{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T04:35:28Z","timestamp":1791434128809,"version":"4.3.3"},"reference-count":62,"publisher":"Wiley","issue":"2","license":[{"start":{"date-parts":[[2026,2,5]],"date-time":"2026-02-05T00:00:00Z","timestamp":1770249600000},"content-version":"vor","delay-in-days":4,"URL":"http:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0\/"},{"start":{"date-parts":[[2026,2,1]],"date-time":"2026-02-01T00:00:00Z","timestamp":1769904000000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/doi.wiley.com\/10.1002\/tdm_license_1.1"}],"funder":[{"DOI":"10.13039\/100019713","name":"VA Pittsburgh Healthcare System","doi-asserted-by":"publisher","award":["#1I01RX003511\u201001A2"],"award-info":[{"award-number":["#1I01RX003511\u201001A2"]}],"id":[{"id":"10.13039\/100019713","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["onlinelibrary.wiley.com"],"crossmark-restriction":true},"short-container-title":["Human Brain Mapping"],"published-print":{"date-parts":[[2026,2,1]]},"abstract":"<jats:title>ABSTRACT<\/jats:title>\n                  <jats:p>Non\u2010primary motor areas, including dorsal premotor cortex (PMd), ventral premotor cortex (PMv), and posterior parietal cortex (PPC), contribute to movement planning, but how these regions differentially shape kinematic features of goal\u2010directed movements, and how this specialization is associated with functional connectivity within the frontoparietal network, remains of interest, particularly in relation to recovery after stroke. We used functional magnetic resonance imaging (fMRI), transcranial magnetic stimulation (TMS), and kinematic assessments to explore how these areas influence reaching performance in neurologically intact adults. Participants performed a goal\u2010directed planar reaching task using the KINARM exoskeleton robot. Brief TMS pulse trains were initiated before movement onset to perturb cortical activity at subthreshold and suprathreshold intensities targeting bilateral PMd, PMv, and dorsomedial superior parietal lobule (SPL) within PPC. Resting\u2010state fMRI quantified functional connectivity among these regions to assess whether connectivity explains stimulation\u2010induced kinematic changes. Relative to the control target within the postcentral sulcus (PCS), subthreshold stimulation of contralateral PMd and PMv reduced reach efficiency and smoothness, while suprathreshold stimulation of contralateral PPC increased deviation error and reduced smoothness. Among ipsilateral targets, PMd showed consistent TMS\u2010induced effects, and was the only target where resting\u2010state connectivity predicted behavioral response. Stronger interhemispheric connectivity in the primary motor cortex and weaker interhemispheric PPC connectivity were associated with greater reductions in straightness and smoothness during subthreshold ipsilateral PMd stimulation. We found that perturbation of premotor and parietal targets led to distinct kinematic effects that varied by site, intensity, and laterality, with premotor stimulation showing the most consistent disruptions at subthreshold intensity and bilateral effects, whereas parietal effects were observed primarily for contralateral stimulation at suprathreshold intensity, and differences in network organization explain variability in behavioral response. Identifying contributions of cortical areas and connectivity patterns may help personalize interventions after stroke.<\/jats:p>\n                  <jats:p>\n                    <jats:bold>Trial Registration:<\/jats:bold>\n                    This study was registered at\n                    <jats:ext-link xmlns:xlink=\"http:\/\/www.w3.org\/1999\/xlink\" xlink:href=\"http:\/\/clinicaltrials.gov\">ClinicalTrials.gov<\/jats:ext-link>\n                    under ID NCT04286516\n                  <\/jats:p>","DOI":"10.1002\/hbm.70465","type":"journal-article","created":{"date-parts":[[2026,2,5]],"date-time":"2026-02-05T13:21:22Z","timestamp":1770297682000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Mapping the Causal Roles of Non\u2010Primary Motor Areas in Human Reach Planning and Execution"],"prefix":"10.1002","volume":"47","author":[{"ORCID":"https:\/\/orcid.org\/0009-0006-6986-3578","authenticated-orcid":false,"given":"Golnaz","family":"Haddadshargh","sequence":"first","affiliation":[{"name":"Rehab Neural Engineering Labs University of Pittsburgh  Pittsburgh Pennsylvania USA"},{"name":"Department of Bioengineering University of Pittsburgh  Pittsburgh Pennsylvania USA"},{"name":"Center for Neural Basis of Cognition, University of Pittsburgh and Carnegie Mellon University  Pittsburgh Pennsylvania USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Roberto M.","family":"de Freitas","sequence":"additional","affiliation":[{"name":"Rehab Neural Engineering Labs University of Pittsburgh  Pittsburgh Pennsylvania USA"},{"name":"Department of Neurological Surgery University of Pittsburgh  Pittsburgh Pennsylvania USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jennifer","family":"Mak","sequence":"additional","affiliation":[{"name":"Rehab Neural Engineering Labs University of Pittsburgh  Pittsburgh Pennsylvania USA"},{"name":"Department of Bioengineering University of Pittsburgh  Pittsburgh Pennsylvania USA"},{"name":"Center for Neural Basis of Cognition, University of Pittsburgh and Carnegie Mellon University  Pittsburgh Pennsylvania USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Amy","family":"Boos","sequence":"additional","affiliation":[{"name":"Rehab Neural Engineering Labs University of Pittsburgh  Pittsburgh Pennsylvania USA"},{"name":"Department of Neurology University of Pittsburgh  Pittsburgh Pennsylvania USA"},{"name":"TECH\u2010GRECC &amp; HERL, VA Pittsburgh Healthcare System  Pittsburgh Pennsylvania USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaoqi","family":"Fang","sequence":"additional","affiliation":[{"name":"Rehab Neural Engineering Labs University of Pittsburgh  Pittsburgh Pennsylvania USA"},{"name":"Department of Neurology University of Pittsburgh  Pittsburgh Pennsylvania USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jennifer L.","family":"Collinger","sequence":"additional","affiliation":[{"name":"Rehab Neural Engineering Labs University of Pittsburgh  Pittsburgh Pennsylvania USA"},{"name":"Department of Bioengineering University of Pittsburgh  Pittsburgh Pennsylvania USA"},{"name":"Center for Neural Basis of Cognition, University of Pittsburgh and Carnegie Mellon University  Pittsburgh Pennsylvania USA"},{"name":"TECH\u2010GRECC &amp; HERL, VA Pittsburgh Healthcare System  Pittsburgh Pennsylvania USA"},{"name":"Department of Physical Medicine and Rehabilitation University of Pittsburgh  Pittsburgh Pennsylvania USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gina","family":"McKernan","sequence":"additional","affiliation":[{"name":"TECH\u2010GRECC &amp; HERL, VA Pittsburgh Healthcare System  Pittsburgh Pennsylvania USA"},{"name":"Department of Physical Medicine and Rehabilitation University of Pittsburgh  Pittsburgh Pennsylvania USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Liang","family":"Zhan","sequence":"additional","affiliation":[{"name":"Department of Bioengineering University of Pittsburgh  Pittsburgh Pennsylvania USA"},{"name":"Department of Electrical and Computer Engineering University of Pittsburgh  Pittsburgh Pennsylvania USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fang","family":"Liu","sequence":"additional","affiliation":[{"name":"Rehab Neural Engineering Labs University of Pittsburgh  Pittsburgh Pennsylvania USA"},{"name":"TECH\u2010GRECC &amp; HERL, VA Pittsburgh Healthcare System  Pittsburgh Pennsylvania USA"},{"name":"Department of Physical Medicine and Rehabilitation University of Pittsburgh  Pittsburgh Pennsylvania USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"George F.","family":"Wittenberg","sequence":"additional","affiliation":[{"name":"Rehab Neural Engineering Labs University of Pittsburgh  Pittsburgh Pennsylvania USA"},{"name":"Department of Bioengineering University of Pittsburgh  Pittsburgh Pennsylvania USA"},{"name":"Center for Neural Basis of Cognition, University of Pittsburgh and Carnegie Mellon University  Pittsburgh Pennsylvania USA"},{"name":"Department of Neurology University of Pittsburgh  Pittsburgh Pennsylvania USA"},{"name":"TECH\u2010GRECC &amp; HERL, VA Pittsburgh Healthcare System  Pittsburgh Pennsylvania USA"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,2,5]]},"reference":[{"key":"e_1_2_12_2_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.neuro.25.112701.142922"},{"key":"e_1_2_12_3_1","volume-title":"TMS Motor Threshold Assessment Tool (MTAT 2.0)","author":"Awiszus F.","year":"2011"},{"key":"e_1_2_12_4_1","doi-asserted-by":"publisher","DOI":"10.1109\/TBME.2011.2179545"},{"key":"e_1_2_12_5_1","doi-asserted-by":"publisher","DOI":"10.1111\/ejn.12742"},{"key":"e_1_2_12_6_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroimage.2007.04.042"},{"key":"e_1_2_12_7_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.2517\u20106161.1995.tb02031.x"},{"key":"e_1_2_12_8_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00011.2012"},{"key":"e_1_2_12_9_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.00456.2006"},{"key":"e_1_2_12_10_1","doi-asserted-by":"publisher","DOI":"10.1152\/jn.91194.2008"},{"key":"e_1_2_12_11_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00429\u2010023\u201002735\u20107"},{"key":"e_1_2_12_12_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1609-22.2022"},{"key":"e_1_2_12_13_1","doi-asserted-by":"publisher","DOI":"10.1093\/cercor\/bhaa224"},{"issue":"1","key":"e_1_2_12_14_1","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1523\/JNEUROSCI.0659-24.2024","article-title":"Role of the Medial Posterior Parietal Cortex in Orchestrating Attention and Reaching","volume":"45","author":"Breveglieri R.","year":"2025","journal-title":"Journal of Neuroscience"},{"key":"e_1_2_12_15_1","doi-asserted-by":"publisher","DOI":"10.1007\/s002210050902"},{"key":"e_1_2_12_16_1","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0004621"},{"key":"e_1_2_12_17_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2009.11.005"},{"key":"e_1_2_12_18_1","doi-asserted-by":"publisher","DOI":"10.1177\/1073858405284255"},{"key":"e_1_2_12_19_1","doi-asserted-by":"publisher","DOI":"10.1177\/1545968309356091"},{"key":"e_1_2_12_20_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00221\u2010003\u20101587\u20101"},{"key":"e_1_2_12_21_1","doi-asserted-by":"publisher","DOI":"10.1006\/cbmr.1996.0014"},{"key":"e_1_2_12_22_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.3386-05.2006"},{"key":"e_1_2_12_23_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.cub.2009.11.063"},{"key":"e_1_2_12_24_1","doi-asserted-by":"publisher","DOI":"10.1038\/9219"},{"key":"e_1_2_12_25_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.4299-12.2012"},{"issue":"3","key":"e_1_2_12_26_1","doi-asserted-by":"crossref","first-page":"342","DOI":"10.1037\/0033-2909.127.3.342","article-title":"A Century Later: Woodworth's (1899) Two\u2010Component Model of Goal\u2010Directed Aiming","volume":"127","author":"Elliott D.","year":"2001","journal-title":"Psychological Bulletin"},{"key":"e_1_2_12_27_1","doi-asserted-by":"publisher","DOI":"10.1177\/1073858410375468"},{"key":"e_1_2_12_28_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroimage.2012.01.021"},{"key":"e_1_2_12_29_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1058-11.2011"},{"issue":"26","key":"e_1_2_12_30_1","doi-asserted-by":"crossref","first-page":"9599","DOI":"10.1523\/JNEUROSCI.0080-11.2011","article-title":"Decoding Action Intentions From Preparatory Brain Activity in Human Parieto\u2010Frontal Networks","volume":"31","author":"Gallivan J. P.","year":"2011","journal-title":"Journal of Neuroscience"},{"issue":"1","key":"e_1_2_12_31_1","doi-asserted-by":"crossref","DOI":"10.1038\/s41598-020-67162-8","article-title":"Individual Differences in Local Functional Brain Connectivity Affect TMS Effects on Behavior","volume":"10","author":"Gie\u00dfing C.","year":"2020","journal-title":"Scientific Reports"},{"key":"e_1_2_12_32_1","doi-asserted-by":"publisher","DOI":"10.1162\/0898929052880066"},{"key":"e_1_2_12_33_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.3923-07.2007"},{"key":"e_1_2_12_34_1","doi-asserted-by":"crossref","first-page":"76","DOI":"10.3389\/fnins.2020.00076","article-title":"Explaining Individual Differences in Motor Behavior by Intrinsic Functional Connectivity and Corticospinal Excitability","volume":"14","author":"Herszage J.","year":"2020","journal-title":"Frontiers in Neuroscience"},{"key":"e_1_2_12_35_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.conb.2007.02.003"},{"key":"e_1_2_12_36_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuron.2012.10.030"},{"key":"e_1_2_12_37_1","doi-asserted-by":"publisher","DOI":"10.1006\/nimg.2001.0856"},{"key":"e_1_2_12_38_1","doi-asserted-by":"publisher","DOI":"10.1006\/nimg.2002.1132"},{"key":"e_1_2_12_39_1","doi-asserted-by":"publisher","DOI":"10.1016\/S1361-8415(01)00036-6"},{"key":"e_1_2_12_40_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1158-06.2006"},{"key":"e_1_2_12_41_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroimage.2010.07.034"},{"issue":"3","key":"e_1_2_12_42_1","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1523\/ENEURO.0008-15.2015","article-title":"Flexible Reference Frames for Grasp Planning in Human Parietofrontal Cortex","volume":"2","author":"Leon\u00e9 F. T.","year":"2015","journal-title":"ENeuro"},{"key":"e_1_2_12_43_1","doi-asserted-by":"publisher","DOI":"10.1007\/PL00005662"},{"key":"e_1_2_12_44_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroimage.2022.119221"},{"key":"e_1_2_12_45_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.2356-15.2016"},{"key":"e_1_2_12_46_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1460\u20109568.2007.05795.x"},{"key":"e_1_2_12_47_1","doi-asserted-by":"publisher","DOI":"10.1098\/rstb.2005.1652"},{"key":"e_1_2_12_48_1","volume-title":"Statistical Parametric Mapping: The Analysis of Functional Brain Images","author":"Penny W. D.","year":"2011"},{"key":"e_1_2_12_49_1","doi-asserted-by":"publisher","DOI":"10.1016\/S0959-4388(01)00266-5"},{"issue":"9","key":"e_1_2_12_50_1","doi-asserted-by":"crossref","first-page":"977","DOI":"10.1093\/cercor\/13.9.977","article-title":"Activation of the Supplementary Motor Area (SMA) During Performance of Visually Guided Movements","volume":"13","author":"Picard N.","year":"2003","journal-title":"Cerebral Cortex"},{"key":"e_1_2_12_51_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroimage.2019.116092"},{"key":"e_1_2_12_52_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41591-022-02202-6"},{"key":"e_1_2_12_53_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroimage.2013.08.048"},{"key":"e_1_2_12_54_1","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1641-06.2006"},{"key":"e_1_2_12_55_1","doi-asserted-by":"publisher","DOI":"10.1162\/089892901750363244"},{"key":"e_1_2_12_56_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.neuroimage.2011.09.016"},{"key":"e_1_2_12_57_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.conb.2006.06.016"},{"issue":"2","key":"e_1_2_12_58_1","doi-asserted-by":"crossref","first-page":"119","DOI":"10.1016\/S0165-0270(99)00053-9","article-title":"Apparatus for Measuring and Perturbing Shoulder and Elbow Joint Positions and Torques During Reaching","volume":"89","author":"Scott S. H.","year":"1999","journal-title":"Journal of Neuroscience Methods"},{"key":"e_1_2_12_59_1","doi-asserted-by":"publisher","DOI":"10.2165\/00019053\u2010200321030\u201000004"},{"key":"e_1_2_12_60_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00221\u2010021\u201006228\u2010z"},{"key":"e_1_2_12_61_1","doi-asserted-by":"publisher","DOI":"10.1007\/s00221\u2010012\u20103158\u20109"},{"key":"e_1_2_12_62_1","doi-asserted-by":"publisher","DOI":"10.1089\/brain.2012.0073"},{"key":"e_1_2_12_63_1","doi-asserted-by":"publisher","DOI":"10.1038\/nmeth.1635"}],"container-title":["Human Brain Mapping"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/hbm.70465","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/full-xml\/10.1002\/hbm.70465","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/pdf\/10.1002\/hbm.70465","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,10]],"date-time":"2026-02-10T07:52:39Z","timestamp":1770709959000},"score":1,"resource":{"primary":{"URL":"https:\/\/onlinelibrary.wiley.com\/doi\/10.1002\/hbm.70465"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,1]]},"references-count":62,"journal-issue":{"issue":"2","published-print":{"date-parts":[[2026,2,1]]}},"alternative-id":["10.1002\/hbm.70465"],"URL":"https:\/\/doi.org\/10.1002\/hbm.70465","archive":["Portico"],"relation":{},"ISSN":["1065-9471","1097-0193"],"issn-type":[{"value":"1065-9471","type":"print"},{"value":"1097-0193","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,1]]},"assertion":[{"value":"2025-09-19","order":0,"name":"received","label":"Received","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-01-25","order":2,"name":"accepted","label":"Accepted","group":{"name":"publication_history","label":"Publication History"}},{"value":"2026-02-05","order":3,"name":"published","label":"Published","group":{"name":"publication_history","label":"Publication History"}}],"article-number":"e70465"}}