Resting state functional connectivity in early blind humans
Identifieur interne : 001F61 ( Pmc/Curation ); précédent : 001F60; suivant : 001F62Resting state functional connectivity in early blind humans
Auteurs : Harold Burton [États-Unis] ; Abraham Z. Snyder [États-Unis] ; Marcus E. Raichle [États-Unis]Source :
- Frontiers in Systems Neuroscience [ 1662-5137 ] ; 2014.
Abstract
Task-based neuroimaging studies in early blind humans (EB) have demonstrated heightened visual cortex responses to non-visual paradigms. Several prior functional connectivity studies in EB have shown altered connections consistent with these task-based results. But these studies generally did not consider behavioral adaptations to lifelong blindness typically observed in EB. Enhanced cognitive abilities shown in EB include greater serial recall and attention to memory. Here, we address the question of the extent to which brain intrinsic activity in EB reflects such adaptations. We performed a resting-state functional magnetic resonance imaging study contrasting 14 EB with 14 age/gender matched normally sighted controls (NS). A principal finding was markedly greater functional connectivity in EB between visual cortex and regions typically associated with memory and cognitive control of attention. In contrast, correlations between visual cortex and non-deprived sensory cortices were significantly lower in EB. Thus, the available data, including that obtained in prior task-based and resting state fMRI studies, as well as the present results, indicate that visual cortex in EB becomes more heavily incorporated into functional systems instantiating episodic recall and attention to non-visual events. Moreover, EB appear to show a reduction in interactions between visual and non-deprived sensory cortices, possibly reflecting suppression of inter-sensory distracting activity.
Url:
DOI: 10.3389/fnsys.2014.00051
PubMed: 24778608
PubMed Central: 3985019
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<front><div type="abstract" xml:lang="en"><p>Task-based neuroimaging studies in early blind humans (EB) have demonstrated heightened visual cortex responses to non-visual paradigms. Several prior functional connectivity studies in EB have shown altered connections consistent with these task-based results. But these studies generally did not consider behavioral adaptations to lifelong blindness typically observed in EB. Enhanced cognitive abilities shown in EB include greater serial recall and attention to memory. Here, we address the question of the extent to which brain intrinsic activity in EB reflects such adaptations. We performed a resting-state functional magnetic resonance imaging study contrasting 14 EB with 14 age/gender matched normally sighted controls (NS). A principal finding was markedly greater functional connectivity in EB between visual cortex and regions typically associated with memory and cognitive control of attention. In contrast, correlations between visual cortex and non-deprived sensory cortices were significantly lower in EB. Thus, the available data, including that obtained in prior task-based and resting state fMRI studies, as well as the present results, indicate that visual cortex in EB becomes more heavily incorporated into functional systems instantiating episodic recall and attention to non-visual events. Moreover, EB appear to show a reduction in interactions between visual and non-deprived sensory cortices, possibly reflecting suppression of inter-sensory distracting activity.</p>
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<pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-ta">Front Syst Neurosci</journal-id>
<journal-id journal-id-type="iso-abbrev">Front Syst Neurosci</journal-id>
<journal-id journal-id-type="publisher-id">Front. Syst. Neurosci.</journal-id>
<journal-title-group><journal-title>Frontiers in Systems Neuroscience</journal-title>
</journal-title-group>
<issn pub-type="epub">1662-5137</issn>
<publisher><publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta><article-id pub-id-type="pmid">24778608</article-id>
<article-id pub-id-type="pmc">3985019</article-id>
<article-id pub-id-type="doi">10.3389/fnsys.2014.00051</article-id>
<article-categories><subj-group subj-group-type="heading"><subject>Neuroscience</subject>
<subj-group><subject>Original Research Article</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group><article-title>Resting state functional connectivity in early blind humans</article-title>
</title-group>
<contrib-group><contrib contrib-type="author"><name><surname>Burton</surname>
<given-names>Harold</given-names>
</name>
<xref ref-type="aff" rid="aff1"><sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup>
</xref>
<xref ref-type="author-notes" rid="fn001"><sup>*</sup>
</xref>
<uri xlink:type="simple" xlink:href="http://community.frontiersin.org/people/u/87580"></uri>
</contrib>
<contrib contrib-type="author"><name><surname>Snyder</surname>
<given-names>Abraham Z.</given-names>
</name>
<xref ref-type="aff" rid="aff2"><sup>2</sup>
</xref>
<uri xlink:type="simple" xlink:href="http://community.frontiersin.org/people/u/11137"></uri>
</contrib>
<contrib contrib-type="author"><name><surname>Raichle</surname>
<given-names>Marcus E.</given-names>
</name>
<xref ref-type="aff" rid="aff2"><sup>2</sup>
</xref>
<uri xlink:type="simple" xlink:href="http://community.frontiersin.org/people/u/11138"></uri>
</contrib>
</contrib-group>
<aff id="aff1"><sup>1</sup>
<institution>Department of Anatomy and Neurobiology, Washington University School of Medicine</institution>
<country>St. Louis, MO, USA</country>
</aff>
<aff id="aff2"><sup>2</sup>
<institution>Department of Radiology, Washington University School of Medicine</institution>
<country>St. Louis, MO, USA</country>
</aff>
<author-notes><fn fn-type="edited-by"><p>Edited by: Gustavo Deco, Universitat Pompeu Fabra, Spain</p>
</fn>
<fn fn-type="edited-by"><p>Reviewed by: Joana R. B. Cabral, Universitat Pompeu Fabra, Spain; James W. Lewis, West Virginia University, USA</p>
</fn>
<corresp id="fn001">*Correspondence: Harold Burton, Department of Anatomy and Neurobiology, Washington University School of Medicine, Campus Box 8108, 660 South Euclid Avenue, St. Louis, MO 63110, USA e-mail: <email xlink:type="simple">harold@pcg.wustl.edu</email>
</corresp>
<fn fn-type="other" id="fn002"><p>This article was submitted to the journal Frontiers in Systems Neuroscience.</p>
</fn>
</author-notes>
<pub-date pub-type="epub"><day>07</day>
<month>4</month>
<year>2014</year>
</pub-date>
<pub-date pub-type="collection"><year>2014</year>
</pub-date>
<volume>8</volume>
<elocation-id>51</elocation-id>
<history><date date-type="received"><day>17</day>
<month>10</month>
<year>2013</year>
</date>
<date date-type="accepted"><day>19</day>
<month>3</month>
<year>2014</year>
</date>
</history>
<permissions><copyright-statement>Copyright © 2014 Burton, Snyder and Raichle.</copyright-statement>
<copyright-year>2014</copyright-year>
<license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by/3.0/"><license-p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</license-p>
</license>
</permissions>
<abstract><p>Task-based neuroimaging studies in early blind humans (EB) have demonstrated heightened visual cortex responses to non-visual paradigms. Several prior functional connectivity studies in EB have shown altered connections consistent with these task-based results. But these studies generally did not consider behavioral adaptations to lifelong blindness typically observed in EB. Enhanced cognitive abilities shown in EB include greater serial recall and attention to memory. Here, we address the question of the extent to which brain intrinsic activity in EB reflects such adaptations. We performed a resting-state functional magnetic resonance imaging study contrasting 14 EB with 14 age/gender matched normally sighted controls (NS). A principal finding was markedly greater functional connectivity in EB between visual cortex and regions typically associated with memory and cognitive control of attention. In contrast, correlations between visual cortex and non-deprived sensory cortices were significantly lower in EB. Thus, the available data, including that obtained in prior task-based and resting state fMRI studies, as well as the present results, indicate that visual cortex in EB becomes more heavily incorporated into functional systems instantiating episodic recall and attention to non-visual events. Moreover, EB appear to show a reduction in interactions between visual and non-deprived sensory cortices, possibly reflecting suppression of inter-sensory distracting activity.</p>
</abstract>
<kwd-group><kwd>blindness</kwd>
<kwd>human</kwd>
<kwd>fMRI</kwd>
<kwd>functional connectivity</kwd>
<kwd>visual cortex</kwd>
</kwd-group>
<counts><fig-count count="7"></fig-count>
<table-count count="2"></table-count>
<equation-count count="0"></equation-count>
<ref-count count="147"></ref-count>
<page-count count="13"></page-count>
<word-count count="10692"></word-count>
</counts>
</article-meta>
</front>
<body><sec sec-type="introduction" id="s1"><title>Introduction</title>
<p>Abundant evidence indicates that “visual cortex”<xref ref-type="fn" rid="fn0001"><sup>1</sup>
</xref>
performs non-visual functions in persons who were born blind or who lost sight in infancy, (e.g., early blind, EB) (Pascual-Leone et al., <xref rid="B102" ref-type="bibr">2005</xref>
; Burton and Mclaren, <xref rid="B19" ref-type="bibr">2008</xref>
; Merabet and Pascual-Leone, <xref rid="B92" ref-type="bibr">2010</xref>
; Kupers and Ptito, <xref rid="B79" ref-type="bibr">2011</xref>
; Ricciardi and Pietrini, <xref rid="B114" ref-type="bibr">2011</xref>
). For example, the observation that EB can acquire Braille alexia following occipital strokes suggests that visual cortex contributes to language cognition. Such strokes do not interfere with general somatosensory awareness nor do they cause aphasia in the context of spoken language (Hamilton et al., <xref rid="B64" ref-type="bibr">2000</xref>
; Maeda and Yasuda, <xref rid="B89" ref-type="bibr">2003</xref>
). Similarly, repetitive TMS applied to visual cortex, disrupts Braille letter identification without eliminating awareness of touching a Braille field (Cohen et al., <xref rid="B31" ref-type="bibr">1997</xref>
).</p>
<p>Despite results such as those described above, the prevailing view is that visual cortex function in EB can be understood in terms of “cross-modal” processing of auditory and haptic stimuli (reviewed in Rauschecker, <xref rid="B109" ref-type="bibr">1995</xref>
; Kujala et al., <xref rid="B77" ref-type="bibr">2000</xref>
; Pascual-Leone and Hamilton, <xref rid="B103" ref-type="bibr">2001</xref>
; Burton, <xref rid="B16" ref-type="bibr">2003</xref>
; Noppeney et al., <xref rid="B96" ref-type="bibr">2003</xref>
; Burton and Mclaren, <xref rid="B19" ref-type="bibr">2008</xref>
; Merabet and Pascual-Leone, <xref rid="B92" ref-type="bibr">2010</xref>
; Sathian and Stilla, <xref rid="B124" ref-type="bibr">2010</xref>
; Dormal and Collignon, <xref rid="B39" ref-type="bibr">2011</xref>
; Frasnelli et al., <xref rid="B50" ref-type="bibr">2011</xref>
; Kupers and Ptito, <xref rid="B79" ref-type="bibr">2011</xref>
; Ricciardi and Pietrini, <xref rid="B114" ref-type="bibr">2011</xref>
; Voss and Zatorre, <xref rid="B139" ref-type="bibr">2012</xref>
). According to this view, visual cortex in EB contributes to a characteristic “perceptual advantage… in processing information of the intact modalities” (p. 118 in Kujala et al., <xref rid="B77" ref-type="bibr">2000</xref>
). Example enhanced abilities include better spatial localization of sounds (Lessard et al., <xref rid="B83" ref-type="bibr">1998</xref>
), sharper pitch discriminations (Gougoux et al., <xref rid="B56" ref-type="bibr">2004</xref>
), and finer perception of tactile grating orientation (Goldreich and Kanics, <xref rid="B55" ref-type="bibr">2003</xref>
). These observations suggest the possibility that visual cortex in EB becomes specialized for the analysis of non-visual modalities, possibly via enhanced functional connectivity with somatosensory and auditory cortices. That these enhanced abilities are attributable to reorganized visual cortex is suggested by the observation that TMS of visual cortex disrupts auditory spatial localization in the EB (Collignon et al., <xref rid="B33" ref-type="bibr">2007</xref>
). However, the “cross-modal” account implies that visual cortex in EB should exhibit enhanced functional connectivity with auditory cortex, and yet this prediction is not supported by prior resting-state functional connectivity experiments (Liu et al., <xref rid="B86" ref-type="bibr">2007</xref>
; Wang et al., <xref rid="B140" ref-type="bibr">2008</xref>
; Yu et al., <xref rid="B147" ref-type="bibr">2008</xref>
; Watkins et al., <xref rid="B141" ref-type="bibr">2012</xref>
). In fact, Liu and colleagues reported abnormally low resting state fMRI correlations between visual and somatomotor, auditory and multisensory cortices (Liu et al., <xref rid="B86" ref-type="bibr">2007</xref>
). The same group also reported generally reduced functional connectivity between primary visual cortex and the rest of the brain (Yu et al., <xref rid="B147" ref-type="bibr">2008</xref>
).</p>
<p>EB exhibit superior abilities in several cognitive domains not explainable in terms of “cross-modal” analysis of the sensorium. These perceptual advantages might be better explained in terms of superior working memory (Hull and Mason, <xref rid="B69" ref-type="bibr">1995</xref>
; Raz et al., <xref rid="B111" ref-type="bibr">2007</xref>
; Rokem and Ahissar, <xref rid="B120" ref-type="bibr">2009</xref>
; Swanson and Luxenberg, <xref rid="B131" ref-type="bibr">2009</xref>
; Withagen et al., <xref rid="B145" ref-type="bibr">2013</xref>
) and better “attentive processing of stimuli” (p. 118 in Kujala et al., <xref rid="B77" ref-type="bibr">2000</xref>
), whether tactile or auditory (Uhl et al., <xref rid="B133" ref-type="bibr">1991</xref>
; Kujala et al., <xref rid="B76" ref-type="bibr">1995a</xref>
,<xref rid="B78" ref-type="bibr">b</xref>
, <xref rid="B75" ref-type="bibr">1997</xref>
; Röder et al., <xref rid="B118" ref-type="bibr">1996</xref>
; Sadato et al., <xref rid="B122" ref-type="bibr">1996</xref>
; Gizewski et al., <xref rid="B53" ref-type="bibr">2003</xref>
). Thus, activated responses in visual cortex are greatest during discrimination tasks that require attention to the stimuli. In contrast, somatosensory stimulation during non-discrimination tasks elicits weaker or even non-significant responses (Sadato et al., <xref rid="B122" ref-type="bibr">1996</xref>
; Burton et al., <xref rid="B25" ref-type="bibr">2002a</xref>
; Gizewski et al., <xref rid="B53" ref-type="bibr">2003</xref>
). Better working memory in EB can be demonstrated with tasks involving verbal material, pitch discrimination, and auditory spatial localization (Park et al., <xref rid="B101" ref-type="bibr">2011</xref>
). EB also show greater recall of rehearsed words (Amedi et al., <xref rid="B3" ref-type="bibr">2003</xref>
; Azulay et al., <xref rid="B5" ref-type="bibr">2009</xref>
) and heightened recollection of studied word lists (Raz et al., <xref rid="B111" ref-type="bibr">2007</xref>
). Similarly, EB retained >60% recognition of studied words after protracted delays (Raz et al., <xref rid="B110" ref-type="bibr">2005</xref>
) and exhibited greater recall accuracy than sighted individuals (Pasqualotto et al., <xref rid="B104" ref-type="bibr">2013</xref>
).</p>
<p>These observations suggest that sensory deprivation-induced brain reorganization (Pascual-Leone and Hamilton, <xref rid="B103" ref-type="bibr">2001</xref>
; Bavelier and Neville, <xref rid="B6" ref-type="bibr">2002</xref>
) in EB leads to a performance advantage not only in discrimination tasks involving intact sensory modalities, but also an advantage in multiple cognitive domains, e.g., enhanced recollection of studied verbal material. A general supposition is that these heightened abilities reflect re-programming of visual cortex for “metamodal” purposes (Pascual-Leone and Hamilton, <xref rid="B103" ref-type="bibr">2001</xref>
; Pascual-Leone et al., <xref rid="B102" ref-type="bibr">2005</xref>
). Supporting this view are multiple neuroimaging experiments in EB demonstrating vigorous responses throughout visual cortex to language tasks involving semantic memory (Burton et al., <xref rid="B25" ref-type="bibr">2002a</xref>
, <xref rid="B17" ref-type="bibr">2003</xref>
; Amedi et al., <xref rid="B3" ref-type="bibr">2003</xref>
; Noppeney et al., <xref rid="B96" ref-type="bibr">2003</xref>
; Raz et al., <xref rid="B110" ref-type="bibr">2005</xref>
; Bedny et al., <xref rid="B7" ref-type="bibr">2011</xref>
). Additionally, EB show evoked activity to other complex cognitive processing normally associated with visual experience such as recognition memories of human action sounds (Lewis et al., <xref rid="B85" ref-type="bibr">2011</xref>
) and even mirroring human actions (Ricciardi et al., <xref rid="B113" ref-type="bibr">2009</xref>
). It is possible that as blind engage normally “visual cortex” when performing such tasks they utilize a memory echo of sensory experiences as previously noted in sighted (Wheeler et al., <xref rid="B144" ref-type="bibr">2000</xref>
).</p>
<p>The issue is whether the functionality represented in visual cortex of EB primarily relates to attention or memory or language, or, indeed, any particular sensory or cognitive domain. To address this question, we performed a resting-state functional connectivity study contrasting 14 EB vs. 14 age/sex matched normally sighted (NS) participants. The underlying hypothesis was that reorganized resting-state networks reflect functionality represented in visual cortex of EB. We assume that the nature of this functionality can be determined by identifying which networks share correlated intrinsic activity with visual cortex in EB. Statistical significance of EB vs. NS functional connectivity differences was assessed using threshold-free cluster enhancement (TFCE) analysis (Smith and Nichols, <xref rid="B129" ref-type="bibr">2009</xref>
; Hill et al., <xref rid="B67" ref-type="bibr">2010</xref>
).</p>
</sec>
<sec sec-type="methods" id="s2"><title>Methods</title>
<sec><title>Participants</title>
<p>The EB group included 14 individuals having a mean age of 46.3, <italic>SD</italic>
± 11.4 (6 female) (Table <xref ref-type="table" rid="T1">1</xref>
). Blindness resulted from retinal pathology in all cases, most commonly retinopathy of prematurity. None could see patterns or objects; 6 claimed slight sensitivity to brightness and 8 had no awareness of light. All EB were Braille literate and most were fluent readers. The NS control group included 14 individuals having a mean age of 48.1, <italic>SD</italic>
± 11.6 (8 female). All participants provided written informed consent in accordance with the Declaration of Helsinki and guidelines approved by the Washington University School of Medicine Human Studies Institutional Review Board.</p>
<table-wrap id="T1" position="float"><label>Table 1</label>
<caption><p><bold>Early blind and sighted characteristics</bold>
.</p>
</caption>
<table frame="hsides" rules="groups"><thead><tr><th align="left" rowspan="1" colspan="1"><bold>Early blind</bold>
</th>
<th align="center" rowspan="1" colspan="1"><bold>Age</bold>
</th>
<th align="center" rowspan="1" colspan="1"><bold>Gender</bold>
</th>
<th align="center" rowspan="1" colspan="1"><bold>Preferred hand</bold>
</th>
<th align="center" rowspan="1" colspan="1"><bold>Braille reading</bold>
</th>
<th align="center" rowspan="1" colspan="1"><bold>wpm</bold>
</th>
<th align="center" rowspan="1" colspan="1"><bold>Onset age</bold>
</th>
<th align="center" rowspan="1" colspan="1"><bold>Light sense</bold>
</th>
<th align="center" rowspan="1" colspan="1"><bold>Cause of blindness<xref ref-type="table-fn" rid="TN1a"><sup>a</sup>
</xref>
</bold>
</th>
</tr>
</thead>
<tbody><tr><td align="left" rowspan="1" colspan="1">EB1</td>
<td align="center" rowspan="1" colspan="1">41</td>
<td align="center" rowspan="1" colspan="1">F</td>
<td align="center" rowspan="1" colspan="1">LH</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">76</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="center" rowspan="1" colspan="1">Some</td>
<td align="center" rowspan="1" colspan="1">ONH</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">EB2</td>
<td align="center" rowspan="1" colspan="1">40</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">BH</td>
<td align="center" rowspan="1" colspan="1">88.7</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="center" rowspan="1" colspan="1">None</td>
<td align="center" rowspan="1" colspan="1">ROP</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">EB3</td>
<td align="center" rowspan="1" colspan="1">53</td>
<td align="center" rowspan="1" colspan="1">F</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">BH</td>
<td align="center" rowspan="1" colspan="1">227.9</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="center" rowspan="1" colspan="1">Some</td>
<td align="center" rowspan="1" colspan="1">ROP</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">EB4</td>
<td align="center" rowspan="1" colspan="1">52</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">113.3</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="center" rowspan="1" colspan="1">None</td>
<td align="center" rowspan="1" colspan="1">ROP</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">EB5</td>
<td align="center" rowspan="1" colspan="1">31</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">LH</td>
<td align="center" rowspan="1" colspan="1">58.7</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="center" rowspan="1" colspan="1">Some</td>
<td align="center" rowspan="1" colspan="1">LCA</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">EB6</td>
<td align="center" rowspan="1" colspan="1">58</td>
<td align="center" rowspan="1" colspan="1">F</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">145.4</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="center" rowspan="1" colspan="1">None</td>
<td align="center" rowspan="1" colspan="1">ROP</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">EB7</td>
<td align="center" rowspan="1" colspan="1">52</td>
<td align="center" rowspan="1" colspan="1">F</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">BH</td>
<td align="center" rowspan="1" colspan="1">185.8</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="center" rowspan="1" colspan="1">None</td>
<td align="center" rowspan="1" colspan="1">ROP</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">EB8</td>
<td align="center" rowspan="1" colspan="1">56</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">152</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="center" rowspan="1" colspan="1">None</td>
<td align="center" rowspan="1" colspan="1">ROP</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">EB9</td>
<td align="center" rowspan="1" colspan="1">62</td>
<td align="center" rowspan="1" colspan="1">F</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">BH</td>
<td align="center" rowspan="1" colspan="1">137</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="center" rowspan="1" colspan="1">Some</td>
<td align="center" rowspan="1" colspan="1">GRP</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">EB10</td>
<td align="center" rowspan="1" colspan="1">54</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">BH</td>
<td align="center" rowspan="1" colspan="1">60.2</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="center" rowspan="1" colspan="1">Some</td>
<td align="center" rowspan="1" colspan="1">ROP</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">EB11</td>
<td align="center" rowspan="1" colspan="1">32</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">BH</td>
<td align="center" rowspan="1" colspan="1">52.7</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="center" rowspan="1" colspan="1">Some</td>
<td align="center" rowspan="1" colspan="1">ROP</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">EB12</td>
<td align="center" rowspan="1" colspan="1">30</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">LH</td>
<td align="center" rowspan="1" colspan="1">60.2</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="center" rowspan="1" colspan="1">None</td>
<td align="center" rowspan="1" colspan="1">ROP</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">EB13</td>
<td align="center" rowspan="1" colspan="1">55</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">n/a</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="center" rowspan="1" colspan="1">None</td>
<td align="center" rowspan="1" colspan="1">Cataract</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">EB14</td>
<td align="center" rowspan="1" colspan="1">32</td>
<td align="center" rowspan="1" colspan="1">F</td>
<td align="center" rowspan="1" colspan="1">RH</td>
<td align="center" rowspan="1" colspan="1">LH</td>
<td align="center" rowspan="1" colspan="1">103.6</td>
<td align="center" rowspan="1" colspan="1">0</td>
<td align="center" rowspan="1" colspan="1">None</td>
<td align="center" rowspan="1" colspan="1">RB</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">Avg</td>
<td align="center" rowspan="1" colspan="1">46.3</td>
<td align="center" rowspan="1" colspan="1">6F/8M</td>
<td align="center" rowspan="1" colspan="1">1L/13R</td>
<td align="center" rowspan="1" colspan="1">3L/5R/6B</td>
<td align="center" rowspan="1" colspan="1">112.4</td>
<td rowspan="1" colspan="1"></td>
<td align="center" rowspan="1" colspan="1">8n/4s</td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">SD</td>
<td align="center" rowspan="1" colspan="1">11.4</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td align="center" rowspan="1" colspan="1">54.7</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"><bold>Sighted</bold>
</td>
<td align="center" rowspan="1" colspan="1"><bold>Age</bold>
</td>
<td align="center" rowspan="1" colspan="1"><bold>Gender</bold>
</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">NS1</td>
<td align="center" rowspan="1" colspan="1">46</td>
<td align="center" rowspan="1" colspan="1">F</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">NS2</td>
<td align="center" rowspan="1" colspan="1">42</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">NS3</td>
<td align="center" rowspan="1" colspan="1">54</td>
<td align="center" rowspan="1" colspan="1">F</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">NS4</td>
<td align="center" rowspan="1" colspan="1">46</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">NS5</td>
<td align="center" rowspan="1" colspan="1">30</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">NS6</td>
<td align="center" rowspan="1" colspan="1">57</td>
<td align="center" rowspan="1" colspan="1">F</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">NS7</td>
<td align="center" rowspan="1" colspan="1">51</td>
<td align="center" rowspan="1" colspan="1">F</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">NS8</td>
<td align="center" rowspan="1" colspan="1">55</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">NS9</td>
<td align="center" rowspan="1" colspan="1">63</td>
<td align="center" rowspan="1" colspan="1">F</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">NS10</td>
<td align="center" rowspan="1" colspan="1">50</td>
<td align="center" rowspan="1" colspan="1">F</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">NS11</td>
<td align="center" rowspan="1" colspan="1">27</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">NS12</td>
<td align="center" rowspan="1" colspan="1">68</td>
<td align="center" rowspan="1" colspan="1">M</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">NS13</td>
<td align="center" rowspan="1" colspan="1">48</td>
<td align="center" rowspan="1" colspan="1">F</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">NS14</td>
<td align="center" rowspan="1" colspan="1">36</td>
<td align="center" rowspan="1" colspan="1">F</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">Avg</td>
<td align="center" rowspan="1" colspan="1">48.1</td>
<td align="center" rowspan="1" colspan="1">8F/6M</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">SD</td>
<td align="center" rowspan="1" colspan="1">11.6</td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
<td rowspan="1" colspan="1"></td>
</tr>
</tbody>
</table>
<table-wrap-foot><fn id="TN1a"><label>a</label>
<p><italic>Abbreviations: ONH, optic nerve hypoplasia; LCA, Leber congenital amaurosis; GRP, genetic retinitis pigmentosa; RB, retinoblastoma; ROP, retinopathy of prematurity</italic>
.</p>
</fn>
</table-wrap-foot>
</table-wrap>
</sec>
<sec><title>Image acquisition</title>
<p>Magnetic resonance imaging was performed with a 3 T Trio scanner (Siemens, Erlangen Germany) equipped with a standard 12 element RF head coil. Resting-state functional connectivity was acquired using a gradient recalled echo-planar sequence (EPI) sensitive to blood oxygen dependent-level (BOLD) contrast [repetition time (<italic>TR</italic>
) = 2200 ms, echo time (<italic>TE</italic>
) = 27 ms, flip angle = 90°, 4-mm cubic voxels]. Whole brain coverage was obtained with 36 interleaved EPI slices (no gap) aligned parallel to the anterior-posterior commissural line. Each fMRI run included 164 volumes (6 min duration). Imaging was in a darkened room and with a blindfold covering the eyes of all participants. NS kept their eyes closed. Four runs were acquired in the EB and three in NS. We checked with the participants between runs to verify that they remained awake.</p>
<p>Structural images included a sagittal T1-weighted magnetization-prepared rapid gradient echo (MP-RAGE) scan [<italic>TR</italic>
= 2100 ms; <italic>TE</italic>
= 3.93 ms; flip angle = 7°; inversion time (<italic>TI</italic>
) = 1000 ms; 1 × 1 × 1.25 mm voxels, 176 slices]. A T2-weighted structural image (<italic>TR</italic>
= 8430 ms, <italic>TE</italic>
= 98 ms, 1.33 × 1.33 × 3 mm voxels, 36 slices) was also acquired to serve as a registration intermediary between the EPI and MP-RAGE (Ojemann et al., <xref rid="B99" ref-type="bibr">1997</xref>
).</p>
</sec>
<sec><title>Image processing</title>
<p>Pre-processing was performed using locally developed software (Shulman et al., <xref rid="B128" ref-type="bibr">2010</xref>
). Asynchronous slice timing was corrected using sinc interpolation. Correction for systematic odd/even slice-dependent intensity differences was as previously described (Hacker et al., <xref rid="B61" ref-type="bibr">2012</xref>
). Head motion within and across runs was corrected using rigid body realignment. There were no systematic group differences in the prevalence of head motion (<italic>EB</italic>
: 0.25 ± 0.14 rms mm vs. <italic>NS</italic>
0.26 ± 0.14 rms mm; two sample <italic>t-test</italic>
, <italic>P</italic>
= 0.73) (Hacker et al., <xref rid="B61" ref-type="bibr">2012</xref>
). Transformation to Talairach atlas space (Talairach and Tournoux, <xref rid="B132" ref-type="bibr">1988</xref>
) was achieved using 12 parameter affine transforms obtained from sequential co-registration between an average from the first frames of each EPI run, T2-weighted and T1-weighted structural images to an atlas template (Shulman et al., <xref rid="B128" ref-type="bibr">2010</xref>
). To eliminate the possibility of group-dependent atlas transformation bias, the atlas representative template was generated from MP-RAGE structural images obtained in EB and NS populations, as previously described (Buckner et al., <xref rid="B13" ref-type="bibr">2004</xref>
). Presently reported atlas coordinates represent Talairach space as defined by the spatial normalization method (Lancaster et al., <xref rid="B80" ref-type="bibr">1995</xref>
).</p>
<p>In preparation for seed-based correlation mapping, the EPI were spatially smoothed (6 mm full width at half-maximum Gaussian kernel in each direction). Temporal low-pass filtering removed frequencies >0.1 Hz. Spurious variance was reduced using linear regression of 9 sources of nuisance waveforms and their associated temporal derivatives. These variables included the six parameters derived by head movement correction, signals in the ventricles and white matter, and a global<xref ref-type="fn" rid="fn0002"><sup>2</sup>
</xref>
whole-brain signal averaged over all voxels in a fixed region of atlas space (Gusnard and Raichle, <xref rid="B60" ref-type="bibr">2001</xref>
; Fox et al., <xref rid="B46" ref-type="bibr">2005</xref>
, <xref rid="B47" ref-type="bibr">2009</xref>
). The ventricle and white matter regressors were extracted from regions individually delineated in each participant using Analyze (Mayo Research Foundation, Rochester, MN) (Burton et al., <xref rid="B27" ref-type="bibr">2012c</xref>
).</p>
</sec>
<sec><title>Correlation computation</title>
<p>Computation of Pearson correlation coefficients was for 16.4 min of resting state fMRI data between paired seed regions for each participant (temporal correlations). Concatenation of three EPI runs provided the time series<xref ref-type="fn" rid="fn0003"><sup>3</sup>
</xref>
. Each run consisted of 164 frames minus the first 15 volumes to assure magnetization equalization and BOLD response adaptation to sequence noises at scan onsets. The signal extracted from each seed region was the average per time point across all voxels within ~1 cm<sup>3</sup>
spheres. A Fischer's r-to-z transformation {<italic>Z</italic>
(<italic>r</italic>
) = 0.5 ln [(1 + <italic>r</italic>
)/(1 − <italic>r</italic>
)]} improved the normality of the correlation coefficients (Jenkins and Watts, <xref rid="B71" ref-type="bibr">1968</xref>
). A two-sample, two-tailed <italic>t</italic>
-test of <italic>Z</italic>
(<italic>r</italic>
), with assumed unequal variances, assessed whether two seed regions were coincidently active.</p>
<p>Temporal correlation computations included all possible combinations of 62 seed regions defined <italic>a priori</italic>
from previous studies (Table <xref ref-type="table" rid="T2">2</xref>
). Seed regions were placed in sensory (vision, audition and touch), attention, cognitive control, and self-referential default mode networks (DMNs). Functional cortical connectivity map computations were restricted to seed regions in temporal correlation pairings with <italic>t</italic>
-test <italic>p</italic>
-values = 0.0005.</p>
<table-wrap id="T2" position="float"><label>Table 2</label>
<caption><p><bold>Talairach atlas coordinates for selected seed regions</bold>
.</p>
</caption>
<table frame="hsides" rules="groups"><thead><tr><th align="left" rowspan="1" colspan="1"><bold>Network</bold>
</th>
<th align="center" rowspan="1" colspan="1"><bold>ROI</bold>
</th>
<th align="left" rowspan="1" colspan="1"><bold>Seed region</bold>
</th>
<th align="center" rowspan="1" colspan="1"><bold><italic>X</italic>
, <italic>Y</italic>
, <italic>Z</italic>
</bold>
</th>
</tr>
</thead>
<tbody><tr><td align="left" rowspan="2" colspan="1">Dorsal attention (DAN)</td>
<td align="left" rowspan="1" colspan="1">1</td>
<td align="left" rowspan="1" colspan="1">L intraparietal sulcus (LIPS)</td>
<td align="left" rowspan="1" colspan="1">−23, −66, 46<xref ref-type="table-fn" rid="TN1"><sup>1</sup>
</xref>
</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">2</td>
<td align="left" rowspan="1" colspan="1">R intraparietal sulcus (RIPS)</td>
<td align="left" rowspan="1" colspan="1">26, −58, 52<xref ref-type="table-fn" rid="TN1"><sup>1</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">3</td>
<td align="left" rowspan="1" colspan="1">L ventral intraparietal sulcus (LVIPS)</td>
<td align="left" rowspan="1" colspan="1">−24, −69, 30<xref ref-type="table-fn" rid="TN2"><sup>2</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">4</td>
<td align="left" rowspan="1" colspan="1">R ventral intraparietal sulcus (RVIPS)</td>
<td align="left" rowspan="1" colspan="1">30, −80, 16<xref ref-type="table-fn" rid="TN2"><sup>2</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">5</td>
<td align="left" rowspan="1" colspan="1">L frontal eye fields (LFEF)</td>
<td align="left" rowspan="1" colspan="1">−25, −12, 49<xref ref-type="table-fn" rid="TN3"><sup>3</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">6</td>
<td align="left" rowspan="1" colspan="1">R frontal eye fields (RFEF)</td>
<td align="left" rowspan="1" colspan="1">25, −12, 50<xref ref-type="table-fn" rid="TN3"><sup>3</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">7</td>
<td align="left" rowspan="1" colspan="1">L superior parietal lobule (LSPL)</td>
<td align="left" rowspan="1" colspan="1">−27, −54, 53<xref ref-type="table-fn" rid="TN4"><sup>4</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">8</td>
<td align="left" rowspan="1" colspan="1">R superior parietal lobule (RSPL)</td>
<td align="left" rowspan="1" colspan="1">22, −58, 52<xref ref-type="table-fn" rid="TN4"><sup>4</sup>
</xref>
</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">Ventral attention (VAN)</td>
<td align="left" rowspan="1" colspan="1">9</td>
<td align="left" rowspan="1" colspan="1">L temporoparietal junction (LTPJ)</td>
<td align="left" rowspan="1" colspan="1">−54, −48, 37<xref ref-type="table-fn" rid="TN4"><sup>4</sup>
</xref>
</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">10</td>
<td align="left" rowspan="1" colspan="1">R temporoparietal junction (RTPJ)</td>
<td align="left" rowspan="1" colspan="1">49, −50, 28<xref ref-type="table-fn" rid="TN4"><sup>4</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">11</td>
<td align="left" rowspan="1" colspan="1">R superior temporal sulcus (RSTS)</td>
<td align="left" rowspan="1" colspan="1">55, −50, 11<xref ref-type="table-fn" rid="TN4"><sup>4</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">12</td>
<td align="left" rowspan="1" colspan="1">R middle frontal gyrus (RMFG)</td>
<td align="left" rowspan="1" colspan="1">39, 12, 34<xref ref-type="table-fn" rid="TN5"><sup>5</sup>
</xref>
</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">Control</td>
<td align="left" rowspan="1" colspan="1">13</td>
<td align="left" rowspan="1" colspan="1">L anterior prefrontal cortex (LAPFC)</td>
<td align="left" rowspan="1" colspan="1">−35, 51, 13<xref ref-type="table-fn" rid="TN3"><sup>3</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">14</td>
<td align="left" rowspan="1" colspan="1">R anterior prefrontal cortex (RAPFC)</td>
<td align="left" rowspan="1" colspan="1">32, 46, 16<xref ref-type="table-fn" rid="TN3"><sup>3</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">15</td>
<td align="left" rowspan="1" colspan="1">L dorsolateral prefrontal cortex (LDLPFC)</td>
<td align="left" rowspan="1" colspan="1">−49, 16, 35<xref ref-type="table-fn" rid="TN3"><sup>3</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">16</td>
<td align="left" rowspan="1" colspan="1">R dorsolateral prefrontal cortex (RDLPFC)</td>
<td align="left" rowspan="1" colspan="1">43, 9, 45<xref ref-type="table-fn" rid="TN3"><sup>3</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">17</td>
<td align="left" rowspan="1" colspan="1">Accessory cingulate cortex (ACC)</td>
<td align="left" rowspan="1" colspan="1">2, 26, 30<xref ref-type="table-fn" rid="TN3"><sup>3</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">18</td>
<td align="left" rowspan="1" colspan="1">Dorsal accessory cingulate cortex (DACC)</td>
<td align="left" rowspan="1" colspan="1">−9, 8, 40<xref ref-type="table-fn" rid="TN6"><sup>6</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">19</td>
<td align="left" rowspan="1" colspan="1">L anterior inferior parietal lobule (LAIPL)</td>
<td align="left" rowspan="1" colspan="1">−51, −50, 43<xref ref-type="table-fn" rid="TN3"><sup>3</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">20</td>
<td align="left" rowspan="1" colspan="1">R anterior inferior parietal lobule (RAIPL)</td>
<td align="left" rowspan="1" colspan="1">49, −48, 45<xref ref-type="table-fn" rid="TN3"><sup>3</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">21</td>
<td align="left" rowspan="1" colspan="1">L anterior insula (LAINS)</td>
<td align="left" rowspan="1" colspan="1">−30, 17, 2<xref ref-type="table-fn" rid="TN3"><sup>3</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">22</td>
<td align="left" rowspan="1" colspan="1">R anterior insula (RAINS)</td>
<td align="left" rowspan="1" colspan="1">29, 17, 3<xref ref-type="table-fn" rid="TN3"><sup>3</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">23</td>
<td align="left" rowspan="1" colspan="1">L inferior frontal gyrus (LIFG)</td>
<td align="left" rowspan="1" colspan="1">−41, 6, 9<xref ref-type="table-fn" rid="TN1"><sup>1</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">24</td>
<td align="left" rowspan="1" colspan="1">R inferior frontal gyrus (RIFG)</td>
<td align="left" rowspan="1" colspan="1">45, −3, 12<xref ref-type="table-fn" rid="TN1"><sup>1</sup>
</xref>
</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">Auditory</td>
<td align="left" rowspan="1" colspan="1">25</td>
<td align="left" rowspan="1" colspan="1">L TE1 (LTE1)</td>
<td align="left" rowspan="1" colspan="1">−41, −32, 9<xref ref-type="table-fn" rid="TN7"><sup>7</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">26</td>
<td align="left" rowspan="1" colspan="1">R TE1 (RTE1)</td>
<td align="left" rowspan="1" colspan="1">37, −29, 14<xref ref-type="table-fn" rid="TN7"><sup>7</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">27</td>
<td align="left" rowspan="1" colspan="1">L TE2 (LTE2)</td>
<td align="left" rowspan="1" colspan="1">−45, −35, 9<xref ref-type="table-fn" rid="TN7"><sup>7</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">28</td>
<td align="left" rowspan="1" colspan="1">L TE3 (LTE3)</td>
<td align="left" rowspan="1" colspan="1">−60, −22, −1<xref ref-type="table-fn" rid="TN7"><sup>7</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">29</td>
<td align="left" rowspan="1" colspan="1">R TE3 (RTE3)</td>
<td align="left" rowspan="1" colspan="1">63, −18, 5<xref ref-type="table-fn" rid="TN7"><sup>7</sup>
</xref>
</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">Vision</td>
<td align="left" rowspan="1" colspan="1">30</td>
<td align="left" rowspan="1" colspan="1">L primary visual (LV1)</td>
<td align="left" rowspan="1" colspan="1">−8, −81, 5<xref ref-type="table-fn" rid="TN8"><sup>8</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">31</td>
<td align="left" rowspan="1" colspan="1">R primary visual (RV1)</td>
<td align="left" rowspan="1" colspan="1">11, −81, 5<xref ref-type="table-fn" rid="TN8"><sup>8</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">32</td>
<td align="left" rowspan="1" colspan="1">L visual area 3A (LV3A)</td>
<td align="left" rowspan="1" colspan="1">−22, −85, 16<xref ref-type="table-fn" rid="TN8"><sup>8</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">33</td>
<td align="left" rowspan="1" colspan="1">R visual area 3A (RV3A)</td>
<td align="left" rowspan="1" colspan="1">18, −88, 18<xref ref-type="table-fn" rid="TN8"><sup>8</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">34</td>
<td align="left" rowspan="1" colspan="1">L visual areas V4-VP (LV4_VP)</td>
<td align="left" rowspan="1" colspan="1">−18, −70, −10<xref ref-type="table-fn" rid="TN8"><sup>8</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">35</td>
<td align="left" rowspan="1" colspan="1">R visual areas V4-VP (RV4_VP)</td>
<td align="left" rowspan="1" colspan="1">23, −80, −10<xref ref-type="table-fn" rid="TN8"><sup>8</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">36</td>
<td align="left" rowspan="1" colspan="1">L lateral occipital cortex (LLO)</td>
<td align="left" rowspan="1" colspan="1">−36, −83, 3<xref ref-type="table-fn" rid="TN8"><sup>8</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">37</td>
<td align="left" rowspan="1" colspan="1">R lateral occipital cortex (RLO)</td>
<td align="left" rowspan="1" colspan="1">38, −80, 3<xref ref-type="table-fn" rid="TN8"><sup>8</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">38</td>
<td align="left" rowspan="1" colspan="1">L middle temporal area (LMT)</td>
<td align="left" rowspan="1" colspan="1">−43, −69, −4<xref ref-type="table-fn" rid="TN3"><sup>3</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">39</td>
<td align="left" rowspan="1" colspan="1">R middle temporal area (RMT)</td>
<td align="left" rowspan="1" colspan="1">48, −70, −3<xref ref-type="table-fn" rid="TN3"><sup>3</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">40</td>
<td align="left" rowspan="1" colspan="1">L visual area 8 (LV8)</td>
<td align="left" rowspan="1" colspan="1">−27, −58, −15<xref ref-type="table-fn" rid="TN9"><sup>9</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">41</td>
<td align="left" rowspan="1" colspan="1">R visual area 8 (RV8)</td>
<td align="left" rowspan="1" colspan="1">32, 61, −15<xref ref-type="table-fn" rid="TN9"><sup>9</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">42</td>
<td align="left" rowspan="1" colspan="1">L parietal-occipital sulcal cortex (LPOCS)</td>
<td align="left" rowspan="1" colspan="1">−21, −68, 15<xref ref-type="table-fn" rid="TN10"><sup>10</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">43</td>
<td align="left" rowspan="1" colspan="1">R parietal-occipital sulcal cortex (RPOCS)</td>
<td align="left" rowspan="1" colspan="1">17, −57, 10<xref ref-type="table-fn" rid="TN10"><sup>10</sup>
</xref>
</td>
</tr>
<tr><td align="left" rowspan="1" colspan="1">Somatosensory</td>
<td align="left" rowspan="1" colspan="1">44</td>
<td align="left" rowspan="1" colspan="1">L primary somatosensory (LS1)</td>
<td align="left" rowspan="1" colspan="1">−56, −18, 37<xref ref-type="table-fn" rid="TN11"><sup>11</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">45</td>
<td align="left" rowspan="1" colspan="1">R primary somatosensory (RS1)</td>
<td align="left" rowspan="1" colspan="1">51, −18, 44<xref ref-type="table-fn" rid="TN11"><sup>11</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">46</td>
<td align="left" rowspan="1" colspan="1">L BA3-trunk (LBA3_Trunk)</td>
<td align="left" rowspan="1" colspan="1">−31, −32, 58<xref ref-type="table-fn" rid="TN11"><sup>11</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">47</td>
<td align="left" rowspan="1" colspan="1">R BA3-trunk (RBA3_Trunk)</td>
<td align="left" rowspan="1" colspan="1">31, −32, 58<xref ref-type="table-fn" rid="TN11"><sup>11</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">48</td>
<td align="left" rowspan="1" colspan="1">L BA3-foot (LBA3_Foot)</td>
<td align="left" rowspan="1" colspan="1">−5, −34, 62<xref ref-type="table-fn" rid="TN11"><sup>11</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">49</td>
<td align="left" rowspan="1" colspan="1">R BA3-foot (RBA3_Foot)</td>
<td align="left" rowspan="1" colspan="1">5, −34, 62<xref ref-type="table-fn" rid="TN11"><sup>11</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">50</td>
<td align="left" rowspan="1" colspan="1">L BA2 (LBA2)</td>
<td align="left" rowspan="1" colspan="1">−43, −27, 44<xref ref-type="table-fn" rid="TN11"><sup>11</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">51</td>
<td align="left" rowspan="1" colspan="1">R BA2 (RBA2)</td>
<td align="left" rowspan="1" colspan="1">43, −27, 44<xref ref-type="table-fn" rid="TN11"><sup>11</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">52</td>
<td align="left" rowspan="1" colspan="1">L second somatosensory (LS2)</td>
<td align="left" rowspan="1" colspan="1">−35, −27, 17<xref ref-type="table-fn" rid="TN12"><sup>12</sup>
</xref>
</td>
</tr>
<tr><td rowspan="1" colspan="1"></td>
<td align="left" rowspan="1" colspan="1">53</td>
<td align="left" rowspan="1" colspan="1">R second somatosensory (RS2)</td>
<td align="left" rowspan="1" colspan="1">48, −14, 16<xref ref-type="table-fn" rid="TN12"><sup>12</sup>
</xref>
</td>
</tr>
</tbody>
</table>
<table-wrap-foot><fn id="TN1"><label>1</label>
<p>Fox et al., <xref rid="B46" ref-type="bibr">2005</xref>
;</p>
</fn>
<fn id="TN2"><label>2</label>
<p>Georgieva et al., <xref rid="B52" ref-type="bibr">2008</xref>
;</p>
</fn>
<fn id="TN3"><label>3</label>
<p>Vincent et al., <xref rid="B137" ref-type="bibr">2008</xref>
;</p>
</fn>
<fn id="TN4"><label>4</label>
<p>Shulman et al., <xref rid="B127" ref-type="bibr">2009</xref>
;</p>
</fn>
<fn id="TN5"><label>5</label>
<p>He et al., <xref rid="B66" ref-type="bibr">2007</xref>
;</p>
</fn>
<fn id="TN6"><label>6</label>
<p>Dosenbach et al., <xref rid="B40" ref-type="bibr">2007</xref>
;</p>
</fn>
<fn id="TN7"><label>7</label>
<p>Burton et al., <xref rid="B18" ref-type="bibr">2012a</xref>
;</p>
</fn>
<fn id="TN8"><label>8</label>
<p>Burton et al., <xref rid="B20" ref-type="bibr">2006</xref>
;</p>
</fn>
<fn id="TN9"><label>9</label>
<p>Noppeney et al., <xref rid="B97" ref-type="bibr">2006</xref>
;</p>
</fn>
<fn id="TN10"><label>10</label>
<p>Kaas et al., <xref rid="B72" ref-type="bibr">2007</xref>
;</p>
</fn>
<fn id="TN11"><label>11</label>
<p>Burton et al., <xref rid="B23" ref-type="bibr">2004</xref>
;</p>
</fn>
<fn id="TN12"><label>12</label>
<p><italic>Burton et al., <xref rid="B24" ref-type="bibr">2008</xref>
</italic>
.</p>
</fn>
</table-wrap-foot>
</table-wrap>
</sec>
<sec><title>Functional connectivity maps</title>
<p>Functional connectivity maps for a selected seed involved computing correlations in each participant between the 16.4 min time series in each 2 mm<sup>3</sup>
brain volume (voxel) and the average across all voxels in a seed region (Fox et al., <xref rid="B46" ref-type="bibr">2005</xref>
, <xref rid="B44" ref-type="bibr">2006</xref>
; Vincent et al., <xref rid="B138" ref-type="bibr">2006</xref>
, <xref rid="B137" ref-type="bibr">2008</xref>
; Burton et al., <xref rid="B27" ref-type="bibr">2012c</xref>
). Volumetric <italic>Z</italic>
(<italic>r</italic>
) maps were registered to a standard, population-average, landmark-surface-based atlas (PALS-B12) (Van Essen, <xref rid="B134" ref-type="bibr">2005</xref>
). The registration process involved using SureFit software to create a surface aligned through the mid-cortical gray matter in each individual (Van Essen et al., <xref rid="B136" ref-type="bibr">2001</xref>
; Van Essen, <xref rid="B134" ref-type="bibr">2005</xref>
; Van Essen and Dierker, <xref rid="B135" ref-type="bibr">2007</xref>
). Next, six selected landmarks per hemisphere on participant surfaces and in the average PALS-B12 atlas guided an algorithm that deformed the distribution of nodes in the native mid-cortical surface to the PALS-B12 atlas spherical surface. The computed deformation matrices directed subsequent registration of volumetric <italic>Z</italic>
(<italic>r</italic>
) maps to PALS-B12 surface space by aligning the coordinates of voxels to corresponding nodes with nearest neighbor coordinates (Nordahl et al., <xref rid="B98" ref-type="bibr">2007</xref>
). Group level surface-based functional connectivity maps for each seed were a computed random effects student's <italic>t</italic>
-test (Holmes and Friston, <xref rid="B68" ref-type="bibr">1998</xref>
) of <italic>Z</italic>
(<italic>r</italic>
) scores at each node. Presently displayed results in PALS-B12 surface space represent two-tailed <italic>t</italic>
-test results with probabilities of 0.05–0.0001 (<italic>t</italic>
= ± 2.16 to 5.5, 13 df). The <italic>t</italic>
-statistic values correspond to NS minus EB <italic>Z</italic>
(<italic>r</italic>
) group differences. Positive correspond to positive NS and/or negative EB correlations; negative <italic>t</italic>
-statistic values correspond to negative NS and/or positive EB correlations.</p>
</sec>
<sec><title>Assessment of statistical significance</title>
<p>The surface <italic>t</italic>
-statistic maps were minimally geodesically smoothed to a FWHM ≤7.4 mm (Hagler et al., <xref rid="B63" ref-type="bibr">2006</xref>
). TFCE (Smith and Nichols, <xref rid="B129" ref-type="bibr">2009</xref>
), adapted for analysis of surface based <italic>t</italic>
-statistic maps (Hill et al., <xref rid="B67" ref-type="bibr">2010</xref>
), was used to determine significant contiguous node clusters (see TFCE Implementation: <ext-link ext-link-type="uri" xlink:href="http://brainvis.wustl.edu/wiki/index.php/Caret:Documentation:Statistics">http://brainvis.wustl.edu/wiki/index.php/Caret:Documentation:Statistics</ext-link>
). Threshold-extent criteria were determined from 5000 permutations of randomly combined participants. The weight given to signal intensity and extent value were fixed at <italic>H</italic>
= 2.0 and <italic>E</italic>
= 1.0. We report clusters significant at <italic>p</italic>
≤ 0.05.</p>
</sec>
</sec>
<sec sec-type="results" id="s3"><title>Results</title>
<sec><title>Temporal correlations assessed in ROI pairs</title>
<p>In the NS group, temporal correlations between 406 ROI pairs within and across sensory systems (auditory, visual, and somatosensory) were consistently positive. Of these, 397 (98%) pairs had <italic>Z</italic>
(<italic>r</italic>
) values in the range 0.01–1.01, mean 0.12 ± 0.01 s.e.m. (Figure <xref ref-type="fig" rid="F1">1A</xref>
, warm hues in right lower portion of matrix). Temporal correlations between 464 ROI pairs between sensory and fronto-parietal control (FPC) and ventral attention network (VAN) ROI were shifted toward negative values, with 361/464 (78%) pairs having <italic>Z</italic>
(<italic>r</italic>
) values in the range −0.01 to −0.12, mean −0.05 ± 0.001 s.e.m. (Figure <xref ref-type="fig" rid="F1">1A</xref>
, cold hues). These findings in NS replicate many previous resting state results (Biswal et al., <xref rid="B9" ref-type="bibr">1995</xref>
; Cordes et al., <xref rid="B38" ref-type="bibr">2000</xref>
; Lowe et al., <xref rid="B87" ref-type="bibr">2000</xref>
; Greicius et al., <xref rid="B58" ref-type="bibr">2003</xref>
, <xref rid="B59" ref-type="bibr">2004</xref>
; Fox et al., <xref rid="B46" ref-type="bibr">2005</xref>
, <xref rid="B47" ref-type="bibr">2009</xref>
; Fransson, <xref rid="B48" ref-type="bibr">2005</xref>
; Hampson et al., <xref rid="B65" ref-type="bibr">2006</xref>
; Vincent et al., <xref rid="B138" ref-type="bibr">2006</xref>
; Buckner and Vincent, <xref rid="B14" ref-type="bibr">2007</xref>
; Dosenbach et al., <xref rid="B40" ref-type="bibr">2007</xref>
; Fair et al., <xref rid="B42" ref-type="bibr">2007</xref>
; Fox and Raichle, <xref rid="B45" ref-type="bibr">2007</xref>
; Fransson et al., <xref rid="B49" ref-type="bibr">2007</xref>
; Vincent et al., <xref rid="B137" ref-type="bibr">2008</xref>
).</p>
<fig id="F1" position="float"><label>Figure 1</label>
<caption><p><bold>Temporal correlation coefficient matrices computed from spontaneous, resting-state activity recorded for >16 min timelines from 53 paired seed regions in sighted (A) and early blind (B) participants</bold>
. See Table <xref ref-type="table" rid="T2">2</xref>
for identification of ROI Numbers. Seed regions were from different networks: dorsal attention, DAN; ventral attention, VAN; cognitive control, Control; auditory system, Aud; visual system, Visual; somatosensory, Somato. Color coded scale for Pearson correlation coefficients [<italic>Z</italic>
(<italic>r</italic>
) values]. Insert histogram of mean and standard error of <italic>Z</italic>
(<italic>r</italic>
) values in each early blind (EB) and normally sighted (NS) across all seed pairings for sensory regions.</p>
</caption>
<graphic xlink:href="fnsys-08-00051-g0001"></graphic>
</fig>
<p>Temporal correlations in several functional systems significantly differed in the EB group (Figure <xref ref-type="fig" rid="F1">1B</xref>
). Two-sided <italic>t</italic>
-tests found significant EB vs. NS group differences in 25 pairs with <italic>p</italic>
< 0.0005, which satisfied a Bonferroni correction (0.05/25 = 0.002). Most commonly were visual ROIs paired with ROIs in other functional systems (Figure <xref ref-type="fig" rid="F2">2A</xref>
). Bar graphs in Figure <xref ref-type="fig" rid="F2">2B</xref>
illustrate significant NS vs. EB <italic>Z</italic>
(<italic>r</italic>
) differences in selected ROI pairs.</p>
<fig id="F2" position="float"><label>Figure 2</label>
<caption><p><bold>Matrix of significant group differences in temporal correlation coefficients (A)</bold>
. Seed pairs correspond to ROI Numbers listed in Table <xref ref-type="table" rid="T2">2</xref>
. Filled cells indicate <italic>p</italic>
-values of <italic>t</italic>
-tests. <bold>(B)</bold>
Bar graphs of mean and standard errors for temporal correlation coefficients in each group. ANOVAs of each bar graph had a significant group effect (<italic>p</italic>
< 0.001) in seed pairs for visual vs. somatosensory, auditory, or control networks.</p>
</caption>
<graphic xlink:href="fnsys-08-00051-g0002"></graphic>
</fig>
<p>Several features are evident in the results shown in Figures <xref ref-type="fig" rid="F1">1</xref>
, <xref ref-type="fig" rid="F2">2</xref>
. (i) Correlations were modestly lower within visual cortex of the EB group, more so for inter-hemispheric than intra-hemispheric correlations, leading to a checkerboard effect in Figure <xref ref-type="fig" rid="F1">1B</xref>
. The greatest reduction in inter-hemispheric correlations was in higher order visual areas as opposed to V1 (Figure <xref ref-type="fig" rid="F3">3</xref>
). The NS vs. EB difference in inter-hemispheric functional connectivity was significant in LO by Bonferroni corrected <italic>t</italic>
-test. (ii) Visual cortex correlations with auditory and somatosensory cortices were consistently lower in the EB group; these differences gave rise to some of the most significant group contrasts in the present dataset. (iii) Correlations between visual cortex and nodes of the FPC network were consistently shifted toward more positive values in the EB group; 8 of these differences were significant at <italic>p</italic>
< 0.0005 and 18 at <italic>p</italic>
< 0.005. (iv) ROI pairs involving the DMN showed no significant group differences. The DMN ROIs are not included in the connectivity mapping results.</p>
<fig id="F3" position="float"><label>Figure 3</label>
<caption><p><bold>Interhemispheric temporal correlations in homotopic seed regions in early blind (EB) and normally sighted (NS)</bold>
. Plotted data values are mean and standard error of interhemispheric <italic>Z</italic>
(<italic>r</italic>
) values. Abbrevations: IFG, inferior frontal gyrus; TE1, core region of primary auditory; V1, primary visual; V3A, visual area 3A; V4-VP, visual areas V4 and VP; LO, lateral occipital cortex; MT, middle-temporal area; V8, visual area 8; POCS, parietal occipital sulcal cortex.</p>
</caption>
<graphic xlink:href="fnsys-08-00051-g0003"></graphic>
</fig>
</sec>
<sec><title>Effects of early blindness assessed in seed-based correlation difference maps</title>
<p>Functional connectivity [i.e., <italic>Z</italic>
(<italic>r</italic>
) maps] were computed for all 25 ROIs yielding a group difference in temporal correlation exceeding a significance level of <italic>p</italic>
= 0.0005 (Figure <xref ref-type="fig" rid="F2">2A</xref>
). Visual cortex seeds in the NS group generated correlation maps characterized by bilateral symmetry, positive <italic>Z</italic>
(<italic>r</italic>
) values in somatomotor, auditory, and within visual cortex, and negative <italic>Z</italic>
(<italic>r</italic>
) values in dorsolateral prefrontal and supramarginal cortex (Figure <xref ref-type="fig" rid="F4">4</xref>
). Differences evident in the EB group included positive rather than negative <italic>Z</italic>
(<italic>r</italic>
) values in left dorsolateral prefrontal and negative rather than positive values in somatomotor and auditory cortex (Figure <xref ref-type="fig" rid="F4">4</xref>
). Within visual cortex, <italic>Z</italic>
(<italic>r</italic>
) maps for positive values were less extensive especially for higher tier visual ROI, e.g., LV3A and LV4-VP (Figures <xref ref-type="fig" rid="F4">4B,C</xref>
, bracketed region). NS vs. EB differences were topographically similar across seed locations in primary (LV1) (Figure <xref ref-type="fig" rid="F4">4A</xref>
).</p>
<fig id="F4" position="float"><label>Figure 4</label>
<caption><p><bold>Resting-state functional connectivity random effect <italic>Z</italic>
(<italic>r</italic>
) maps in normally sighted (NS) and early blind (EB) associated with selected sensory seed regions identified in Table <xref ref-type="table" rid="T2">2</xref>
</bold>
. In NS, all seed regions in visual cortex <bold>(A–D)</bold>
showed positively correlated connectivity with other sensory cortical regions. In EB, these same seed regions showed positive correlations only within visual cortex and negatively correlated connectivity with non-visual somatosensory and auditory regions. Seed regions in non-visual sensory cortex, e.g., auditory RTE1, <bold>(E)</bold>
similarly showed positive correlations with other sensory cortex in NS. In EB, connectivity correlations with occipital cortex were negative but positive elsewhere as in NS. An arrow points at a green sphere that marks the location of the seed region in each map. Orange-Yellow painted in cortex with positive correlations having <italic>p</italic>
-values of 0.05 to <0.0001. Dark to light blue painted in cortex with negative correlations having <italic>p</italic>
-values of 0.05 to <0.0001. Black borders surround significant contiguous clusters identified using the threshold-free cluster enhancement assay (see Figure <xref ref-type="fig" rid="F5">5</xref>
).</p>
</caption>
<graphic xlink:href="fnsys-08-00051-g0004"></graphic>
</fig>
<p>TFCE analysis of correlation maps obtained with visual cortex seeds revealed significant NS vs. EB clusters in extensive regions of somatomotor, auditory, and higher tier visual cortex as well as prefrontal and parietal regions. In greater detail, visual cortex correlations with dorsolateral frontal cortex (DLPFC), posterior intraparietal sulcal (pIPS) and left fusiform gyrus were significantly more positive in the EB group. Conversely, visual cortex correlations with somatomotor, auditory and higher tier visual (Figures <xref ref-type="fig" rid="F5">5B,C</xref>
bracketed region) cortex were significantly less positive in the EB group. Figures <xref ref-type="fig" rid="F4">4</xref>
, <xref ref-type="fig" rid="F5">5A–D</xref>
show results obtained with 4 representative visual cortex seeds. Similar results were obtained with other visual cortex seeds (RV1, RV3A, RV4-VP, RLO, LLO, LV8, RV8).</p>
<fig id="F5" position="float"><label>Figure 5</label>
<caption><p><bold>Contrast of NS compared to EB functional connectivity based on two-tailed <italic>t</italic>
-test for selected sensory seed regions (A–E) identified in Table <xref ref-type="table" rid="T2">2</xref>
</bold>
. The <italic>t</italic>
-statistic maps show positive <italic>t</italic>
-test values (orange to yellow) where NS had greater positive correlations and/or EB had greater negative correlations. The <italic>t</italic>
-statistic maps show negative <italic>t</italic>
-test values (dark to light blue) where NS had larger negative correlations and/or EB had larger positive correlations. Black borders surround significant contiguous clusters identified using the threshold-free cluster enhancement assay (see Methods). An arrow points at a green sphere that marks the location of the seed region in each map.</p>
</caption>
<graphic xlink:href="fnsys-08-00051-g0005"></graphic>
</fig>
<p>TFCE analysis of correlation maps obtained with auditory and somatosensory cortex seeds revealed significant NS vs. EB clusters throughout visual cortex. Correlations with visual cortex were significantly more positive in the NS compared to the EB group. Figures <xref ref-type="fig" rid="F4">4</xref>
, <xref ref-type="fig" rid="F5">5E</xref>
show results obtained with a representative auditory cortex seed, RTE1. Other auditory cortex seeds (LTE1, LTE2, LTE3, RTE3) and somatosensory cortex seeds (LS1, RS1, LBA3, RBA3, LBA2, RBA2, LS2, RS2) yielded similar results.</p>
<p>Figures <xref ref-type="fig" rid="F6">6</xref>
, <xref ref-type="fig" rid="F7">7A–C</xref>
present results for three seed ROIs in cognitive-control cortex. Each had significantly more positive visual cortex correlations in the EB as compared to the NS group. The latter effect was most prominent in extra-striate visual areas, i.e., areas V2 and V3/VP rather than V1. For the seed in left anterior intraparietal sulcus (LAIPL, Figure <xref ref-type="fig" rid="F6">6C</xref>
), significantly more positive correlations extended to areas in infero-temporal cortex, probably corresponding to the V4 and V8 areas. Additional regions of significant difference included portions of parietal cortex and posterior temporal cortex in a distribution corresponding to the posterior components of the dorsal attention network (DAN). TFCE analysis of correlation maps obtained with cognitive-control cortex seeds revealed significant NS vs. EB clusters throughout visual cortex (Figures <xref ref-type="fig" rid="F7">7A–C</xref>
).</p>
<fig id="F6" position="float"><label>Figure 6</label>
<caption><p><bold>Resting-state functional connectivity random effect <italic>Z</italic>
(<italic>r</italic>
) maps in normally sighted (NS) and early blind (EB) associated with selected seed regions (A–C) identified in Table <xref ref-type="table" rid="T2">2</xref>
</bold>
. A green sphere marks the location of the seed region in each map. See text for Figure <xref ref-type="fig" rid="F4">4</xref>
.</p>
</caption>
<graphic xlink:href="fnsys-08-00051-g0006"></graphic>
</fig>
<fig id="F7" position="float"><label>Figure 7</label>
<caption><p><bold>Contrast of NS compared to EB functional connectivity based on two-tailed <italic>t</italic>
-test for selected seed regions (A–C) identified in Table <xref ref-type="table" rid="T2">2</xref>
</bold>
. See text for Figure <xref ref-type="fig" rid="F5">5</xref>
.</p>
</caption>
<graphic xlink:href="fnsys-08-00051-g0007"></graphic>
</fig>
</sec>
</sec>
<sec sec-type="discussion" id="s4"><title>Discussion</title>
<sec><title>Overview</title>
<p>EB compared to NS had: (1) decreased functional connectivity between visual and somatosensory or auditory cortices; (2) inter-hemispheric visual cortex temporal correlations of lower magnitude in higher tier visual areas; and (3) increased functional connectivity between visual cortex and regions in frontal and parietal cortex associated with cognitive control.</p>
</sec>
<sec><title>Sensory network functional connectivity</title>
<p>The “cross-modal” view of functional reorganization in EB implies greater functional connectivity between visual cortex from non-deprived sensory cortices (Wittenberg et al., <xref rid="B146" ref-type="bibr">2004</xref>
). Results obtained using dynamic causal modeling (DCM) support the “cross-modal” view (Noppeney et al., <xref rid="B95" ref-type="bibr">2005</xref>
; Ptito et al., <xref rid="B108" ref-type="bibr">2008</xref>
; Klinge et al., <xref rid="B74" ref-type="bibr">2010</xref>
; Bedny et al., <xref rid="B7" ref-type="bibr">2011</xref>
; Collignon et al., <xref rid="B35" ref-type="bibr">2011</xref>
, <xref rid="B32" ref-type="bibr">2013</xref>
; Kupers and Ptito, <xref rid="B79" ref-type="bibr">2011</xref>
; Ma and Han, <xref rid="B88" ref-type="bibr">2011</xref>
; Park et al., <xref rid="B101" ref-type="bibr">2011</xref>
; Ricciardi and Pietrini, <xref rid="B114" ref-type="bibr">2011</xref>
; Leo et al., <xref rid="B82" ref-type="bibr">2012</xref>
). These DCM results are consistent with fiber tracings in primates (Falchier et al., <xref rid="B43" ref-type="bibr">2002</xref>
; Rockland and Ojima, <xref rid="B117" ref-type="bibr">2003</xref>
) and diffusion tensor tractography data in humans showing fiber tracts between auditory and visual cortex (Beer et al., <xref rid="B8" ref-type="bibr">2011</xref>
). However, DCM is limited to a few ROIs at one time, selected <italic>a priori</italic>
to study directed influences. Therefore, DCM studies, by design, cannot detect spatial reorganization in an unbiased fashion. Also, DCM targets “effective connectivity,” i.e., correlated responses induced by tasks or exogenous stimuli (as in the TMS study of Wittenberg et al., <xref rid="B146" ref-type="bibr">2004</xref>
). Functional connectivity based on embedded “rest” intervals is also subject to contamination from persistent responses evoked by task paradigms. Unfortunately, results obtained both by DCM and correlation analysis in the context of task performance are often referred to as “functional connectivity” (e.g., Bedny et al., <xref rid="B7" ref-type="bibr">2011</xref>
; Collignon et al., <xref rid="B35" ref-type="bibr">2011</xref>
). Such results are not directly comparable to true resting state results, as reported here.</p>
<p>The presented results indicate that EB compared to NS show decreased resting state functional connectivity between visual and somatosensory/auditory cortex, which is contrary to expectations based on a “cross-modal” hypothesis. Prior fMRI studies of resting-state activity also indicate decreased functional connectivity between visual and non-deprived sensory cortex (Liu et al., <xref rid="B86" ref-type="bibr">2007</xref>
; Yu et al., <xref rid="B147" ref-type="bibr">2008</xref>
). These resting state results are concordant with diffusion tensor imaging data showing absent/reduced axonal connectivity between visual and other sensory cortices (Shu et al., <xref rid="B126" ref-type="bibr">2009</xref>
).</p>
<p>The physiological substrate supporting enhanced EB visual cortex responses to non-visual tasks still needs identification. One possibility is through intra-visual cortex connections. Most extra-striate areas are multisensory in sighted individuals. In EB, non-visual perceptual tasks activate extra-striate regions that, in NS, typically respond to visual stimulation. Examples include lateral occipital cortex activation in response to tactile object recognition (Amedi et al., <xref rid="B1" ref-type="bibr">2001</xref>
, <xref rid="B4" ref-type="bibr">2005</xref>
, <xref rid="B2" ref-type="bibr">2010</xref>
; Pietrini et al., <xref rid="B105" ref-type="bibr">2004</xref>
); medial temporal area (MT) responses to perceived motion of non-visual stimuli (Hagen et al., <xref rid="B62" ref-type="bibr">2002</xref>
; Poirier et al., <xref rid="B106" ref-type="bibr">2006</xref>
; Ricciardi et al., <xref rid="B115" ref-type="bibr">2007</xref>
; Sani et al., <xref rid="B123" ref-type="bibr">2010</xref>
), and superior occipital cortex responses to spatial localization of sounds (Gougoux et al., <xref rid="B57" ref-type="bibr">2005</xref>
; Collignon et al., <xref rid="B33" ref-type="bibr">2007</xref>
, <xref rid="B35" ref-type="bibr">2011</xref>
, <xref rid="B32" ref-type="bibr">2013</xref>
) or touch (Ricciardi et al., <xref rid="B112" ref-type="bibr">2006</xref>
; Bonino et al., <xref rid="B10" ref-type="bibr">2008</xref>
). These cross-modal responses can be mediated by existing connections, as shown by reversible cross-modal activation in visual cortex of NS volunteers after relatively short periods of blindfolding (Kauffman et al., <xref rid="B73" ref-type="bibr">2002</xref>
; Weisser et al., <xref rid="B143" ref-type="bibr">2005</xref>
; Merabet et al., <xref rid="B93" ref-type="bibr">2007</xref>
, <xref rid="B91" ref-type="bibr">2008</xref>
; Lazzouni et al., <xref rid="B81" ref-type="bibr">2012</xref>
). Consequently, cross-modal activity in visual cortex of EB possibly involves native connections. However, like Yu and colleagues (Yu et al., <xref rid="B147" ref-type="bibr">2008</xref>
), we observed generally reduced intra-visual cortex functional connectivity in EB. The enhancement of cross-modal activity might then be a consequence of excitability changes resulting from visual deprivation, which even short periods of blindfolding elicit in NS (Boroojerdi et al., <xref rid="B12" ref-type="bibr">2000</xref>
, <xref rid="B11" ref-type="bibr">2001</xref>
; Merabet et al., <xref rid="B91" ref-type="bibr">2008</xref>
). Thus, cross-modal reorganization of visual cortex in EB is possibly a strengthened experience-dependent plasticity. Alternatively, these effects might arise from deprivation-induced developmental changes, not dependent on experience or skill acquisition (Bavelier and Neville, <xref rid="B6" ref-type="bibr">2002</xref>
). Adjudicating between these alternatives will require animal experiments.</p>
<p>The significance of reduced interhemispheric functional connectivity in higher-tier visual areas is uncertain. Similar findings were reported in patients with anophthalmia (Watkins et al., <xref rid="B141" ref-type="bibr">2012</xref>
). These results coincide with evidence of reduced white matter in the ventral splenium of EB (Shimony et al., <xref rid="B125" ref-type="bibr">2006</xref>
). Less inter-visual connectivity might explain previously described differences between EB and sighted in spatially discriminating tactile inputs (Röder et al., <xref rid="B119" ref-type="bibr">2004</xref>
). EB were better and faster than sighted at detecting the temporal order between two successive tactile stimuli applied to hands crossed over the midline (Röder et al., <xref rid="B119" ref-type="bibr">2004</xref>
). In sighted persons, vision subjugates localization of tactile events to a visual space (Eimer, <xref rid="B41" ref-type="bibr">2004</xref>
). Consequently, touching crossed hands in sighted provoked slower reactions from a conflict between visual and hand somatosensory space. Reaction times were faster in EB with only a tactile, proprioceptive coordinate space. Potentially further reducing confusion might be lower inter-occipital functional connectivity between higher tier visual areas like those in the superior occipital gyrus that serve tactile spatial working memory in EB (Bonino et al., <xref rid="B10" ref-type="bibr">2008</xref>
).</p>
</sec>
<sec><title>Visual to cognitive network functional connectivity</title>
<p>Blindness requires greater reliance on remembering. Even blind children show compensatory, adaptive strengthening of memory skills (Withagen et al., <xref rid="B145" ref-type="bibr">2013</xref>
). These effects suggest long-term changes in synaptic efficacy. A consequence might be the finding of increased functional connectivity between visual and dorsolateral frontal cortex in EB for a persistently used network (Dosenbach et al., <xref rid="B40" ref-type="bibr">2007</xref>
; Fair et al., <xref rid="B42" ref-type="bibr">2007</xref>
; Lewis et al., <xref rid="B84" ref-type="bibr">2009</xref>
). The dorsolateral prefrontal cortex is a known contributor to recognition memory involving familiarity and remembering (Iidaka et al., <xref rid="B70" ref-type="bibr">2000</xref>
; Mcdermott et al., <xref rid="B90" ref-type="bibr">2000</xref>
; Gold and Buckner, <xref rid="B54" ref-type="bibr">2002</xref>
; Gallo et al., <xref rid="B51" ref-type="bibr">2010</xref>
). A further implication of increased functional connectivity with dorsolateral prefrontal cortex is that visual cortex contributes to memory processes in EB. A finding consistent with this notion is activity throughout visual cortex in EB for semantic tasks that engage lexical memory (Burton et al., <xref rid="B25" ref-type="bibr">2002a</xref>
,<xref rid="B26" ref-type="bibr">b</xref>
, <xref rid="B17" ref-type="bibr">2003</xref>
). As suggested in a previous study involving recognition memory for studied words in EB, visual cortex activity possibly generates a recollection heuristic that augments remembering as opposed to familiarity (Burton et al., <xref rid="B21" ref-type="bibr">2012b</xref>
).</p>
<p>A prior functional connectivity study suggested that left visual cortex specifically contributes to semantic and lexical processing in EB, based on greater connectivity with left inferior frontal cortex language areas (Bedny et al., <xref rid="B7" ref-type="bibr">2011</xref>
). The current study observed neither a left hemisphere functional connectivity bias nor a significant TFCE cluster confined to left inferior frontal language areas. This negative result is consistent with diffusion tensor tractography evidence of balanced bilateral inferior frontal-occipital (IFO) fasciculi (Shimony et al., <xref rid="B125" ref-type="bibr">2006</xref>
; Shu et al., <xref rid="B126" ref-type="bibr">2009</xref>
).</p>
<p>The notion of a language cognition role for visual cortex in EB was invoked to explain the results of TMS experiments (Cohen et al., <xref rid="B31" ref-type="bibr">1997</xref>
). Further support was obtained from the observation that Braille alexia may follow visual cortex strokes (Hamilton et al., <xref rid="B64" ref-type="bibr">2000</xref>
; Maeda and Yasuda, <xref rid="B89" ref-type="bibr">2003</xref>
). An alternative explanation is that the visual cortex lesions might have disturbed attention to memory processes. Thus, patients with Braille alexia might be unable to attend to tactile features during fluent braille reading, which is an attention demanding perceptual task focused on lateral shearing produced by the pattern of dot-gaps within braille cells (Millar, <xref rid="B94" ref-type="bibr">1987</xref>
; Pring, <xref rid="B107" ref-type="bibr">1994</xref>
).</p>
<p>Exceptional “attentive processing of stimuli” (p. 118 in Kujala et al., <xref rid="B77" ref-type="bibr">2000</xref>
) probably reflect learned behaviors in EB. Visual cortex activation during auditory spatial localization tasks (Kujala et al., <xref rid="B77" ref-type="bibr">2000</xref>
; Collignon et al., <xref rid="B34" ref-type="bibr">2006</xref>
; Weaver and Stevens, <xref rid="B142" ref-type="bibr">2007</xref>
) have been conventionally interpreted as reflecting “cross-modal” processes, but they might be alternatively interpreted as reflecting enhanced top-down attentional mechanisms. EB show activation of visual and parietal attention areas during a tactile working memory task (Burton et al., <xref rid="B22" ref-type="bibr">2010</xref>
). These adaptive behaviors in EB might alter synaptic efficacies that result in increased functional connectivity between visual and cortical regions concerned with attention. Present results supporting this interpretation include increased functional connectivity between visual and anterior inferior parietal cortex, superior part of the supramarginal gyrus, anterior insula, and posterior middle frontal gyrus. Anterior inferior parietal cortex contributes to attention modulated task switching (Vincent et al., <xref rid="B137" ref-type="bibr">2008</xref>
). Anterior insula serves cognitive switching in conflict situations (Roberts and Hall, <xref rid="B116" ref-type="bibr">2008</xref>
; Sridharan et al., <xref rid="B130" ref-type="bibr">2008</xref>
). Supramarginal gyrus is active during attention to memory (Cabeza, <xref rid="B28" ref-type="bibr">2008</xref>
; Cabeza et al., <xref rid="B29" ref-type="bibr">2008</xref>
; Ciaramelli et al., <xref rid="B30" ref-type="bibr">2008</xref>
; Olson and Berryhill, <xref rid="B100" ref-type="bibr">2009</xref>
). Similar supramarginal gyrus responses can be seen in responses to working, long-term and episodic memory paradigms (Buckner and Wheeler, <xref rid="B15" ref-type="bibr">2001</xref>
). Posterior middle frontal gyrus responses are correlates of orienting to salient, yet unexpected stimuli (Corbetta and Shulman, <xref rid="B36" ref-type="bibr">2002</xref>
).</p>
</sec>
</sec>
<sec><title>Conclusions and study limitations</title>
<p>A crucial accommodation to blindness is reliance on memory and facile attentional switching between preserved non-visual modalities of the sensorium. Therefore, understanding possible changes in synaptic efficacies that might underlie functional connectivity differences in EB requires awareness of these adaptive behaviors. Persistent behavioral adaptations can strengthen synaptic efficacy in functionally connected networks (Lewis et al., <xref rid="B84" ref-type="bibr">2009</xref>
). Furthermore, greater correlations occur within networks serving identifiable functions. In EB, resting state fMRI differences included increased functional connectivity between visual cortex and cognitive control networks concerned with memory, attention, or task switching in frontal and parietal cortex. In contrast, there was decreased functional connectivity between visual and non-visual sensory networks and within extra-striate visual cortex. Most probably, extra-striate areas with resident multisensory inputs conveyed cross-modal inputs through existing connections with non-visual cortex. The enhancement of cross-modal activity might then result from adapting to visual deprivation.</p>
<p>Crucial to functional connectivity analyses is isolating brain-derived signals from artifact (Cordes et al., <xref rid="B37" ref-type="bibr">2001</xref>
). The current methodology relies on regression of nuisance signals, including the global signal (GSR) (Fox et al., <xref rid="B47" ref-type="bibr">2009</xref>
). It has been pointed out that GSR can artificially induce group differences if the global signal is unequally distributed in the two groups (Saad et al., <xref rid="B121" ref-type="bibr">2012</xref>
), which was not the case between the EB and NS groups.</p>
<p>A principal limitation of this study is that the functionality represented in the visual cortex of EB depended entirely on the extant literature reporting task-based fMRI responses to memory and attention paradigms. However, we submit that this literature is sufficiently mature to support such use. The relatively small participant sample size was limited by requiring that all EB were blind from peripheral pathologies and be Braille literate from childhood. Echo planar imaging data also had to meet standards of minimal movement (Hacker et al., <xref rid="B61" ref-type="bibr">2012</xref>
). A larger sample of younger participants with a range of Braille literacy may permit identification of age and literacy effects on resting-state activity.</p>
<sec><title>Conflict of interest statement</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
</sec>
</body>
<back><ack><p>The content is solely the responsibility of the authors and does not necessarily represent the official views of the National Institute of Neurological Disorders and Stroke or the National Institutes of Health. Contract grant sponsor: NIH; Contract grant numbers: NS06833 (Marcus E. Raichle, Abraham Z. Snyder), P30NS048056 (Abraham Z. Snyder) and NS37237 (Harold Burton). We especially are grateful for the assistance of Dr. Jill Firszt and Tim Holden for imaging data from normally sighted individuals who were participants in an R01 supported by the National Institutes of Deafness and Other Communication Disorders: Contract grant number DC009010 (JBF).</p>
</ack>
<fn-group><fn id="fn0001"><p><sup>1</sup>
We use “visual cortex” as an anatomic descriptor to indicate cortex that normally is associated with visual function in sighted persons.</p>
</fn>
<fn id="fn0002"><p><sup>2</sup>
The average signal values for the global regressor were not significantly different between the groups.</p>
</fn>
<fn id="fn0003"><p><sup>3</sup>
We used three of four runs obtained from EB that showed the least head movement.</p>
</fn>
</fn-group>
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