkeys! with! lesions! in! the pre-SMA/anterior! SMA! area! still! successfully! perform! vocal! operant! conditioning! tasks! (Sutton! et! al.,! 1985).! As! such, ! reported! that! lesions! invading the! anterior,! but! not! the! posterior! part! of! the! SMA, 1982. ,
, SMA! have! been! associated! both! with! orofacial! and! tongue! movements! as! well! as! vocalizations!, 1991.
! As! such,! the! cortical! orofacial! region! in! NHPs! appears! to! be critical! for! the! volitional! control! of! tongue! and! orofacial! musculature! but! not! vocalizations. As! in! NHPs,! the! human! motor! cortex! has! direct! projections! to the! various! articulatory! nuclei! implicated! in! vocalizations! (Kuypers, 1958b).! However,! additional! direct! corticobulbar! projections! to the! nucleus! amibiguus! are! found! in! the! human! motor! cortex, which! enable! direct! neocortical! control! over! laryngeal! musculature! (Simonyan! and! Horwitz, ! These! vocalization-triggering! sites! were! clustered cortex! showed! a! severe! weakening! of! lower! facial! and! tongue! muscles,! impairments! in! biting,! chewing! and! licking! movements,! but were! still! able! to! produce! calls!, 1951. ,
! Interestingly,! the! laryngeal! representation! in the! human! primary! motor! cortex! is! more! caudally! located! in! the precentral! gyrus! relative! to! NHPs!, 1949. ,
, (area! 6)! into! the! primary! motor! cortex! (area! 4)! across primate! evolution,! which! provided! increased! fine! motor! control over! both! articulatory! and! laryngeal! muscles! that! is! necessary! for human! verbalization! (Simonyan,! 2014).! Lesions! of! the! orofacial motor! region! in! humans! leads! to! the! loss! of! voluntary! control over! orofacial! muscles! and! speech! production,! while! monkeys! with comparable! lesions! can! still! vocalize! voluntarily, This! suggests! a! shift! of! neocortical! laryngeal! control! from! premotor! regions!, 1982.
, areas The! orofacial! motor! region! which! lies! in! the! lower! part! of! the precentral! gyrus! comprises! the! primary! motor! cortex! (area! 4)! and two! premotor! areas:! ventrocaudal! area! 6! (6VC)! and! ventrorostral area! 6! (6VR)! (see! Petrides,! 2014).! Anterior! to! these! motor/premotor areas! lies! a! transitional! prefrontal! cytoarchitectonic! area! 44! and! a fully! prefrontal! area! 45! that! are! often! referred! to! as! Broca's! area, !, 1999.
, , 1994.
, ! we! use! the! label! 'PMv'! to! refer! to! both! areas 6VC! and! 6VR;! and! 'vlPFC'! to! refer! to! transitional! area! 44! (Fig.! 2a, ! the! current! review
! immediately! caudal! to! this! premotor region! and! extending! into! the! anterior! bank! of! the! central! sulcus,! is! the! orofacial! and! laryngeal! representation! of! the! primary motor! cortex! (area! 4).! The! transitional! vlPFC! is! situated! in! the! pars opercularis,! immediately! rostral! to! the! precentral! gyrus! followed further! rostrally! by! ventrolateral! prefrontal! area! 45! that! occupies the! pars! triangularis! (Amunts! et! al.,! 1999;! Petrides, ! the! human! brain,! the! PMv! occupies! most! of! the! ventral! surface of! the! precentral! gyrus! and, 1994. ,
, !, 2010.
! In! addition! to! the direct! projections! to! the! motor! nuclei! controlling! tongue,! jaw! and lip! movements,! the! human! PMv! connects! directly! to! the! nucleus ambiguus,! the! laryngeal! motor! nuclei! (Kuypers,! 1958b).! In! contrast, the! monkey! PMv! is! indirectly! linked! to! the! nucleus! ambiguus! via the! reticular! formation! in! the! brainstem! (Jürgens,! 2002).! Consistent! with! this! observation,! stimulation! of! the! PMv! evoked! orofacial movements! in! both! humans! and! monkeys! but! vocalizations!, !, 1949. ,
, , 2005.
! Located! between! higher-order! cognitive! multimodal! and! premotor! regions,! Petrides! (2005)! suggested that! the! transitional! vlPFC,! namely! area! 44,! is! well-positioned! to exert! high-level! control! over! orofacial! and! vocal! actions! and! to mediate! prefrontal! cortical! influences! on! such! motor! acts. Both! the! PMv! and! transitional! vlPFC! are! associated! with! voluntary! vocal! control! in! NHPs.! As! mentioned,! these! regions! project directly! to! the! orofacial! motor! nuclei! (Kuypers,! 1958a)! and! have been! associated! with! orofacial! movements! but! not! call! productions! upon! electrical! stimulations!, ! such! as! the! adjacent! area! 45! and! the mid-dorsolateral! area! 9/46v.! This! transitional! vlPFC! also! receives somatosensory! inputs! from! the! parietal! operculum,! insula! and! inferior! parietal! lobule! and! is! also! connected! to! premotor! regions! such as! the! PMv,! the! cingulate! (area! 23! and! 24)! and! supplementary motor! areas!, 1974. ,
! This! suggested! that PMv! in! NHPs! is! involved,! but! not! critical! for! conditioned! vocal! productions. The! volitional! control! of! vocalizations! in! humans! is! also! associated! with! the! PMv! and! the! vlPFC.! As! in! the! caudally! adjacent orofacial! motor! area,! these! regions! elicit! lip,! jaw! and! tongue! movements! and! vocalizations! upon! electrical! stimulation!, !, 1974. ,
! motor/premotor! PMv-evoked! speech arrests,! patients! reported! having! the! word! or! number! in! mind! but being! unable! to! vocalize! them! due! to! a! loss! of! control! over! articulatory! muscles.! With! vlPFC-evoked! speech! arrest,! patients! rather felt! they! had! muscular! control! but! were! disrupted! in! their! thoughts and! lost! track! of! the! content! to! be! vocalized.! Breshears! et! al.! (2015) further! showed! that! the! peak! probabilities! of! evoking! lips,! jaw, tongue! and! throat! movements! were! situated! in! the! PMv,! whereas the! peak! probability! of! evoking! speech! arrest! was! located! in! the vlPFC.! Long! et! al.! (2016)! demonstrated! that! focal! cortical! cooling in! awake! patients! of! the! PMv! impaired! the! articulatory! quality but! not! the! temporal! structure! of! verbalizations! (i.e.,! the! duration and! temporal! spacing! of! speech! elements),! whereas! the! opposite pattern! was! observed! with! vlPFC! deactivations.! Taken! together, rect! responses.! This! finding! indicates! that! the! MCC! is! involved! in the! processing! and! discrimination! of! outcomes! relevant! for! behavioral! adaptation.! Adopting! the! same! experimental! paradigm,! Amiez et! al.! (2012b,! 2013)! obtained! the! same! set! of! findings! in! an! fMRI study! with! human! subjects:! increased! BOLD! signal, ! However,! language interference! was! evoked! more! frequently! and! extensively! from stimulation! in! vlPFC! compared! to! stimulation! of! the! orofacial! motor region.! Penfield! and! Rasmussen! (1949)! reported! that! after! precentral! gyrus! stimulation,! i, 1949. ,
, ! the! MCC! is! also! involved! in! motivating! speech production! since! patients! with! MCC! lesions! exhibit! a! reduced! motivation! to! produce! speech! spontaneously.! As! discussed! earlier,! the MCC! is! critical! for! the! volitional! production! of! calls! in! NHPs! since! it constitutes! a! crucial! link! between! the! cognitive! brain! regions! and the! PAG.! Human! verbalizations, ! the! context! of! vocal! production,! the! MCC! could! be! involved In! humans
, ! In front! of! the! precentral! gyrus! (area! 4! and! caudal! area! 6)! that! is directly! involved! in! the! motor! control! and! execution! of! sensorimotor! associations,! the! caudal! frontal! region! (rostral! area! 6! and area! 8)! is! critical! for! the! conditional! rule-based! selection! between competing! alternative! responses.! Importantly,! particular! parts! of the! caudal! frontal! region! appear! to! be! involved! in! different! types! of these! conditional! if-then! selections! between! competing! responses based! on! the! motor! output! region! they! have! access! to! (Petrides, 2005).! For! instance,! area! 8,! which! is! linked! to! oculomotor! and! visuospatial! processing! regions,! was! shown! to! be! critical! for! selecting between! alternative! visual! stimuli! in! the! spatial! environment! based on! learned! if-then! rules,! while! the! dorsal! part! of! rostral! area! 6, which! is! linked! to! the! hand! motor! region! and! the! parietal! areas that! represent! the! hand! position! in! space,! was! critical! for! selecting between! different! motor! actions! based! on! pre-learned! if-then! conditional! rules.! By! the! same! reasoning,! the! ventral! caudal! prefrontal region! (i.e.! the! transitional! vlPFC),! adjacent! to! the! part! of! the! precentral! gyrus! that! is! strongly! associated! with! orofacial/vocal! motor control! (i.e.! PMv),! could! play! an! important! role! in! the! conditional selection! of! orofacial/vocal! responses! (Petrides, ! a! series! of! experiments! involving! selective! lesions! in! the! monkey,! Petrides! (2005)! demonstrated! the! existence! of! a! rostral-caudal functional! gradient! within! the! lateral! prefrontal! cortex! (Fig.! 2), 2006.
! via! intracortical recordings,! demonstrated! that,! as! human! subjects! adjusted! their vocal! productions! in! response! to! acoustic! perturbations,! the! PMv reflected! compensatory! activity! changes! that! were! correlated! with both! the! activity! associated! with! auditory! processing! as! well! as the! magnitude! of! vocal! pitch! adjustment.! Functional! neuroimaging investigations! also! revealed! that! human! area! 44! is! activated! for! both the! processing! of! articulatory/phonological! information! and! production! of! verbalizations!, !, 1999. ,
! the! primate! VLF! region! appears! to! be! involved! in! the higher-order! control! of! orofacial/vocal! responses! based! on! incoming! sensory! information.! This! functional! contribution! is! congruent with! the! VLF! region's! strong! anatomical! connections! with! temporoparietal! regions! that! provide! access! to! multisensory! inputs! and! the motor! regions! that! provide! control! over! orofacial/vocal! movements, 2006. ,
! found! mirror! neurons! in! the! anterior! part! of! area! 6VR that! responded! specifically! to! the! performance! and! observation of! orofacial! actions,! such! as! feeding! and! communicative! movements! (i.e.! lip-smacking).! These! orofacial! mirror! neurons! could contribute! to! the! imitative! learning! of! orofacial! actions! by! forming an! internal! representation! of! observed! orofacial! actions! via!, Within! area! 6VR! (often! referred! to! as! F5! in! macaque! monkeys) of! the! PMv! (Fig.! 2c), 1996. ,
, ! With! reduced! direct! laryngeal! access! in! the! primary! orofacial! motor! area,! the! PMv! in! NHPs! might! be! less! disposed! to! generate orofacial-laryngeal! motor! representations! of! heard! vocalizations
This! could! explain! why! vocal! imitative! abilities! are! limited! in! NHPs, !, 2014. ,
! In! contrast, with! direct! access! to! both! orofacial! and! laryngeal! motor! output, the! human! PMv! can! 'mirror'! heard! vocalizations! via! the! orofaciallaryngeal! motor! circuitry.! Congruent! with! this! account, 2015. ,
, ! In! this! section,! we! propose! the! existence! of! a! primate VLF-DMF! network! and! its! generic! contributions! to! the! cognitive control! of! vocalizations! in! both! human! and! NHPs.! Anatomically,! the primate! VLF-DMF! regions! are! interconnected! (Fig.! 1):! The! anterior MCC! (the! part! associated! with! vocalizations)! has! strong!, We! have! argued! that! a! comparable! set! of! primate! VLF-DMF regions! may! contribute! to! voluntary! vocal! control! in! both! NHPs and! humans, 1993.
! 1993).! Interestingly,! within this! primate! VLF-DMF! network, ! which! show! strong! interconnections! and! are! involved! in! vocal! control! in! both! the! human! and! NHP brain! (Fig.! 1), 1994. ,
! could! be! involved! in! linking! vocal/orofacial responses! to! different! contexts! via! conditional! if-then! rules.! With access! to! both! multimodal! contextual! information! and! vocalorofacial! motor! outputs,! the! vlPFC! is! well-placed! to! establish context-orofacial/vocal! response! conditional! operations! that! are then! executed! via! the! PMv! (Petrides,! 2006).! In! both! humans! and monkeys, regions! appear! to! perform! generic functions! in! NHPs! and! humans.! Primate! VLF! regions,! including the! vlPFC! and! PMv ,
, ! The! observation! that! pre-SMA! lesions! increased! the! latencies! of! voluntary! call! and! speech! initiations! in! both! monkeys! and NBR-2676
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! the! MCC region! in! both! monkeys! and! humans! is! associated! with! the! evaluation! of! action! outcomes! to! adapt! future! behaviors.! In! the! context of! vocalizations,! it! is! important! for! motivating! the! initiation! of! new vocalizations! or! adjustments! to! current! vocal! behaviors! based! on the! outcomes! of! previous! actions.! Indeed,! MCC! lesions! result! in reduced! volitional! vocalizations! in! both! NHPs! and! humans.! With strong! links! to! emotional! processing! regions! and! the! PAG,! the! MCC also! contributes! to! affective! vocalizations! in! both! humans! and! monkeys. Based! on! the! above! discussion,! we! propose! the! following! relative roles! of! the! primate! VLF-DMF! regions! that! are! generic! to! cognitive! control! of! vocalizations! in! both! humans! and! NHPs:! Whereas VLF! areas! contribute! to! the! conditional! selection! of! orofacial/vocal responses! based! on! sensorimotor! associations,! the! MCC! is! involved in! the! evaluation! of! orofacial/vocal! action! outcomes! to! drive! future orofacial/vocal! responses.! The! pre-SMA! is! generally! involved! in! the initiation! and! adjustments! of! orofacial/vocal! responses, ! strongly! suggests! that! the! primate! pre-SMA! generally! contributes! to! the! volitional! initiation! of! vocalizations.! Finally, 2016. ,
! One! possible! way! to test! our! proposed! functions! of! the! VLF! and! DMF! regions! in! parallel! would! be! via! a! conditional! associative! learning! protocol! (see Amiez! et! al.,! 2012a)! that! involves! both! the! learning! and! the! production! of! conditional! associations! between! arbitrary! sensory, ! these! regions! might! constitute! the! core! network! for! the primate! cognitive! control! of! vocalizations ,
DMF! network! relative! to! NHPs.! Firstly, as! suggested! by! Simonyan! (2014),! the! emergence! of! a! laryngeal! representation! in! the! human! orofacial! motor! area! afforded! increased fine-motor! control! over! oro-laryngeal! movements.! As! such,! areas! in the! human! VLF! which! are! directly! connected! to! the! orofacial! motor area,! would! be! able! to! construct! and! exercise! conditional! sensoryvocal! motor! associations! not! only! at! the! level! of! well-formed orofacial/vocal! responses!, ! How! has! the! VLF-DMF! network! changed! during! primate evolution? In! NHPs,! the! MCC! serves! as! the! crucial! link! between! the! PAG! and the! other! DMF! and! VLF! cognitive! brain! regions,! and! is! thus! critical for! the! production! of! affective! vocalizations! in! NHPs.! The! MCC! is also! likely! involved! in! the! limited! adjustments! of! affective! calls! (i.e. amplitude, 2013. ,
, ! apart! from! the! core! primate VLF,! pre-SMA! and! MCC! regions,! the! cognitive! control! of! human vocalizations! additionally! recruits! the! SMA.! As! discussed! earlier, the! SMA! is! associated! with! the! sequencing! of! motor! actions! and could! be! vital! in! the! sequencing! of! vocal/orofacial! movements! into novel! vocal! patterns! via! the! human! orofacial! motor! area,! which! has direct! access! to! laryngeal! and! orofacial! control.! These! novel! vocal patterns! could! then! be! conditionally! linked! with! contextual! representations! in! the! VLF,! adjusted! based! on! vocal! behavioral! feedback analyses! in! the! MCC,! and!, This! could! account! for! the! increased! capacity! of! humans! to! generate and! modify! flexibly! their! vocal! patterns! based! on! contextual! rules or! vocal! behavioral! feedback.! Secondly
, ! etc.)! that! involve! complex! combinations! of! oro-laryngeal! movements! as! well! as! the! acoustic! variations of! these! elements! (e.g.! pitch,! amplitude,! timbre,! etc.).! These! speech units,! along! with! their! acoustic! variations,! could! then! be! selectively produced! in! different! contexts! based! on! first-order! conditional! sensorimotor! rules! (i.e.! if! stimulus! A,! then! speech! pattern! X)! in! the! VLF (Petrides,! 2006).! Beyond! the! VLF,! rostral! lateral! frontal! regions! (i.e. areas! 45,! 9,! 46! and! 10)! might! be! implicated! in! higher-order! monitoring! of! selections! of! speech! actions! (Petrides, The! increased! fine-motor! control! over! oro-laryngeal! musculature! and! the! recruitment! of! the! SMA! for! oro-laryngeal! movement sequencing! have! enabled! the! generation! of! speech! elements! (e.g. phonemes,! words,! phrases, 2009.
! ventrolateral! prefrontal! area! 47/12,! and! frontal! pole! area! 10)! may select! these! context-sensorimotor! rules! based! on! episodic! representations,! such! as! past! experiences! and! internal! goals.! Along! with this! rostral-caudal! functional! gradient,! we! have! shown! earlier! that, within! the! VLF! region,! the! vlPFC! and! PMv! are! associated! with! higher and! lower! aspects! of! vocal! sensorimotor! control,! respectively.! Furthermore,! regions! rostral! to! the! VLF! have! consistently! been! found! to be! involved! in! higher! aspects! of! speech! production! (Bourguignon, ! Thus,! in! the! lateral! frontal! cortex,! the! most caudal! premotor! region! (area! 6)! represent! basic! sensorimotor! associations,! middle! regions! (areas! 44, vol.45, 2003. ,
, As! in! the! lateral! frontal! regions,! a! rostro-caudal! organization! has also! been! proposed! for! the! medial! frontal! cortex! (Domenech! and Koechlin, !, 2015.
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