550 lines
No EOL
98 KiB
HTML
550 lines
No EOL
98 KiB
HTML
<?xml version="1.0" encoding="utf-8"?>
|
|
<!DOCTYPE html PUBLIC "-//W3C//DTD XHTML 1.0 Transitional//EN" "http://www.w3.org/TR/xhtml1/DTD/xhtml1-transitional.dtd">
|
|
<html xmlns="http://www.w3.org/1999/xhtml" xml:lang="en" lang="en">
|
|
|
|
<head><meta http-equiv="Content-Type" content="text/html; charset=utf-8" />
|
|
<!-- AppResources meta begin -->
|
|
<meta name="paf-app-resources" content="" />
|
|
<script type="text/javascript">var ncbi_startTime = new Date();</script>
|
|
|
|
<!-- AppResources meta end -->
|
|
|
|
<!-- TemplateResources meta begin -->
|
|
<meta name="paf_template" content="" />
|
|
|
|
<!-- TemplateResources meta end -->
|
|
|
|
<!-- Logger begin -->
|
|
<meta name="ncbi_db" content="books" /><meta name="ncbi_pdid" content="book-part" /><meta name="ncbi_acc" content="NBK11536" /><meta name="ncbi_domain" content="webvision" /><meta name="ncbi_report" content="record" /><meta name="ncbi_type" content="fulltext" /><meta name="ncbi_objectid" content="" /><meta name="ncbi_pcid" content="/NBK11536/" /><meta name="ncbi_pagename" content="Inner Plexiform Layer - Webvision - NCBI Bookshelf" /><meta name="ncbi_bookparttype" content="chapter" /><meta name="ncbi_app" content="bookshelf" />
|
|
<!-- Logger end -->
|
|
|
|
<title>Inner Plexiform Layer - Webvision - NCBI Bookshelf</title>
|
|
|
|
<!-- AppResources external_resources begin -->
|
|
<link rel="stylesheet" href="/core/jig/1.15.2/css/jig.min.css" /><script type="text/javascript" src="/core/jig/1.15.2/js/jig.min.js"></script>
|
|
|
|
<!-- AppResources external_resources end -->
|
|
|
|
<!-- Page meta begin -->
|
|
<meta name="robots" content="INDEX,FOLLOW,NOARCHIVE" /><meta name="citation_inbook_title" content="Webvision: The Organization of the Retina and Visual System [Internet]" /><meta name="citation_title" content="Inner Plexiform Layer" /><meta name="citation_publisher" content="University of Utah Health Sciences Center" /><meta name="citation_date" content="2012/03/25" /><meta name="citation_author" content="Helga Kolb" /><meta name="citation_pmid" content="21413394" /><meta name="citation_fulltext_html_url" content="https://www.ncbi.nlm.nih.gov/books/NBK11536/" /><link rel="schema.DC" href="http://purl.org/DC/elements/1.0/" /><meta name="DC.Title" content="Inner Plexiform Layer" /><meta name="DC.Type" content="Text" /><meta name="DC.Publisher" content="University of Utah Health Sciences Center" /><meta name="DC.Contributor" content="Helga Kolb" /><meta name="DC.Date" content="2012/03/25" /><meta name="DC.Identifier" content="https://www.ncbi.nlm.nih.gov/books/NBK11536/" /><meta name="description" content="The axonal endings of bipolar cells bring information from the outer plexiform layer (OPL) to the neuropil of the inner plexiform layer (IPL). Here bipolar cells talk to different varieties of functionally specialized amacrine cells and to dendrites of the various ganglion cells (Figs 1 and 2). The neuropil is a confusing network of interconnecting profiles that, to be understood, has to be investigated at the higher magnification afforded by the electron microscope over, the light microscope, and with knowledge gained from Golgi-staining for morphology, and intracellular electrophysiology for function of individual cells in the network." /><meta name="og:title" content="Inner Plexiform Layer" /><meta name="og:type" content="book" /><meta name="og:description" content="The axonal endings of bipolar cells bring information from the outer plexiform layer (OPL) to the neuropil of the inner plexiform layer (IPL). Here bipolar cells talk to different varieties of functionally specialized amacrine cells and to dendrites of the various ganglion cells (Figs 1 and 2). The neuropil is a confusing network of interconnecting profiles that, to be understood, has to be investigated at the higher magnification afforded by the electron microscope over, the light microscope, and with knowledge gained from Golgi-staining for morphology, and intracellular electrophysiology for function of individual cells in the network." /><meta name="og:url" content="https://www.ncbi.nlm.nih.gov/books/NBK11536/" /><meta name="og:site_name" content="NCBI Bookshelf" /><meta name="og:image" content="https://www.ncbi.nlm.nih.gov/corehtml/pmc/pmcgifs/bookshelf/thumbs/th-webvision-lrg.png" /><meta name="twitter:card" content="summary" /><meta name="twitter:site" content="@ncbibooks" /><meta name="bk-non-canon-loc" content="/books/n/webvision/ch06ipl/" /><link rel="canonical" href="https://www.ncbi.nlm.nih.gov/books/NBK11536/" /><link rel="stylesheet" href="/corehtml/pmc/css/figpopup.css" type="text/css" media="screen" /><link rel="stylesheet" href="/corehtml/pmc/css/bookshelf/2.26/css/books.min.css" type="text/css" /><link rel="stylesheet" href="/corehtml/pmc/css/bookshelf/2.26/css/books_print.min.css" type="text/css" media="print" /><style type="text/css">p a.figpopup{display:inline !important} .bk_tt {font-family: monospace} .first-line-outdent .bk_ref {display: inline} .body-content h2, .body-content .h2 {border-bottom: 1px solid #97B0C8} .body-content h2.inline {border-bottom: none} a.page-toc-label , .jig-ncbismoothscroll a {text-decoration:none;border:0 !important} .temp-labeled-list .graphic {display:inline-block !important} .temp-labeled-list img{width:100%}</style><script type="text/javascript" src="/corehtml/pmc/js/jquery.hoverIntent.min.js"> </script><script type="text/javascript" src="/corehtml/pmc/js/common.min.js?_=3.18"> </script><script type="text/javascript" src="/corehtml/pmc/js/large-obj-scrollbars.min.js"> </script><script type="text/javascript">window.name="mainwindow";</script><script type="text/javascript" src="/corehtml/pmc/js/bookshelf/2.26/book-toc.min.js"> </script><script type="text/javascript" src="/corehtml/pmc/js/bookshelf/2.26/books.min.js"> </script><meta name="book-collection" content="NONE" />
|
|
|
|
<!-- Page meta end -->
|
|
<link rel="shortcut icon" href="//www.ncbi.nlm.nih.gov/favicon.ico" /><meta name="ncbi_phid" content="CE8C4DB17C9A458100000000003F0033.m_13" />
|
|
<meta name='referrer' content='origin-when-cross-origin'/><link type="text/css" rel="stylesheet" href="//static.pubmed.gov/portal/portal3rc.fcgi/4216699/css/3852956/3985586/3808861/4121862/3974050/3917732/251717/4216701/14534/45193/4113719/3849091/3984811/3751656/4033350/3840896/3577051/3852958/4008682/4207974/4206132/4062871/12930/3964959/3854974/36029/4128070/9685/3549676/3609192/3609193/3609213/3395586.css" /><link type="text/css" rel="stylesheet" href="//static.pubmed.gov/portal/portal3rc.fcgi/4216699/css/3411343/3882866.css" media="print" /></head>
|
|
<body class="book-part">
|
|
<div class="grid">
|
|
<div class="col twelve_col nomargin shadow">
|
|
<!-- System messages like service outage or JS required; this is handled by the TemplateResources portlet -->
|
|
<div class="sysmessages">
|
|
<noscript>
|
|
<p class="nojs">
|
|
<strong>Warning:</strong>
|
|
The NCBI web site requires JavaScript to function.
|
|
<a href="/guide/browsers/#enablejs" title="Learn how to enable JavaScript" target="_blank">more...</a>
|
|
</p>
|
|
</noscript>
|
|
</div>
|
|
<!--/.sysmessage-->
|
|
<div class="wrap">
|
|
<div class="page">
|
|
<div class="top">
|
|
<div id="universal_header">
|
|
<section class="usa-banner">
|
|
<div class="usa-accordion">
|
|
<header class="usa-banner-header">
|
|
<div class="usa-grid usa-banner-inner">
|
|
<img src="https://www.ncbi.nlm.nih.gov/coreutils/uswds/img/favicons/favicon-57.png" alt="U.S. flag" />
|
|
<p>An official website of the United States government</p>
|
|
<button class="non-usa-accordion-button usa-banner-button" aria-expanded="false" aria-controls="gov-banner-top" type="button">
|
|
<span class="usa-banner-button-text">Here's how you know</span>
|
|
</button>
|
|
</div>
|
|
</header>
|
|
<div class="usa-banner-content usa-grid usa-accordion-content" id="gov-banner-top" aria-hidden="true">
|
|
<div class="usa-banner-guidance-gov usa-width-one-half">
|
|
<img class="usa-banner-icon usa-media_block-img" src="https://www.ncbi.nlm.nih.gov/coreutils/uswds/img/icon-dot-gov.svg" alt="Dot gov" />
|
|
<div class="usa-media_block-body">
|
|
<p>
|
|
<strong>The .gov means it's official.</strong>
|
|
<br />
|
|
Federal government websites often end in .gov or .mil. Before
|
|
sharing sensitive information, make sure you're on a federal
|
|
government site.
|
|
</p>
|
|
</div>
|
|
</div>
|
|
<div class="usa-banner-guidance-ssl usa-width-one-half">
|
|
<img class="usa-banner-icon usa-media_block-img" src="https://www.ncbi.nlm.nih.gov/coreutils/uswds/img/icon-https.svg" alt="Https" />
|
|
<div class="usa-media_block-body">
|
|
<p>
|
|
<strong>The site is secure.</strong>
|
|
<br />
|
|
The <strong>https://</strong> ensures that you are connecting to the
|
|
official website and that any information you provide is encrypted
|
|
and transmitted securely.
|
|
</p>
|
|
</div>
|
|
</div>
|
|
</div>
|
|
</div>
|
|
</section>
|
|
<div class="usa-overlay"></div>
|
|
<header class="ncbi-header" role="banner" data-section="Header">
|
|
|
|
<div class="usa-grid">
|
|
<div class="usa-width-one-whole">
|
|
|
|
<div class="ncbi-header__logo">
|
|
<a href="/" class="logo" aria-label="NCBI Logo" data-ga-action="click_image" data-ga-label="NIH NLM Logo">
|
|
<img src="https://www.ncbi.nlm.nih.gov/coreutils/nwds/img/logos/AgencyLogo.svg" alt="NIH NLM Logo" />
|
|
</a>
|
|
</div>
|
|
|
|
<div class="ncbi-header__account">
|
|
<a id="account_login" href="https://account.ncbi.nlm.nih.gov" class="usa-button header-button" style="display:none" data-ga-action="open_menu" data-ga-label="account_menu">Log in</a>
|
|
<button id="account_info" class="header-button" style="display:none" aria-controls="account_popup" type="button">
|
|
<span class="fa fa-user" aria-hidden="true">
|
|
<svg xmlns="http://www.w3.org/2000/svg" viewBox="0 0 24 24" width="20px" height="20px">
|
|
<g style="fill: #fff">
|
|
<ellipse cx="12" cy="8" rx="5" ry="6"></ellipse>
|
|
<path d="M21.8,19.1c-0.9-1.8-2.6-3.3-4.8-4.2c-0.6-0.2-1.3-0.2-1.8,0.1c-1,0.6-2,0.9-3.2,0.9s-2.2-0.3-3.2-0.9 C8.3,14.8,7.6,14.7,7,15c-2.2,0.9-3.9,2.4-4.8,4.2C1.5,20.5,2.6,22,4.1,22h15.8C21.4,22,22.5,20.5,21.8,19.1z"></path>
|
|
</g>
|
|
</svg>
|
|
</span>
|
|
<span class="username desktop-only" aria-hidden="true" id="uname_short"></span>
|
|
<span class="sr-only">Show account info</span>
|
|
</button>
|
|
</div>
|
|
|
|
<div class="ncbi-popup-anchor">
|
|
<div class="ncbi-popup account-popup" id="account_popup" aria-hidden="true">
|
|
<div class="ncbi-popup-head">
|
|
<button class="ncbi-close-button" data-ga-action="close_menu" data-ga-label="account_menu" type="button">
|
|
<span class="fa fa-times">
|
|
<svg xmlns="http://www.w3.org/2000/svg" viewBox="0 0 48 48" width="24px" height="24px">
|
|
<path d="M38 12.83l-2.83-2.83-11.17 11.17-11.17-11.17-2.83 2.83 11.17 11.17-11.17 11.17 2.83 2.83 11.17-11.17 11.17 11.17 2.83-2.83-11.17-11.17z"></path>
|
|
</svg>
|
|
</span>
|
|
<span class="usa-sr-only">Close</span></button>
|
|
<h4>Account</h4>
|
|
</div>
|
|
<div class="account-user-info">
|
|
Logged in as:<br />
|
|
<b><span class="username" id="uname_long">username</span></b>
|
|
</div>
|
|
<div class="account-links">
|
|
<ul class="usa-unstyled-list">
|
|
<li><a id="account_myncbi" href="/myncbi/" class="set-base-url" data-ga-action="click_menu_item" data-ga-label="account_myncbi">Dashboard</a></li>
|
|
<li><a id="account_pubs" href="/myncbi/collections/bibliography/" class="set-base-url" data-ga-action="click_menu_item" data-ga-label="account_pubs">Publications</a></li>
|
|
<li><a id="account_settings" href="/account/settings/" class="set-base-url" data-ga-action="click_menu_item" data-ga-label="account_settings">Account settings</a></li>
|
|
<li><a id="account_logout" href="/account/signout/" class="set-base-url" data-ga-action="click_menu_item" data-ga-label="account_logout">Log out</a></li>
|
|
</ul>
|
|
</div>
|
|
</div>
|
|
</div>
|
|
|
|
</div>
|
|
</div>
|
|
</header>
|
|
<div role="navigation" aria-label="access keys">
|
|
<a id="nws_header_accesskey_0" href="https://www.ncbi.nlm.nih.gov/guide/browsers/#ncbi_accesskeys" class="usa-sr-only" accesskey="0" tabindex="-1">Access keys</a>
|
|
<a id="nws_header_accesskey_1" href="https://www.ncbi.nlm.nih.gov" class="usa-sr-only" accesskey="1" tabindex="-1">NCBI Homepage</a>
|
|
<a id="nws_header_accesskey_2" href="/myncbi/" class="set-base-url usa-sr-only" accesskey="2" tabindex="-1">MyNCBI Homepage</a>
|
|
<a id="nws_header_accesskey_3" href="#maincontent" class="usa-sr-only" accesskey="3" tabindex="-1">Main Content</a>
|
|
<a id="nws_header_accesskey_4" href="#" class="usa-sr-only" accesskey="4" tabindex="-1">Main Navigation</a>
|
|
</div>
|
|
<section data-section="Alerts">
|
|
<div class="ncbi-alerts-placeholder"></div>
|
|
</section>
|
|
</div>
|
|
<div class="header">
|
|
<div class="res_logo"><h1 class="res_name"><a href="/books/" title="Bookshelf home">Bookshelf</a></h1><h2 class="res_tagline"></h2></div>
|
|
<div class="search"><form method="get" action="/books/"><div class="search_form"><label for="database" class="offscreen_noflow">Search database</label><select id="database"><optgroup label="Recent"><option value="books" selected="selected" data-ac_dict="bookshelf-search">Books</option><option value="snp">SNP</option><option value="nlmcatalog">NLM Catalog</option><option value="pcsubstance" class="last">PubChem Substance</option></optgroup><optgroup label="All"><option value="gquery">All Databases</option><option value="assembly">Assembly</option><option value="biocollections">Biocollections</option><option value="bioproject">BioProject</option><option value="biosample">BioSample</option><option value="books" data-ac_dict="bookshelf-search">Books</option><option value="clinvar">ClinVar</option><option value="cdd">Conserved Domains</option><option value="gap">dbGaP</option><option value="dbvar">dbVar</option><option value="gene">Gene</option><option value="genome">Genome</option><option value="gds">GEO DataSets</option><option value="geoprofiles">GEO Profiles</option><option value="gtr">GTR</option><option value="ipg">Identical Protein Groups</option><option value="medgen">MedGen</option><option value="mesh">MeSH</option><option value="nlmcatalog">NLM Catalog</option><option value="nuccore">Nucleotide</option><option value="omim">OMIM</option><option value="pmc">PMC</option><option value="protein">Protein</option><option value="proteinclusters">Protein Clusters</option><option value="protfam">Protein Family Models</option><option value="pcassay">PubChem BioAssay</option><option value="pccompound">PubChem Compound</option><option value="pcsubstance">PubChem Substance</option><option value="pubmed">PubMed</option><option value="snp">SNP</option><option value="sra">SRA</option><option value="structure">Structure</option><option value="taxonomy">Taxonomy</option><option value="toolkit">ToolKit</option><option value="toolkitall">ToolKitAll</option><option value="toolkitbookgh">ToolKitBookgh</option></optgroup></select><div class="nowrap"><label for="term" class="offscreen_noflow" accesskey="/">Search term</label><div class="nowrap"><input type="text" name="term" id="term" title="Search Books. Use up and down arrows to choose an item from the autocomplete." value="" class="jig-ncbiclearbutton jig-ncbiautocomplete" data-jigconfig="dictionary:'bookshelf-search',disableUrl:'NcbiSearchBarAutoComplCtrl'" autocomplete="off" data-sbconfig="ds:'no',pjs:'no',afs:'no'" /></div><button id="search" type="submit" class="button_search nowrap" cmd="go">Search</button></div></div></form><ul class="searchlinks inline_list"><li>
|
|
<a href="/books/browse/">Browse Titles</a>
|
|
</li><li>
|
|
<a href="/books/advanced/">Advanced</a>
|
|
</li><li class="help">
|
|
<a href="/books/NBK3833/">Help</a>
|
|
</li><li class="disclaimer">
|
|
<a target="_blank" data-ga-category="literature_resources" data-ga-action="link_click" data-ga-label="disclaimer_link" href="https://www.ncbi.nlm.nih.gov/books/about/disclaimer/">Disclaimer</a>
|
|
</li></ul></div>
|
|
</div>
|
|
|
|
|
|
|
|
<!--<component id="Page" label="headcontent"/>-->
|
|
|
|
</div>
|
|
<div class="content">
|
|
<!-- site messages -->
|
|
<!-- Custom content 1 -->
|
|
<div class="col1">
|
|
|
|
</div>
|
|
|
|
<div class="container">
|
|
<div id="maincontent" class="content eight_col col">
|
|
<!-- Custom content in the left column above book nav -->
|
|
<div class="col2">
|
|
|
|
</div>
|
|
|
|
<!-- Book content -->
|
|
|
|
|
|
<!-- Custom content between navigation and content -->
|
|
<div class="col3">
|
|
|
|
</div>
|
|
|
|
<div class="document">
|
|
<div class="pre-content"><div><div class="bk_prnt"><p class="small">NCBI Bookshelf. A service of the National Library of Medicine, National Institutes of Health.</p><p>Kolb H, Fernandez E, Jones B, et al., editors. Webvision: The Organization of the Retina and Visual System [Internet]. Salt Lake City (UT): University of Utah Health Sciences Center; 1995-. </p></div><div class="iconblock clearfix whole_rhythm no_top_margin bk_noprnt"><a class="img_link icnblk_img" title="Table of Contents Page" href="/books/n/webvision/"><img class="source-thumb" src="/corehtml/pmc/pmcgifs/bookshelf/thumbs/th-webvision-lrg.png" alt="Cover of Webvision" height="100px" width="80px" /></a><div class="icnblk_cntnt eight_col"><h2>Webvision: The Organization of the Retina and Visual System [Internet].</h2><a data-jig="ncbitoggler" href="#__NBK11536_dtls__">Show details</a><div style="display:none" class="ui-widget" id="__NBK11536_dtls__"><div>Kolb H, Fernandez E, Jones B, et al., editors.</div><div>Salt Lake City (UT): <a href="http://webvision.med.utah.edu/" ref="pagearea=page-banner&targetsite=external&targetcat=link&targettype=publisher">University of Utah Health Sciences Center</a>; 1995-.</div></div><div class="half_rhythm"><ul class="inline_list"><li style="margin-right:1em"><a class="bk_cntns" href="/books/n/webvision/">Contents</a></li></ul></div><div class="bk_noprnt"><form method="get" action="/books/n/webvision/" id="bk_srch"><div class="bk_search"><label for="bk_term" class="offscreen_noflow">Search term</label><input type="text" title="Search this book" id="bk_term" name="term" value="" data-jig="ncbiclearbutton" /> <input type="submit" class="jig-ncbibutton" value="Search this book" submit="false" style="padding: 0.1em 0.4em;" /></div></form></div></div><div class="icnblk_cntnt two_col"><div class="pagination bk_noprnt"><a class="active page_link prev" href="/books/n/webvision/A207/" title="Previous page in this title">< Prev</a><a class="active page_link next" href="/books/n/webvision/ch07gc1/" title="Next page in this title">Next ></a></div></div></div></div></div>
|
|
<div class="main-content lit-style" itemscope="itemscope" itemtype="http://schema.org/CreativeWork"><div class="meta-content fm-sec"><h1 id="_NBK11536_"><span class="title" itemprop="name">Inner Plexiform Layer</span></h1><p class="contrib-group"><span itemprop="author">Helga Kolb</span>, PhD.</p><a data-jig="ncbitoggler" href="#__NBK11536_ai__" style="border:0;text-decoration:none">Author Information and Affiliations</a><div style="display:none" class="ui-widget" id="__NBK11536_ai__"><p class="contrib-group"><h4>Authors</h4><span itemprop="author">Helga Kolb</span>, PhD<sup><img src="/corehtml/pmc/pmcgifs/corrauth.gif" alt="corresponding author" /></sup><sup>1</sup>.</p><h4>Affiliations</h4><div class="affiliation"><sup>1</sup> Moran Eye Institute, University of Utah School of Medicine, Salt Lake City<div><span class="email-label">Email: </span><a href="mailto:dev@null" data-email="ude.hatu.csh@blok.agleH" class="oemail">ude.hatu.csh@blok.agleH</a></div></div><div><sup><img src="/corehtml/pmc/pmcgifs/corrauth.gif" alt="corresponding author" /></sup>Corresponding author.</div></div><p class="small">Last Update: <span itemprop="dateModified">March 25, 2012</span>.</p></div><div class="jig-ncbiinpagenav body-content whole_rhythm" data-jigconfig="allHeadingLevels: ['h2'],smoothScroll: false" itemprop="text"><div id="ch06ipl.sec1"><h2 id="_ch06ipl_sec1_">1. Bipolar, amacrine and ganglion cells interact in the inner plexiform layer.</h2><p>The axonal endings of bipolar cells bring information from the outer plexiform layer (OPL) to the neuropil of the inner plexiform layer (IPL). Here bipolar cells talk to different varieties of functionally specialized amacrine cells and to dendrites of the various ganglion cells (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F1/?report=objectonly" target="object" rid-figpopup="figch06iplF1" rid-ob="figobch06iplF1">Figs 1</a> and <a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F2/?report=objectonly" target="object" rid-figpopup="figch06iplF2" rid-ob="figobch06iplF2">2</a>). The neuropil is a confusing network of interconnecting profiles that, to be understood, has to be investigated at the higher magnification afforded by the electron microscope over, the light microscope, and with knowledge gained from Golgi-staining for morphology, and intracellular electrophysiology for function of individual cells in the network.</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF1" co-legend-rid="figlgndch06iplF1"><a href="/books/NBK11536/figure/ch06ipl.F1/?report=objectonly" target="object" title="Fig. 1. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF1" rid-ob="figobch06iplF1"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image002.gif" src-large="/books/NBK11536/bin/ch06ipl-Image002.jpg" alt="Fig. 1. " /></a><div class="icnblk_cntnt" id="figlgndch06iplF1"><h4 id="ch06ipl.F1"><a href="/books/NBK11536/figure/ch06ipl.F1/?report=objectonly" target="object" rid-ob="figobch06iplF1">Fig. 1. </a></h4><p class="float-caption no_bottom_margin">3-D block of retina with IPL highlighted </p></div></div><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF2" co-legend-rid="figlgndch06iplF2"><a href="/books/NBK11536/figure/ch06ipl.F2/?report=objectonly" target="object" title="Fig. 2. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF2" rid-ob="figobch06iplF2"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image003.gif" src-large="/books/NBK11536/bin/ch06ipl-Image003.jpg" alt="Fig. 2. " /></a><div class="icnblk_cntnt" id="figlgndch06iplF2"><h4 id="ch06ipl.F2"><a href="/books/NBK11536/figure/ch06ipl.F2/?report=objectonly" target="object" rid-ob="figobch06iplF2">Fig. 2. </a></h4><p class="float-caption no_bottom_margin">Light micrograph of a vertical section through the IPL </p></div></div></div><div id="ch06ipl.sec2"><h2 id="_ch06ipl_sec2_">2. Ultrastructure of the neuropil of the inner plexiform layer.</h2><p>A view of a small part of the neuropil of the inner plexiform layer is shown in <a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F3/?report=objectonly" target="object" rid-figpopup="figch06iplF3" rid-ob="figobch06iplF3">Fig. 3</a>. Bipolar cell axon terminals (red profiles) are vesicle-filled profiles, containing irregular, long mitochondria and neurotubules. Their synapses are typified by a small synaptic ribbon pointing into a wedge with two post synaptic profiles (known as a dyad) at the apex (arrows in <a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F3/?report=objectonly" target="object" rid-figpopup="figch06iplF3" rid-ob="figobch06iplF3">Fig. 3</a> and red dots in bipolar cell profiles, <a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F4a/?report=objectonly" target="object" rid-figpopup="figch06iplF4a" rid-ob="figobch06iplF4a">Fig. 4a</a>) (<a class="bk_pop" href="#ch06ipl.REF.dowling.1966">Dowling and Boycott, 1966</a>). Amacrine cell dendrites are also vesicle-filled and have round mitochondria and sometimes neurofilaments as well as neurotubules (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F3/?report=objectonly" target="object" rid-figpopup="figch06iplF3" rid-ob="figobch06iplF3">Fig. 3</a>). Amacrine profiles vary between being very small cross sections through thin straight tubes, and larger varicosities budding off the dendrites.</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF3" co-legend-rid="figlgndch06iplF3"><a href="/books/NBK11536/figure/ch06ipl.F3/?report=objectonly" target="object" title="Fig. 3. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF3" rid-ob="figobch06iplF3"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image004.gif" src-large="/books/NBK11536/bin/ch06ipl-Image004.jpg" alt="Fig. 3. " /></a><div class="icnblk_cntnt" id="figlgndch06iplF3"><h4 id="ch06ipl.F3"><a href="/books/NBK11536/figure/ch06ipl.F3/?report=objectonly" target="object" rid-ob="figobch06iplF3">Fig. 3. </a></h4><p class="float-caption no_bottom_margin">Electron micrograph of the neuropil of the IPL </p></div></div><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF4a" co-legend-rid="figlgndch06iplF4a"><a href="/books/NBK11536/figure/ch06ipl.F4a/?report=objectonly" target="object" title="Fig. 4a. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF4a" rid-ob="figobch06iplF4a"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image005.gif" src-large="/books/NBK11536/bin/ch06ipl-Image005.jpg" alt="Fig. 4a. " /></a><div class="icnblk_cntnt" id="figlgndch06iplF4a"><h4 id="ch06ipl.F4a"><a href="/books/NBK11536/figure/ch06ipl.F4a/?report=objectonly" target="object" rid-ob="figobch06iplF4a">Fig. 4a. </a></h4><p class="float-caption no_bottom_margin">Electron micrograph of reciprocal synapses in the IPL </p></div></div><p>Typically amacrine cells synapse upon other profiles (bipolar axons, amacrine cells or ganglion cell dendrites) in the enlarged varicosities at, what is known as a conventional synapse: the synapse consists of synaptic vesicles clustered at a pre- and post membrane density (blue arrows in <a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F3/?report=objectonly" target="object" rid-figpopup="figch06iplF3" rid-ob="figobch06iplF3">Fig. 3</a>, yellow dots in amacrine profile, <a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F4a/?report=objectonly" target="object" rid-figpopup="figch06iplF4a" rid-ob="figobch06iplF4a">Fig. 4a</a>) (<a class="bk_pop" href="#ch06ipl.REF.dowling.1966">Dowling and Boycott, 1966</a>). Ganglion cell dendrites (yellow profiles, <a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F3/?report=objectonly" target="object" rid-figpopup="figch06iplF3" rid-ob="figobch06iplF3">Figs. 3</a>) are recognizable as profiles lacking synaptic vesicles but containing ribosomes, neurotubules and filaments. Ganglion cell dendrites are seen to be postsynaptic to bipolar ribbon synapses and amacrine conventional synapses (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F4a/?report=objectonly" target="object" rid-figpopup="figch06iplF4a" rid-ob="figobch06iplF4a">Figs. 4a and b</a>). Bipolar cells are also postsynaptic to many amacrine synapses. Often amacrine cells make, what are called reciprocal synapses, to the bipolar cell axons from which they receive ribbon synapses (yellow dots, <a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F4a/?report=objectonly" target="object" rid-figpopup="figch06iplF4a" rid-ob="figobch06iplF4a">Fig. 4a</a>).</p><p>In the last ten years many studies have shown that different glutamate receptors (glutamate being the excitatory amino acid that is the neurotransmitter of the bipolar cells, See chapter on glutamate) is present on the postsynaptic dendrites of amacrine and ganglion cells. These include AMPA, NMDA and GluRs of different types (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F4b/?report=objectonly" target="object" rid-figpopup="figch06iplF4b" rid-ob="figobch06iplF4b">Fig. 4b</a>) (<a class="bk_pop" href="#ch06ipl.REF.grunert.2002.138">Grunert et al., 2002</a>). The postsynaptic receptors of IPL processes for the inhibitory neurotransmitter glycine, of amacrine cells, are the various glycine receptors (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F4b/?report=objectonly" target="object" rid-figpopup="figch06iplF4b" rid-ob="figobch06iplF4b">Fig. 4b</a>). For the inhibitory transmitter GABA, of other amacrine cells, the receptors are typically GABA<sub>A</sub> subtypes (<a class="bk_pop" href="#ch06ipl.REF.grunert.2000.130">Grunert, 2000</a>; <a class="bk_pop" href="#ch06ipl.REF.kneussel.2000.1">Kneussel and Betz, 2000</a>), but in some cases, are GABA<sub>c</sub> subtypes (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F4b/?report=objectonly" target="object" rid-figpopup="figch06iplF4b" rid-ob="figobch06iplF4b">Fig. 4b</a>). The GABA<sub>c</sub> receptors are typical of the reciprocal synapses from the wide field amacrine cell (A17) at the ribbon synapse in the rod bipolar axon terminal (<a class="bk_pop" href="#ch06ipl.REF.chavez.2010.2330">Chavez at al., 2010</a>).</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF4b" co-legend-rid="figlgndch06iplF4b"><a href="/books/NBK11536/figure/ch06ipl.F4b/?report=objectonly" target="object" title="Fig. 4b. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF4b" rid-ob="figobch06iplF4b"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image006.gif" src-large="/books/NBK11536/bin/ch06ipl-Image006.jpg" alt="Fig. 4b. . Diagram of the bipolar synapses to ganglion and amacrine cells to show types of receptors that are now thought to be typical at the postsynaptic sites." /></a><div class="icnblk_cntnt" id="figlgndch06iplF4b"><h4 id="ch06ipl.F4b"><a href="/books/NBK11536/figure/ch06ipl.F4b/?report=objectonly" target="object" rid-ob="figobch06iplF4b">Fig. 4b. </a></h4><p class="float-caption no_bottom_margin">Diagram of the bipolar synapses to ganglion and amacrine cells to show types of receptors that are now thought to be typical at the postsynaptic sites. </p></div></div><p>This reciprocal synapse is in essence a feed-back synapse to the bipolar cell axon in the inner plexiform layer, with the same strategic importance as the horizontal cell dendrite making a feed-back synapse at the photoreceptor ribbon synapse, in the outer plexiform layer. It will be remembered that such a local circuit in the outer plexiform layer was thought to provide the bipolar cell response with a center-surround organization. It has been suggested that another surround mechanism coming from the amacrine cells could be added at this reciprocal (feed-back) synapse in the inner plexiform layer.</p></div><div id="ch06ipl.sec3"><h2 id="_ch06ipl_sec3_">3. Different morphological types of amacrine and ganglion cells.</h2><p>It is clear that there are many different kinds of amacrine cell and ganglion cell branching in the IPL of the human retina. From Golgi staining studies we know that there are at least 25 different amacrine cell types in the monkey and human retina (<a class="bk_pop" href="#ch06ipl.REF.mariani.1990.382">Mariani, 1990</a>; <a class="bk_pop" href="#ch06ipl.REF.kolb.1992.147">Kolb et al. 1992</a>). Some drawings of different amacrine cells as seen after Golgi staining in wholemounts of human retina, are shown below (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F5/?report=objectonly" target="object" rid-figpopup="figch06iplF5" rid-ob="figobch06iplF5">Fig. 5</a>).</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF5" co-legend-rid="figlgndch06iplF5"><a href="/books/NBK11536/figure/ch06ipl.F5/?report=objectonly" target="object" title="Fig. 5. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF5" rid-ob="figobch06iplF5"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image007.gif" src-large="/books/NBK11536/bin/ch06ipl-Image007.jpg" alt="Fig. 5. " /></a><div class="icnblk_cntnt" id="figlgndch06iplF5"><h4 id="ch06ipl.F5"><a href="/books/NBK11536/figure/ch06ipl.F5/?report=objectonly" target="object" rid-ob="figobch06iplF5">Fig. 5. </a></h4><p class="float-caption no_bottom_margin">Amacrine cells in human retina </p></div></div><p>The amacrine cells are classified into different types on morphological characteristics of dendritic tree size such as small, medium and large, branching characteristics (i.e. tufted, varicose, linear, beaded and radiate) and, most importantly, on the stratification of their dendrites in the IPL (<a class="bk_pop" href="#ch06ipl.REF.mariani.1990.382">Mariani, 1990</a>; <a class="bk_pop" href="#ch06ipl.REF.kolb.1992.147">Kolb et al., 1992</a>). The neuropil of the IPL was arbitrarily divided into 5 strata by <a class="bk_pop" href="#ch06ipl.REF.cajal.1892">Cajal (1892)</a>, because he appreciated the fact that cells branching in disparate strata could <b>not</b> make synaptic interactions while those that costratified could. This 5 strata descriptive classification scheme has been used by morphologists since to classify retinal cells on their dendritic branching level. See, for example in Fgigure 6 below, some amacrine cells have dendrites in strata 1 and 2 (called broadly stratified types, cell 8 below), or strata 1 and 5 (called bistratified types, cell 1, below), stratum 1 or any of the 5 strata (called monostratified, cell 13, below) or dendrite through all strata 1 to 5 (called diffuse types, cells 12 and 14, <a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F6/?report=objectonly" target="object" rid-figpopup="figch06iplF6" rid-ob="figobch06iplF6">Fig. 6</a>).</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF6" co-legend-rid="figlgndch06iplF6"><a href="/books/NBK11536/figure/ch06ipl.F6/?report=objectonly" target="object" title="Fig. 6. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF6" rid-ob="figobch06iplF6"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image008.gif" src-large="/books/NBK11536/bin/ch06ipl-Image008.jpg" alt="Fig. 6. " /></a><div class="icnblk_cntnt" id="figlgndch06iplF6"><h4 id="ch06ipl.F6"><a href="/books/NBK11536/figure/ch06ipl.F6/?report=objectonly" target="object" rid-ob="figobch06iplF6">Fig. 6. </a></h4><p class="float-caption no_bottom_margin">Stratification of amacrine cells in human retina </p></div></div><p>The above amacrine cells have been stained by the Golgi procedure which, as we have seen, is very good for showing the isolated complete shape and size of a nerve cell. Newer techniques are also providing valuable information on cell shapes and sizes: techniques such as using intracellular staining through a microelectrode and immunocytochemistry to stain cells with antibodies to their neurotransmitter or synthesising enzymes.<a href="http://webvision.med.utah.edu/imageswv/starburs.jpeg" ref="pagearea=body&targetsite=external&targetcat=link&targettype=uri">http://webvision.med.utah.edu/imageswv/starburs.jpeg</a></p><p>These are some different amacrine cell types stained by injection of dyes through a microelectrode inserted into the cell body. The cells that look like exploding fireworks, are called “starburst” cells (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F7/?report=objectonly" target="object" rid-figpopup="figch06iplF7" rid-ob="figobch06iplF7">Fig. 7</a>) and are known to use acetylcholine as their neurotransmitter (<a class="bk_pop" href="#ch06ipl.REF.famiglietti.1983.138">Famiglietti, 1983</a>; <a class="bk_pop" href="#ch06ipl.REF.masland.1988.405">Masland 1988</a>; <a class="bk_pop" href="#ch06ipl.REF.vaney.1990.49">Vaney, 1990</a>).</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF7" co-legend-rid="figlgndch06iplF7"><a href="/books/NBK11536/figure/ch06ipl.F7/?report=objectonly" target="object" title="Fig.7. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF7" rid-ob="figobch06iplF7"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image009.gif" src-large="/books/NBK11536/bin/ch06ipl-Image009.jpg" alt="Fig.7. . Starbust amacrine cellshttp://webvision." /></a><div class="icnblk_cntnt" id="figlgndch06iplF7"><h4 id="ch06ipl.F7"><a href="/books/NBK11536/figure/ch06ipl.F7/?report=objectonly" target="object" rid-ob="figobch06iplF7">Fig.7. </a></h4><p class="float-caption no_bottom_margin">Starbust amacrine cellshttp://webvision.med.utah.edu/imageswv/seroton.jpeg </p></div></div><p>Another type of amacrine cell, thought to contain GABA as a neurotransmitter, and can be well stained with serotonin antibodies has been stained by the same technique (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F8/?report=objectonly" target="object" rid-figpopup="figch06iplF8" rid-ob="figobch06iplF8">Fig. 8</a>) (<a class="bk_pop" href="#ch06ipl.REF.vaney.1986.444">Vaney, 1986</a>). This is the A17 cell of the rod system that makes reciprocal synapses at the rod bipolar ribbon (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F4b/?report=objectonly" target="object" rid-figpopup="figch06iplF4b" rid-ob="figobch06iplF4b">Fig. 4b</a>).</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF8" co-legend-rid="figlgndch06iplF8"><a href="/books/NBK11536/figure/ch06ipl.F8/?report=objectonly" target="object" title="Fig.8. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF8" rid-ob="figobch06iplF8"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image010.gif" src-large="/books/NBK11536/bin/ch06ipl-Image010.jpg" alt="Fig.8. " /></a><div class="icnblk_cntnt" id="figlgndch06iplF8"><h4 id="ch06ipl.F8"><a href="/books/NBK11536/figure/ch06ipl.F8/?report=objectonly" target="object" rid-ob="figobch06iplF8">Fig.8. </a></h4><p class="float-caption no_bottom_margin">Serotonin-containing amacrine cells </p></div></div><p>Using similar morphological techniques of Golgi staining, immunocytochemisry and intracellular staining, the ganglion cells of the retina have been classified into various different types.</p></div><div id="ch06ipl.sec4"><h2 id="_ch06ipl_sec4_">4. Different morphological types of ganglion cells.</h2><p>Using similar morphological techniques of Golgi staining, immunocytochemisry and intracellular staining, the ganglion cells of the retina have been classified into various different types.</p><p>As many as 25 different types exist in mammalian and human retinas, a few of which are shown in Fig.s 9 and 10, in both wholemount view and sectioned view. As with the amacrine cells, ganglion cells are classified onto the different types by cell body size, dendritic tree spread, branching patterns (e.g. radiate or tufted) and branching level in the 5 strata of the IPL. In both cat and human retinas ganglion cells can be from small to large, diffuse, bistratified and various monostratified large-field types with different stratification levels in the inner plexiform layer (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F9/?report=objectonly" target="object" rid-figpopup="figch06iplF9" rid-ob="figobch06iplF9">Figs. 9</a> and <a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F10/?report=objectonly" target="object" rid-figpopup="figch06iplF10" rid-ob="figobch06iplF10">10</a>).</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF9" co-legend-rid="figlgndch06iplF9"><a href="/books/NBK11536/figure/ch06ipl.F9/?report=objectonly" target="object" title="Fig. 9. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF9" rid-ob="figobch06iplF9"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image011.gif" src-large="/books/NBK11536/bin/ch06ipl-Image011.jpg" alt="Fig. 9. " /></a><div class="icnblk_cntnt" id="figlgndch06iplF9"><h4 id="ch06ipl.F9"><a href="/books/NBK11536/figure/ch06ipl.F9/?report=objectonly" target="object" rid-ob="figobch06iplF9">Fig. 9. </a></h4><p class="float-caption no_bottom_margin">Sectioned view of ganglion cells </p></div></div><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF10" co-legend-rid="figlgndch06iplF10"><a href="/books/NBK11536/figure/ch06ipl.F10/?report=objectonly" target="object" title="Fig. 10. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF10" rid-ob="figobch06iplF10"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image012.gif" src-large="/books/NBK11536/bin/ch06ipl-Image012.jpg" alt="Fig. 10. " /></a><div class="icnblk_cntnt" id="figlgndch06iplF10"><h4 id="ch06ipl.F10"><a href="/books/NBK11536/figure/ch06ipl.F10/?report=objectonly" target="object" rid-ob="figobch06iplF10">Fig. 10. </a></h4><p class="float-caption no_bottom_margin">Wholemount view of golgi stained ganglion cells in primate retina </p></div></div><p>Both amacrine and ganglion cells increase in dendritic tree span with eccentricity from the fovea. The very smallest dendritic fields for all cells types being possessed by cells of the fovea (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F9/?report=objectonly" target="object" rid-figpopup="figch06iplF9" rid-ob="figobch06iplF9">Fig. 9</a>) or area centralis (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F11/?report=objectonly" target="object" rid-figpopup="figch06iplF11" rid-ob="figobch06iplF11">Fig. 11</a>) for the common ganglion cells of the cat retina (alpha and beta types). The same ganglion cell types are ten times the size in dendritic tree spread in peripheral retina (<a class="bk_pop" href="#ch06ipl.REF.boycott.1974.397">Boycott and Wassle, 1974</a>; <a class="bk_pop" href="#ch06ipl.REF.kolb.1981.1081">Kolb et al., 1981</a>). For this reason classification of amacrines and ganglion cells has to always take eccentricity from the fovea into account and compare cells in similar areas of retina to be sure that they are indeed different cell types (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F11/?report=objectonly" target="object" rid-figpopup="figch06iplF11" rid-ob="figobch06iplF11">Fig. 11</a>).</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF11" co-legend-rid="figlgndch06iplF11"><a href="/books/NBK11536/figure/ch06ipl.F11/?report=objectonly" target="object" title="Fig. 11. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF11" rid-ob="figobch06iplF11"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image013.gif" src-large="/books/NBK11536/bin/ch06ipl-Image013.jpg" alt="Fig. 11. " /></a><div class="icnblk_cntnt" id="figlgndch06iplF11"><h4 id="ch06ipl.F11"><a href="/books/NBK11536/figure/ch06ipl.F11/?report=objectonly" target="object" rid-ob="figobch06iplF11">Fig. 11. </a></h4><p class="float-caption no_bottom_margin">Cat ganglion cells </p></div></div><p>In the human retina, the commonest ganglion cell types are the parasol and the midget ganglion cells (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F12/?report=objectonly" target="object" rid-figpopup="figch06iplF12" rid-ob="figobch06iplF12">Fig. 12</a>) (<a class="bk_pop" href="#ch06ipl.REF.polyak.1941">Polyak, 1941</a>; <a class="bk_pop" href="#ch06ipl.REF.rodieck.1985.115">Rodieck et al., 1985</a>; <a class="bk_pop" href="#ch06ipl.REF.kolb.1992.147">Kolb et al., 1992</a>; <a class="bk_pop" href="#ch06ipl.REF.dacey.1993.5334">Dacey, 1993</a>). They are also known as P cells (midget ganglion cells) because of their projection to the parvocellular layers, and M cells (parasol ganglion cells) because of their projection to the magnocellular layers of the lateral geniculate nucleus (Shapley and Perry, 1982). We shall return to these ganglion cell types in a later chapter.</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF12" co-legend-rid="figlgndch06iplF12"><a href="/books/NBK11536/figure/ch06ipl.F12/?report=objectonly" target="object" title="Fig. 12. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF12" rid-ob="figobch06iplF12"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image014.gif" src-large="/books/NBK11536/bin/ch06ipl-Image014.jpg" alt="Fig. 12. " /></a><div class="icnblk_cntnt" id="figlgndch06iplF12"><h4 id="ch06ipl.F12"><a href="/books/NBK11536/figure/ch06ipl.F12/?report=objectonly" target="object" rid-ob="figobch06iplF12">Fig. 12. </a></h4><p class="float-caption no_bottom_margin">Primate ganglion cells </p></div></div></div><div id="ch06ipl.sec5"><h2 id="_ch06ipl_sec5_">5. Stratification of amacrine and ganglion cells in relationship to bipolar cell axons.</h2><p>One may ask why amacrine cells and ganglion cells should be classified on stratification level of their dendrites, particularly if they look the same in other respects such as field size, cell body size and dendritic morphology. It was clear more than 100 years ago to Cajal (1894) that dendrites of amacrine and ganglion cells and axons of bipolar cells were arranged to costratify in some meaningful manner. The meaning of costratifications and orderly arrangements of branching levels, other than the obvious of making sure certain cell types made synapses with certain other cell types, escaped him though.<a href="http://webvision.med.utah.edu/imageswv/luciferelec.jpeg" ref="pagearea=body&targetsite=external&targetcat=link&targettype=uri">http://webvision.med.utah.edu/imageswv/luciferelec.jpeg</a></p><p>We had to wait for 70 years before the answer was found in the physiological responses of the different nerve cells of the retina. In the 1970s it became possible to make intracellular recordings from the neurons of the retina with dye filled glass microelectrodes. Physiologists were able to record both the response of the impaled cell to light and to fill the cell with a fluorescent dye such as lucifer to see what type of cell it was (see <a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F13/?report=objectonly" target="object" rid-figpopup="figch06iplF13" rid-ob="figobch06iplF13">Fig. 13</a>, left) (<a class="bk_pop" href="#ch06ipl.REF.kaneko.1970.623">Kaneko, 1970</a>; <a class="bk_pop" href="#ch06ipl.REF.nelson.1978.427">Nelson et al., 1978</a>).</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF13" co-legend-rid="figlgndch06iplF13"><a href="/books/NBK11536/figure/ch06ipl.F13/?report=objectonly" target="object" title="Fig.13. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF13" rid-ob="figobch06iplF13"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image015.gif" src-large="/books/NBK11536/bin/ch06ipl-Image015.jpg" alt="Fig.13. " /></a><div class="icnblk_cntnt" id="figlgndch06iplF13"><h4 id="ch06ipl.F13"><a href="/books/NBK11536/figure/ch06ipl.F13/?report=objectonly" target="object" rid-ob="figobch06iplF13">Fig.13. </a></h4><p class="float-caption no_bottom_margin">Experimental procedure for impaling ganglion cell in the retina </p></div></div><p>
|
|
<a href="/books/NBK11536/bin/electrode.mp4">CLICK HERE TO SEE AN ANIMATION OF THE IONTOPHORESIS OF LUCIFER DYE INTO A GANGLION CELL VIA A MICROLECTRODE (MP4 MOVIE)</a>
|
|
</p><p>
|
|
<a href="/books/NBK11536/bin/schema.mp4">CLICK HERE TO SEE AN ANIMATION OF THE PHYSIOLOGICAL RECORDING PROCEDURE (MP4 MOVIE)</a>
|
|
</p><p>In the fifties and sixties it became apparant from ganglion cell recordings in the mammalian retina that the visual message leaving the retina was in the form of ganglion cell axon spike discharges that occurred either when a spot of light stimulated the retina (ON discharge to light), or when the spot of light was turned off (OFF discharge). Ganglion cells were responding to either one or the other of the change of state of light. The one group was stimulated by the light brighter than background and the other by light darker than background (<a class="bk_pop" href="#ch06ipl.REF.kuffler.1953.37">Kuffler, 1953</a>; <a class="bk_pop" href="#ch06ipl.REF.enrothcugell.1966.517">Enroth-Cugell and Robson, 1966</a>). Next came intracellular recordings from the five basic cell types in a mudpuppy retina, which revealed that cells preceding ganglion cells also fell into the two physiological types i.e. bipolar and amacrine cells could give ON or OFF discharges (<a class="bk_pop" href="#ch06ipl.REF.werblin.1969.339">Werblin and Dowling, 1969</a>; <a class="bk_pop" href="#ch06ipl.REF.kaneko.1970.623">Kaneko, 1970</a>). Knowing now that bipolar cells could be ON- or OFF-center to a spot of light (they could have the opposite response in a surround concentrically around their central receptive field; mentioned in the chapter on the outer plexiform layer and dealt with in detail in later chapters on circuitry for rod and cone signals), it was reasonable to suggest that OFF-center bipolar chains excited OFF ganglion cells and that ON-center bipolar cell chains excited ON ganglion cells. The typical bipolar cell types that might form the bipolar chains driving ganglion cells of the primate retina are shown below.</p><p>These bipolar cells, as described in the previous section on the outer plexiform layer, make different types of synapses with cone pedicles and rod spherules. Some types make basal contacts on the surface of the cone pedicles and others make invaginating contact to end close to the synaptic ribbons in both rod spherules and cone pedicles.</p><p>It is now thought that bipolar cells respond to light just like the photoreceptor with a slow hyperpolarization when the bipolar dendrite is in the basal junction position. In contrast, the other type of contact, invaginating to the ribbon synapse, causes the bipolar to respond to light with an inverted sign compared with the photoreceptor. It gives a slow depolarizing response. Thus the nature of the postsynaptic membrane channels on the cone bipolar cell dendrite is the important designator of the response sign the bipolar will have. The hyperpolarizing bipolar types are the start of OFF-center channels and the depolarizing types are the start of ON-center channels through the retina.</p><p>The bipolar cells shown above (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F14/?report=objectonly" target="object" rid-figpopup="figch06iplF14" rid-ob="figobch06iplF14">Fig. 14</a>), will be noticed to have different axonal ending levels in the inner plexiform layer e.g. DB1 has an axon ending in stratum 1 neuropil close to the amacrine cell layer of the inner nucelar layer, while imb has an axon ending at the opposite side of the inner plexiform layer in straum 5 against the ganglion cell bodies. It seems obvious now, but it took a lot of painstaking serial section recontruction to show (<a class="bk_pop" href="#ch06ipl.REF.kolb.1979.295">Kolb, 1979</a>; <a class="bk_pop" href="#ch06ipl.REF.kolb.1991.617">Kolb and DeKorver, 1991</a>) that DB1 and imb made contacts only with ganglion cells that had dendritic stratification in the same neuropil as the respective bipolar axonal terminals. At the level of information transfer between bipolar cells and ganglion cells in the IPL, only excitatory channels are present (<a class="bk_pop" href="#ch06ipl.REF.raviola.1982.233">Raviola and Raviola, 1982</a>) so the type of signal transmitted to the ganglion cell, as either ON- or OFF-center, is essentially determined by the bipolar cells contacting it.</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF14" co-legend-rid="figlgndch06iplF14"><a href="/books/NBK11536/figure/ch06ipl.F14/?report=objectonly" target="object" title="Fig. 14. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF14" rid-ob="figobch06iplF14"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image016.gif" src-large="/books/NBK11536/bin/ch06ipl-Image016.jpg" alt="Fig. 14. " /></a><div class="icnblk_cntnt" id="figlgndch06iplF14"><h4 id="ch06ipl.F14"><a href="/books/NBK11536/figure/ch06ipl.F14/?report=objectonly" target="object" rid-ob="figobch06iplF14">Fig. 14. </a></h4><p class="float-caption no_bottom_margin">Bipolar cell types in primate retina </p></div></div><p>In order to keep the ON and OFF channels separate through the ganglion cells to the brain, the inner plexiform layer is divided into two functionally discrete sublaminae, called a (the 2 strata below the amacrine cell bodies) andb (the other 3 strata stretching to the ganglion cell bodies) (Famigletti and Kolb, 1976). Interactions are only allowed between basal-contacting cone bipolar types and one set of ganglion cells in sublamina a, while invaginating-contacting cone bipolar cells can only interact with another set of ganglion cells branching in sublamina b (see above). <a class="bk_pop" href="#ch06ipl.REF.gouras.1971.397">Gouras (1971)</a> was the first to suggest that this specificity of bipolar to ganglion cell contacts underlay ON-center and OFF-center midget ganglion cell responses in monkey.</p><p>
|
|
<a href="http://webvision.med.utah.edu/wp-content/uploads/2011/03/GourasIPL1.jpg" class="img_link" ref="pagearea=body&targetsite=external&targetcat=link&targettype=uri">
|
|
<span class="graphic"><img src="/books/NBK11536/bin/ch06ipl-Image001.jpg" alt="Image ch06ipl-Image001.jpg" /></span>
|
|
</a>
|
|
</p><p>Peter Gouras was the first to record from midget ganglion cells in the monkey retina and report on their spectrally opponent receptive field organization (1968). He then suggested in 1971 that specificity of biplar to ganglion cell contacts in the IPL underlay ON-center and OFF-center responses of midget ganglion cells in primate retina.</p><p>Later Nelson (<a class="bk_pop" href="#ch06ipl.REF.nelson.1978.427">Nelson, et al., 1978</a>) conclusively proved this hypothesis by means of intracellular recording and marking experiments in ganglion cells of cat (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F15/?report=objectonly" target="object" rid-figpopup="figch06iplF15" rid-ob="figobch06iplF15">Figs 15</a> annd 16).</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF15" co-legend-rid="figlgndch06iplF15"><a href="/books/NBK11536/figure/ch06ipl.F15/?report=objectonly" target="object" title="Fig. 15. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF15" rid-ob="figobch06iplF15"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image017.gif" src-large="/books/NBK11536/bin/ch06ipl-Image017.jpg" alt="Fig. 15. " /></a><div class="icnblk_cntnt" id="figlgndch06iplF15"><h4 id="ch06ipl.F15"><a href="/books/NBK11536/figure/ch06ipl.F15/?report=objectonly" target="object" rid-ob="figobch06iplF15">Fig. 15. </a></h4><p class="float-caption no_bottom_margin">Organization of ON- and OFF-center ganglion cells </p></div></div><p>
|
|
<a href="/books/NBK11536/bin/betaON-OFF.mov">CLICK HERE TO SEE AN ANIMATION OF THE PHYSIOLOGICAL RECORDING OF ON AND OFF BETA GANGLION CELLS (QUICKTIME MOVIE)</a>
|
|
</p><p><a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F16/?report=objectonly" target="object" rid-figpopup="figch06iplF16" rid-ob="figobch06iplF16">Fig. 16</a> shows the intracellular recordings made from two different ganglion cells of Nelson and coauthors’ study (1978). One cell proves to be ON-center, giving a burst of spikes riding on a depolarization of the membrane as soon as the light flash goes on (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F16/?report=objectonly" target="object" rid-figpopup="figch06iplF16" rid-ob="figobch06iplF16">Fig. 16</a>, left). In contrast, the other cell is OFF-center, giving a hyperpolarization to the membrane when the light flash is on, but a burst of spikes riding on the depolarization when the light flash is over (<a class="figpopup" href="/books/NBK11536/figure/ch06ipl.F16/?report=objectonly" target="object" rid-figpopup="figch06iplF16" rid-ob="figobch06iplF16">Fig. 16</a> right). The ON-center ganglion cell has dendrites restricted to branching in sublamina <b>b</b> and has synaptic input from invaginating types of bipolar cell. The OFF-center cell has dendrites reaching higher to branch only in sublamina <b>a</b> and gets synapses from basal junction contacting types of bipolar cell.</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch06iplF16" co-legend-rid="figlgndch06iplF16"><a href="/books/NBK11536/figure/ch06ipl.F16/?report=objectonly" target="object" title="Fig. 16. " class="img_link icnblk_img figpopup" rid-figpopup="figch06iplF16" rid-ob="figobch06iplF16"><img class="small-thumb" src="/books/NBK11536/bin/ch06ipl-Image018.gif" src-large="/books/NBK11536/bin/ch06ipl-Image018.jpg" alt="Fig. 16. " /></a><div class="icnblk_cntnt" id="figlgndch06iplF16"><h4 id="ch06ipl.F16"><a href="/books/NBK11536/figure/ch06ipl.F16/?report=objectonly" target="object" rid-ob="figobch06iplF16">Fig. 16. </a></h4><p class="float-caption no_bottom_margin">Intracellular recordings of ON-center and OFF-center ganglion cells </p></div></div><p>Thus it is, that there is a definite functional architecture to the inner plexiform layer of the mammalian retina and, in fact, to all vertebrate retinas. Stratification of the neuropil is formed by specific levels of branching of bipolar, amacrine and ganglion cells so that specialized circuits of interactions are set up. In addition there has, during the course of evolution, been an imposition of a broad division of the two halves of the inner plexiform layer into the top half allowing only interactions for the OFF-center ganglion cell pathways and the bottom half, only for the ON-center ganglion cell pathways. Occasionally ON center bipolars pass by ON ganglion cells (melanopsin or ipRGC cells) and amacrine cells (transient dopaminergic amacrine cells) and direct synapses to them thereby “breaking the rule” (<a class="bk_pop" href="#ch06ipl.REF.hoshi.2009.8875">Hoshi et al., 2009</a>). However on the whole, bipolar and ganglion cell are responsible for the bisublaminar ON and OFF channel organization in the IPL. Many amacrine cells are diffuse or bistratified and serve to connect up the ON- and OFF-center neuropils, while still others stratify in such a manner at the border between the two functional neuropils, to receive both ON and OFF inputs to drive them. Such amacrine cells may be the majority in the vertebrate retina and are thought to be involved more in temporal facets of retinal performance. They may be important for fast transfer of information, i.e. for speeding up signals. The roles of some of these amacrine cells will become clearer in later chapters.</p></div><div id="ch06ipl.sec5___1"><h2 id="_ch06ipl_sec5___1_">5. References.</h2><ol><li><div class="bk_ref" id="ch06ipl.REF.boycott.1974.397">Boycott, B.B. and Wassle, H. (1974) The morphological types of ganglion cells of the domestic cat’s retina.
|
|
J. Physiol. (Lond.), 240, 397-419.
|
|
[<a href="/pmc/articles/PMC1331022/" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pmc">PMC free article<span class="bk_prnt">: PMC1331022</span></a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/4422168" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 4422168</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.cajal.1892">Cajal, S.R. (1892) The Structure of the Retina. (Transl. Thorpe, S.A. and Glickstein, M.), Thomas, Springfield, Il., 1972.</div></li><li><div class="bk_ref" id="ch06ipl.REF.chavez.2010.2330">Chavez
|
|
AE, Grimes
|
|
WN, Diamond
|
|
JS (2010) Mechanisms underlying lateral GABAergic feedback onto rod bipolar cells in rat retina.
|
|
J Neurosci. 30, 2330-9.
|
|
[<a href="/pmc/articles/PMC2836865/" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pmc">PMC free article<span class="bk_prnt">: PMC2836865</span></a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/20147559" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 20147559</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.dacey.1993.5334">Dacey, D.M. (1993) The mosaic of midget ganglion cells in the human retina.
|
|
J. Neurosci. 13, 5334-5355.
|
|
[<a href="/pmc/articles/PMC6576399/" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pmc">PMC free article<span class="bk_prnt">: PMC6576399</span></a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/8254378" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 8254378</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.dowling.1966">Dowling, J.E. and Boycott, B.B. (1966) Organization of the primate retina: electron microscopy. Proc. R. Soc., B, (Lond.) <strong>166</strong>, 80-111. [<a href="https://pubmed.ncbi.nlm.nih.gov/4382694" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 4382694</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.enrothcugell.1966.517">Enroth-Cugell, C. and Robson, J.G. (1966) The contrast sensitivity of retinal ganglion cells of the cat.
|
|
J. Physiol. (Lond.), 187, 517-552.
|
|
[<a href="/pmc/articles/PMC1395960/" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pmc">PMC free article<span class="bk_prnt">: PMC1395960</span></a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/16783910" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 16783910</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.famiglietti.1983.138">Famiglietti, E.V. (1983) ‘Starburst’ amacrine cells and cholinergic neurons: mirror-symmetric ON and OFF amacrine cells of rabbit retina.
|
|
Brain Res. 261, 138-144.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/6301622" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 6301622</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.famiglietti.1976.193">Famiglietti, E.V. and Kolb, H. (1976) Structural basis for ON- and OFF-center responses in retinal ganglion cells.
|
|
Science, 194, 193-195.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/959847" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 959847</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.gouras.1971.397">Gouras, P. (1971) The function of the midget system in primate color vision.
|
|
Vision Res. (Suppl) 3, 397-410.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/5293881" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 5293881</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.grunert.2000.130">Grunert
|
|
U. (2000) Distribution of GABA and glycine receptors on bipolar and ganglion cells in the mammalian retina.
|
|
Microsc Res Tech.50, 130-40.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/10891877" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 10891877</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.grunert.2002.138">Grunert
|
|
U, Haverkamp
|
|
S, Fletcher
|
|
EL, Wassle
|
|
H. (2002) Synaptic distribution of ionotropic glutamate receptors in the inner plexiform layer of the primate retina.
|
|
J Comp Neurol. 447, 138-51.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/11977117" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 11977117</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.hoshi.2009.8875">Hoshi
|
|
H, Liu
|
|
W-L, Massey
|
|
SC and Mills
|
|
SL (2009) ON inputs to the OFF layer: Bipolar cells that break the stratification rules of the retina.
|
|
J Neurosci 29, 8875-8883.
|
|
[<a href="/pmc/articles/PMC2724754/" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pmc">PMC free article<span class="bk_prnt">: PMC2724754</span></a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/19605625" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 19605625</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.kaneko.1970.623">Kaneko, A. (1970) Physiological and morphological identification of horizontal, bipolar and amacrine cells in goldfish retina.
|
|
J. Physiol. (Lond), 207, 623-633.
|
|
[<a href="/pmc/articles/PMC1348731/" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pmc">PMC free article<span class="bk_prnt">: PMC1348731</span></a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/5499739" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 5499739</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.kolb.1979.295">Kolb, H. (1979) The inner plexiform layer in the retina of the cat: electron microscopic observations.
|
|
J. Neurocytol. 8, 295-329.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/490185" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 490185</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.kolb.1991.617">Kolb, H. and DeKorver, L. (1991) Midget ganglion cells of the parafovea of the human retina: A study by electron microscopy and serial section reconstructions.
|
|
J. Comp. Neurol., 303, 617-636.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/1707423" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 1707423</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.kolb.1992.147">Kolb, H., Linberg, K.A. and Fisher, S.K. (1992) The neurons of the human retina: a Golgi study.
|
|
J. Comp. Neurol. 318, 147-187.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/1374766" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 1374766</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.kolb.1981.1081">Kolb, H., Nelson, R. and Mariani, A. (1981) Amacrine cells, bipolar cells and ganglion cells of the cat retina: A Golgi study.
|
|
Vision Res. 21, 1081-1114.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/7314489" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 7314489</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.kneussel.2000.1">Kneussel
|
|
M, Betz
|
|
H. (2000) Receptors, gephyrin and gephyrin-associated proteins: novel insights into the assembly of inhibitory postsynaptic membrane specializations.
|
|
J Physiol.525,1-9.
|
|
[<a href="/pmc/articles/PMC2269938/" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pmc">PMC free article<span class="bk_prnt">: PMC2269938</span></a>] [<a href="https://pubmed.ncbi.nlm.nih.gov/10811719" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 10811719</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.kuffler.1953.37">Kuffler, S.W. (1953) Discharge patterns and functional organization of mammalian retina.
|
|
J. Neurophysiol. 16, 37-68.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/13035466" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 13035466</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.mariani.1990.382">Mariani, A.P. (1990) Amacrine cells of the rhesus monkey retina.
|
|
J. Comp. Neurol. 301, 382-400.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/2262597" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 2262597</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.masland.1988.405">Masland, R.H. (1988) Amacrine Cells.
|
|
Trends in Neuroscience 11, 405-410.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/2469207" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 2469207</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.nelson.1978.427">Nelson, R., Famiglietti, E.V. and Kolb, H.(1978) Intracellular staining reveals different levels of stratification for on-center and off-center ganglion cells in the cat retina.
|
|
J. Neurophysiol. 4, 427-483. [<a href="https://pubmed.ncbi.nlm.nih.gov/650277" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 650277</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.polyak.1941">Polyak, S.L. (1941) The Retina. The University of Chicago Press, Chicago, Ill.</div></li><li><div class="bk_ref" id="ch06ipl.REF.raviola.1982.233">Raviola, E. and Raviola, G. (1982) Structure of the synaptic membranes in the inner plexiform layer of the retina: A freeze-fracture study in monkeys and rabbits.
|
|
J. Comp. Neurol. 209, 233-248.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/7130454" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 7130454</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.rodieck.1985.115">Rodieck, R.W., Binmoeller, K.F. and Dineen, J. (1985) Parasol and midget ganglion cells of the human retina.
|
|
J. Comp. Neurol. 233, 115-132.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/3980768" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 3980768</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.shapley.1986.229">Shapley, R. and Perry, V.H. (1986) Cat and monkey retinal ganglion cells and their visual functional roles.
|
|
Trends in Neuroscience 9, 229-235.</div></li><li><div class="bk_ref" id="ch06ipl.REF.vaney.1986.444">Vaney, D.I. (1986) Morphological identification of serotonin-accumulating neurons in the living retina.
|
|
Science 233, 444-446.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/3726538" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 3726538</span></a>]</div></li><li><div class="bk_ref" id="ch06ipl.REF.vaney.1990.49">Vaney, D.I. (1990) The mosaic of amacrine cells in the mammalian retina.
|
|
Prog. Ret. Res. 9, 49-100.</div></li><li><div class="bk_ref" id="ch06ipl.REF.werblin.1969.339">Werblin, F.S. and Dowling, J.E. (1969) Organization of the retina of the mudpuppy, Necturus maculosus. II. Intracellular recording.
|
|
J. Neurophysiol. 32, 339-355.
|
|
[<a href="https://pubmed.ncbi.nlm.nih.gov/4306897" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 4306897</span></a>]</div></li></ol></div><div id="bk_toc_contnr"></div></div></div>
|
|
<div class="post-content"><div><div class="half_rhythm"><a href="/books/about/copyright/">Copyright</a>: © 2025 Webvision .<p class="small">All copyright for chapters belongs to the individual authors who created them. However, for non-commercial, academic purposes, images and content from the chapters portion of Webvision may be used with a non-exclusive rights under a Attribution, <a href="https://creativecommons.org/licenses/by-nc/4.0/" ref="pagearea=meta&targetsite=external&targetcat=link&targettype=uri">Noncommercial 4.0 International (CC BY-NC) Creative Commons license</a>. Cite Webvision, http://webvision.med.utah.edu/ as the source. Commercial applications need to obtain license permission from the administrator of Webvision and are generally declined unless the copyright owner can/wants to donate or license material. Use online should be accompanied by a link back to the original source of the material. All imagery or content associated with blog posts belong to the authors of said posts, except where otherwise noted.</p></div><div class="small"><span class="label">Bookshelf ID: NBK11536</span><span class="label">PMID: <a href="https://pubmed.ncbi.nlm.nih.gov/21413394" title="PubMed record of this page" ref="pagearea=meta&targetsite=entrez&targetcat=link&targettype=pubmed">21413394</a></span></div><div style="margin-top:2em" class="bk_noprnt"><a class="bk_cntns" href="/books/n/webvision/">Contents</a><div class="pagination bk_noprnt"><a class="active page_link prev" href="/books/n/webvision/A207/" title="Previous page in this title">< Prev</a><a class="active page_link next" href="/books/n/webvision/ch07gc1/" title="Next page in this title">Next ></a></div></div></div></div>
|
|
|
|
</div>
|
|
|
|
<!-- Custom content below content -->
|
|
<div class="col4">
|
|
|
|
</div>
|
|
|
|
|
|
<!-- Book content -->
|
|
|
|
<!-- Custom contetnt below bottom nav -->
|
|
<div class="col5">
|
|
|
|
</div>
|
|
</div>
|
|
|
|
<div id="rightcolumn" class="four_col col last">
|
|
<!-- Custom content above discovery portlets -->
|
|
<div class="col6">
|
|
<div id="ncbi_share_book"><a href="#" class="ncbi_share" data-ncbi_share_config="popup:false,shorten:true" ref="id=NBK11536&db=books">Share</a></div>
|
|
|
|
</div>
|
|
<div xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance"></div><div class="portlet"><div class="portlet_head"><div class="portlet_title"><h3><span>Views</span></h3></div><a name="Shutter" sid="1" href="#" class="portlet_shutter" title="Show/hide content" remembercollapsed="true" pgsec_name="PDF_download" id="Shutter"></a></div><div class="portlet_content"><ul xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="simple-list"><li><a href="/books/NBK11536/?report=reader">PubReader</a></li><li><a href="/books/NBK11536/?report=printable">Print View</a></li><li><a data-jig="ncbidialog" href="#_ncbi_dlg_citbx_NBK11536" data-jigconfig="width:400,modal:true">Cite this Page</a><div id="_ncbi_dlg_citbx_NBK11536" style="display:none" title="Cite this Page"><div class="bk_tt">Kolb H. Inner Plexiform Layer. [Updated 2012 Mar 25]. In: Kolb H, Fernandez E, Jones B, et al., editors. Webvision: The Organization of the Retina and Visual System [Internet]. Salt Lake City (UT): University of Utah Health Sciences Center; 1995-. <span class="bk_cite_avail"></span></div></div></li><li><a href="/books/NBK11536/pdf/Bookshelf_NBK11536.pdf">PDF version of this page</a> (2.0M)</li><li><a href="/books/n/webvision/pdf/">PDF version of this title</a> (235M)</li></ul></div></div><div class="portlet"><div class="portlet_head"><div class="portlet_title"><h3><span>In this Page</span></h3></div><a name="Shutter" sid="1" href="#" class="portlet_shutter" title="Show/hide content" remembercollapsed="true" pgsec_name="page-toc" id="Shutter"></a></div><div class="portlet_content"><ul xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="simple-list"><li><a href="#ch06ipl.sec1" ref="log$=inpage&link_id=inpage">Bipolar, amacrine and ganglion cells interact in the inner plexiform layer.</a></li><li><a href="#ch06ipl.sec2" ref="log$=inpage&link_id=inpage">Ultrastructure of the neuropil of the inner plexiform layer.</a></li><li><a href="#ch06ipl.sec3" ref="log$=inpage&link_id=inpage">Different morphological types of amacrine and ganglion cells.</a></li><li><a href="#ch06ipl.sec4" ref="log$=inpage&link_id=inpage">Different morphological types of ganglion cells.</a></li><li><a href="#ch06ipl.sec5" ref="log$=inpage&link_id=inpage">Stratification of amacrine and ganglion cells in relationship to bipolar cell axons.</a></li><li><a href="#ch06ipl.sec5___1" ref="log$=inpage&link_id=inpage">References.</a></li></ul></div></div><div class="portlet"><div class="portlet_head"><div class="portlet_title"><h3><span>Related Items in Bookshelf</span></h3></div><a name="Shutter" sid="1" href="#" class="portlet_shutter" title="Show/hide content" remembercollapsed="true" pgsec_name="source-links" id="Shutter"></a></div><div class="portlet_content"><ul xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="simple-list"><li><a href="https://www.ncbi.nlm.nih.gov/books?term=%22reference%20works%22%5BResource%20Type%5D" ref="pagearea=source-links&targetsite=external&targetcat=link&targettype=uri">All Reference Works</a></li><li><a href="https://www.ncbi.nlm.nih.gov/books?term="textbooks"%5BResource%20Type%5D" ref="pagearea=source-links&targetsite=external&targetcat=link&targettype=uri">All Textbooks</a></li></ul></div></div><div class="portlet"><div class="portlet_head"><div class="portlet_title"><h3><span>Related information</span></h3></div><a name="Shutter" sid="1" href="#" class="portlet_shutter" title="Show/hide content" remembercollapsed="true" pgsec_name="discovery_db_links" id="Shutter"></a></div><div class="portlet_content"><ul><li class="brieflinkpopper"><a class="brieflinkpopperctrl" href="/books/?Db=pmc&DbFrom=books&Cmd=Link&LinkName=books_pmc_refs&IdsFromResult=1652408" ref="log$=recordlinks">PMC</a><div class="brieflinkpop offscreen_noflow">PubMed Central citations</div></li><li class="brieflinkpopper"><a class="brieflinkpopperctrl" href="/books/?Db=pubmed&DbFrom=books&Cmd=Link&LinkName=books_pubmed_refs&IdsFromResult=1652408" ref="log$=recordlinks">PubMed</a><div class="brieflinkpop offscreen_noflow">Links to PubMed</div></li></ul></div></div><div class="portlet"><div class="portlet_head"><div class="portlet_title"><h3><span>Similar articles in PubMed</span></h3></div><a name="Shutter" sid="1" href="#" class="portlet_shutter" title="Show/hide content" remembercollapsed="true" pgsec_name="PBooksDiscovery_RA" id="Shutter"></a></div><div class="portlet_content"><ul><li class="brieflinkpopper two_line"><a class="brieflinkpopperctrl" href="/pubmed/39139018" ref="ordinalpos=1&linkpos=1&log$=relatedarticles&logdbfrom=pubmed">The thickness of the retinal nerve fiber layer, macula, and ganglion cell-inner plexiform layer in people with drug-resistant epilepsy.</a><span class="source">[Epilepsia Open. 2024]</span><div class="brieflinkpop offscreen_noflow">The thickness of the retinal nerve fiber layer, macula, and ganglion cell-inner plexiform layer in people with drug-resistant epilepsy.<div class="brieflinkpopdesc"><em xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="author">Chen Y, Xiong W, Lu L, Wu X, Cao L, Chen J, Xiao Y, Sander JW, Wu B, Zhou D. </em><em xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="cit">Epilepsia Open. 2024 Oct; 9(5):1783-1792. Epub 2024 Aug 14.</em></div></div></li><li class="brieflinkpopper two_line"><a class="brieflinkpopperctrl" href="/pubmed/2487094" ref="ordinalpos=1&linkpos=2&log$=relatedarticles&logdbfrom=pubmed">Localization of GABA- and GAD-like immunoreactivity in the turtle retina.</a><span class="source">[Vis Neurosci. 1989]</span><div class="brieflinkpop offscreen_noflow">Localization of GABA- and GAD-like immunoreactivity in the turtle retina.<div class="brieflinkpopdesc"><em xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="author">Hurd LB 2nd, Eldred WD. </em><em xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="cit">Vis Neurosci. 1989 Jul; 3(1):9-20. </em></div></div></li><li class="brieflinkpopper two_line"><a class="brieflinkpopperctrl" href="/pubmed/1674751" ref="ordinalpos=1&linkpos=3&log$=relatedarticles&logdbfrom=pubmed">Dopaminergic interplexiform cells and centrifugal fibres in the Xenopus retina.</a><span class="source">[J Neurocytol. 1991]</span><div class="brieflinkpop offscreen_noflow">Dopaminergic interplexiform cells and centrifugal fibres in the Xenopus retina.<div class="brieflinkpopdesc"><em xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="author">Schütte M, Witkovsky P. </em><em xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="cit">J Neurocytol. 1991 Mar; 20(3):195-207. </em></div></div></li><li class="brieflinkpopper two_line"><a class="brieflinkpopperctrl" href="/pubmed/33953540" ref="ordinalpos=1&linkpos=4&log$=relatedarticles&logdbfrom=pubmed">Relationship Between Central Corneal Thickness and Ganglionic-Inner Plexiform Cell Layer and Retinal Nerve Fibre Layer Thickness in Normal Subjects.</a><span class="source">[Clin Ophthalmol. 2021]</span><div class="brieflinkpop offscreen_noflow">Relationship Between Central Corneal Thickness and Ganglionic-Inner Plexiform Cell Layer and Retinal Nerve Fibre Layer Thickness in Normal Subjects.<div class="brieflinkpopdesc"><em xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="author">Al Saad M, Shehadeh A, Meqbil J, Elubous K, Qablawi M, Abu-Yaghi N. </em><em xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="cit">Clin Ophthalmol. 2021; 15:1809-1812. Epub 2021 Apr 28.</em></div></div></li><li class="brieflinkpopper two_line"><a class="brieflinkpopperctrl" href="/pubmed/9302106" ref="ordinalpos=1&linkpos=5&log$=relatedreviews&logdbfrom=pubmed"><span xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="invert">Review</span> A Golgi study of the short-axon interneurons of the cell layer and inner plexiform layer of the medial cortex of the lizard Podarcis hispanica.</a><span class="source">[J Comp Neurol. 1997]</span><div class="brieflinkpop offscreen_noflow"><span xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="invert">Review</span> A Golgi study of the short-axon interneurons of the cell layer and inner plexiform layer of the medial cortex of the lizard Podarcis hispanica.<div class="brieflinkpopdesc"><em xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="author">Luis de la Iglesia JA, Lopez-Garcia C. </em><em xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="cit">J Comp Neurol. 1997 Sep 8; 385(4):565-98. </em></div></div></li></ul><a class="seemore" href="/sites/entrez?db=pubmed&cmd=link&linkname=pubmed_pubmed_reviews&uid=21413394" ref="ordinalpos=1&log$=relatedreviews_seeall&logdbfrom=pubmed">See reviews...</a><a class="seemore" href="/sites/entrez?db=pubmed&cmd=link&linkname=pubmed_pubmed&uid=21413394" ref="ordinalpos=1&log$=relatedarticles_seeall&logdbfrom=pubmed">See all...</a></div></div><div class="portlet"><div class="portlet_head"><div class="portlet_title"><h3><span>Recent Activity</span></h3></div><a name="Shutter" sid="1" href="#" class="portlet_shutter" title="Show/hide content" remembercollapsed="true" pgsec_name="recent_activity" id="Shutter"></a></div><div class="portlet_content"><div xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" id="HTDisplay" class=""><div class="action"><a href="javascript:historyDisplayState('ClearHT')">Clear</a><a href="javascript:historyDisplayState('HTOff')" class="HTOn">Turn Off</a><a href="javascript:historyDisplayState('HTOn')" class="HTOff">Turn On</a></div><ul id="activity"><li class="ra_rcd ralinkpopper two_line"><a class="htb ralinkpopperctrl" ref="log$=activity&linkpos=1" href="/portal/utils/pageresolver.fcgi?recordid=67c9a72fb15b832ebc78169b">Inner Plexiform Layer - Webvision</a><div class="ralinkpop offscreen_noflow">Inner Plexiform Layer - Webvision<div class="brieflinkpopdesc"></div></div><div class="tertiary"></div></li><li class="ra_rcd ralinkpopper two_line"><a class="htb ralinkpopperctrl" ref="log$=activity&linkpos=2" href="/portal/utils/pageresolver.fcgi?recordid=67c9a72eb15b832ebc780be0">OUTER PLEXIFORM LAYER - Webvision</a><div class="ralinkpop offscreen_noflow">OUTER PLEXIFORM LAYER - Webvision<div class="brieflinkpopdesc"></div></div><div class="tertiary"></div></li><li class="ra_rcd ralinkpopper two_line"><a class="htb ralinkpopperctrl" ref="log$=activity&linkpos=3" href="/portal/utils/pageresolver.fcgi?recordid=67c9a72d84f3725e59a59bdc">Photoreceptors - Webvision</a><div class="ralinkpop offscreen_noflow">Photoreceptors - Webvision<div class="brieflinkpopdesc"></div></div><div class="tertiary"></div></li><li class="ra_rcd ralinkpopper two_line"><a class="htb ralinkpopperctrl" ref="log$=activity&linkpos=4" href="/portal/utils/pageresolver.fcgi?recordid=67c9a72bf4a390645ea2f7c8">Part II: Anatomy and Physiology of the retina - Webvision</a><div class="ralinkpop offscreen_noflow">Part II: Anatomy and Physiology of the retina - Webvision<div class="brieflinkpopdesc"></div></div><div class="tertiary"></div></li><li class="ra_rcd ralinkpopper two_line"><a class="htb ralinkpopperctrl" ref="log$=activity&linkpos=5" href="/portal/utils/pageresolver.fcgi?recordid=67c9a72ba68b6b5afcdd9fb4">The Retinal Pigment Epithelium - Webvision</a><div class="ralinkpop offscreen_noflow">The Retinal Pigment Epithelium - Webvision<div class="brieflinkpopdesc"></div></div><div class="tertiary"></div></li></ul><p class="HTOn">Your browsing activity is empty.</p><p class="HTOff">Activity recording is turned off.</p><p id="turnOn" class="HTOff"><a href="javascript:historyDisplayState('HTOn')">Turn recording back on</a></p><a class="seemore" href="/sites/myncbi/recentactivity">See more...</a></div></div></div>
|
|
|
|
<!-- Custom content below discovery portlets -->
|
|
<div class="col7">
|
|
|
|
</div>
|
|
</div>
|
|
</div>
|
|
|
|
<!-- Custom content after all -->
|
|
<div class="col8">
|
|
|
|
</div>
|
|
<div class="col9">
|
|
|
|
</div>
|
|
|
|
<script type="text/javascript" src="/corehtml/pmc/js/jquery.scrollTo-1.4.2.js"></script>
|
|
<script type="text/javascript">
|
|
(function($){
|
|
$('.skiplink').each(function(i, item){
|
|
var href = $($(item).attr('href'));
|
|
href.attr('tabindex', '-1').addClass('skiptarget'); // ensure the target can receive focus
|
|
$(item).on('click', function(event){
|
|
event.preventDefault();
|
|
$.scrollTo(href, 0, {
|
|
onAfter: function(){
|
|
href.focus();
|
|
}
|
|
});
|
|
});
|
|
});
|
|
})(jQuery);
|
|
</script>
|
|
</div>
|
|
<div class="bottom">
|
|
|
|
<div id="NCBIFooter_dynamic">
|
|
<!--<component id="Breadcrumbs" label="breadcrumbs"/>
|
|
<component id="Breadcrumbs" label="helpdesk"/>-->
|
|
|
|
</div>
|
|
|
|
<div class="footer" id="footer">
|
|
<section class="icon-section">
|
|
<div id="icon-section-header" class="icon-section_header">Follow NCBI</div>
|
|
<div class="grid-container container">
|
|
<div class="icon-section_container">
|
|
<a class="footer-icon" id="footer_twitter" href="https://twitter.com/ncbi" aria-label="Twitter"><svg xmlns="http://www.w3.org/2000/svg" data-name="Layer 1" viewBox="0 0 300 300">
|
|
<defs>
|
|
<style>
|
|
.cls-11 {
|
|
fill: #737373;
|
|
}
|
|
</style>
|
|
</defs>
|
|
<title>Twitter</title>
|
|
<path class="cls-11" d="M250.11,105.48c-7,3.14-13,3.25-19.27.14,8.12-4.86,8.49-8.27,11.43-17.46a78.8,78.8,0,0,1-25,9.55,39.35,39.35,0,0,0-67,35.85,111.6,111.6,0,0,1-81-41.08A39.37,39.37,0,0,0,81.47,145a39.08,39.08,0,0,1-17.8-4.92c0,.17,0,.33,0,.5a39.32,39.32,0,0,0,31.53,38.54,39.26,39.26,0,0,1-17.75.68,39.37,39.37,0,0,0,36.72,27.3A79.07,79.07,0,0,1,56,223.34,111.31,111.31,0,0,0,116.22,241c72.3,0,111.83-59.9,111.83-111.84,0-1.71,0-3.4-.1-5.09C235.62,118.54,244.84,113.37,250.11,105.48Z">
|
|
</path>
|
|
</svg></a>
|
|
<a class="footer-icon" id="footer_facebook" href="https://www.facebook.com/ncbi.nlm" aria-label="Facebook"><svg xmlns="http://www.w3.org/2000/svg" data-name="Layer 1" viewBox="0 0 300 300">
|
|
<title>Facebook</title>
|
|
<path class="cls-11" d="M210.5,115.12H171.74V97.82c0-8.14,5.39-10,9.19-10h27.14V52l-39.32-.12c-35.66,0-42.42,26.68-42.42,43.77v19.48H99.09v36.32h27.24v109h45.41v-109h35Z">
|
|
</path>
|
|
</svg></a>
|
|
<a class="footer-icon" id="footer_linkedin" href="https://www.linkedin.com/company/ncbinlm" aria-label="LinkedIn"><svg xmlns="http://www.w3.org/2000/svg" data-name="Layer 1" viewBox="0 0 300 300">
|
|
<title>LinkedIn</title>
|
|
<path class="cls-11" d="M101.64,243.37H57.79v-114h43.85Zm-22-131.54h-.26c-13.25,0-21.82-10.36-21.82-21.76,0-11.65,8.84-21.15,22.33-21.15S101.7,78.72,102,90.38C102,101.77,93.4,111.83,79.63,111.83Zm100.93,52.61A17.54,17.54,0,0,0,163,182v61.39H119.18s.51-105.23,0-114H163v13a54.33,54.33,0,0,1,34.54-12.66c26,0,44.39,18.8,44.39,55.29v58.35H198.1V182A17.54,17.54,0,0,0,180.56,164.44Z">
|
|
</path>
|
|
</svg></a>
|
|
<a class="footer-icon" id="footer_github" href="https://github.com/ncbi" aria-label="GitHub"><svg xmlns="http://www.w3.org/2000/svg" data-name="Layer 1" viewBox="0 0 300 300">
|
|
<defs>
|
|
<style>
|
|
.cls-11,
|
|
.cls-12 {
|
|
fill: #737373;
|
|
}
|
|
|
|
.cls-11 {
|
|
fill-rule: evenodd;
|
|
}
|
|
</style>
|
|
</defs>
|
|
<title>GitHub</title>
|
|
<path class="cls-11" d="M151.36,47.28a105.76,105.76,0,0,0-33.43,206.1c5.28,1,7.22-2.3,7.22-5.09,0-2.52-.09-10.85-.14-19.69-29.42,6.4-35.63-12.48-35.63-12.48-4.81-12.22-11.74-15.47-11.74-15.47-9.59-6.56.73-6.43.73-6.43,10.61.75,16.21,10.9,16.21,10.9,9.43,16.17,24.73,11.49,30.77,8.79,1-6.83,3.69-11.5,6.71-14.14C108.57,197.1,83.88,188,83.88,147.51a40.92,40.92,0,0,1,10.9-28.39c-1.1-2.66-4.72-13.42,1-28,0,0,8.88-2.84,29.09,10.84a100.26,100.26,0,0,1,53,0C198,88.3,206.9,91.14,206.9,91.14c5.76,14.56,2.14,25.32,1,28a40.87,40.87,0,0,1,10.89,28.39c0,40.62-24.74,49.56-48.29,52.18,3.79,3.28,7.17,9.71,7.17,19.58,0,14.15-.12,25.54-.12,29,0,2.82,1.9,6.11,7.26,5.07A105.76,105.76,0,0,0,151.36,47.28Z">
|
|
</path>
|
|
<path class="cls-12" d="M85.66,199.12c-.23.52-1.06.68-1.81.32s-1.2-1.06-.95-1.59,1.06-.69,1.82-.33,1.21,1.07.94,1.6Zm-1.3-1">
|
|
</path>
|
|
<path class="cls-12" d="M90,203.89c-.51.47-1.49.25-2.16-.49a1.61,1.61,0,0,1-.31-2.19c.52-.47,1.47-.25,2.17.49s.82,1.72.3,2.19Zm-1-1.08">
|
|
</path>
|
|
<path class="cls-12" d="M94.12,210c-.65.46-1.71,0-2.37-.91s-.64-2.07,0-2.52,1.7,0,2.36.89.65,2.08,0,2.54Zm0,0"></path>
|
|
<path class="cls-12" d="M99.83,215.87c-.58.64-1.82.47-2.72-.41s-1.18-2.06-.6-2.7,1.83-.46,2.74.41,1.2,2.07.58,2.7Zm0,0">
|
|
</path>
|
|
<path class="cls-12" d="M107.71,219.29c-.26.82-1.45,1.2-2.64.85s-2-1.34-1.74-2.17,1.44-1.23,2.65-.85,2,1.32,1.73,2.17Zm0,0">
|
|
</path>
|
|
<path class="cls-12" d="M116.36,219.92c0,.87-1,1.59-2.24,1.61s-2.29-.68-2.3-1.54,1-1.59,2.26-1.61,2.28.67,2.28,1.54Zm0,0">
|
|
</path>
|
|
<path class="cls-12" d="M124.42,218.55c.15.85-.73,1.72-2,1.95s-2.37-.3-2.52-1.14.73-1.75,2-2,2.37.29,2.53,1.16Zm0,0"></path>
|
|
</svg></a>
|
|
<a class="footer-icon" id="footer_blog" href="https://ncbiinsights.ncbi.nlm.nih.gov/" aria-label="Blog">
|
|
<svg xmlns="http://www.w3.org/2000/svg" id="Layer_1" data-name="Layer 1" viewBox="0 0 40 40">
|
|
<defs><style>.cls-1{fill:#737373;}</style></defs>
|
|
<title>NCBI Insights Blog</title>
|
|
<path class="cls-1" d="M14,30a4,4,0,1,1-4-4,4,4,0,0,1,4,4Zm11,3A19,19,0,0,0,7.05,15a1,1,0,0,0-1,1v3a1,1,0,0,0,.93,1A14,14,0,0,1,20,33.07,1,1,0,0,0,21,34h3a1,1,0,0,0,1-1Zm9,0A28,28,0,0,0,7,6,1,1,0,0,0,6,7v3a1,1,0,0,0,1,1A23,23,0,0,1,29,33a1,1,0,0,0,1,1h3A1,1,0,0,0,34,33Z"></path>
|
|
</svg>
|
|
</a>
|
|
</div>
|
|
</div>
|
|
</section>
|
|
|
|
<section class="container-fluid bg-primary">
|
|
<div class="container pt-5">
|
|
<div class="row mt-3">
|
|
<div class="col-lg-3 col-12">
|
|
<p><a class="text-white" href="https://www.nlm.nih.gov/socialmedia/index.html">Connect with NLM</a></p>
|
|
<ul class="list-inline social_media">
|
|
<li class="list-inline-item"><a href="https://twitter.com/NLM_NIH" aria-label="Twitter" target="_blank" rel="noopener noreferrer"><svg xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink" version="1.1" x="0px" y="0px" viewBox="0 0 249 249" style="enable-background:new 0 0 249 249;" xml:space="preserve">
|
|
<style type="text/css">
|
|
.st20 {
|
|
fill: #FFFFFF;
|
|
}
|
|
|
|
.st30 {
|
|
fill: none;
|
|
stroke: #FFFFFF;
|
|
stroke-width: 8;
|
|
stroke-miterlimit: 10;
|
|
}
|
|
</style>
|
|
<title>Twitter</title>
|
|
<g>
|
|
<g>
|
|
<g>
|
|
<path class="st20" d="M192.9,88.1c-5,2.2-9.2,2.3-13.6,0.1c5.7-3.4,6-5.8,8.1-12.3c-5.4,3.2-11.4,5.5-17.6,6.7 c-10.5-11.2-28.1-11.7-39.2-1.2c-7.2,6.8-10.2,16.9-8,26.5c-22.3-1.1-43.1-11.7-57.2-29C58,91.6,61.8,107.9,74,116 c-4.4-0.1-8.7-1.3-12.6-3.4c0,0.1,0,0.2,0,0.4c0,13.2,9.3,24.6,22.3,27.2c-4.1,1.1-8.4,1.3-12.5,0.5c3.6,11.3,14,19,25.9,19.3 c-11.6,9.1-26.4,13.2-41.1,11.5c12.7,8.1,27.4,12.5,42.5,12.5c51,0,78.9-42.2,78.9-78.9c0-1.2,0-2.4-0.1-3.6 C182.7,97.4,189.2,93.7,192.9,88.1z"></path>
|
|
</g>
|
|
</g>
|
|
<circle class="st30" cx="124.4" cy="128.8" r="108.2"></circle>
|
|
</g>
|
|
</svg></a></li>
|
|
<li class="list-inline-item"><a href="https://www.facebook.com/nationallibraryofmedicine" aria-label="Facebook" rel="noopener noreferrer" target="_blank">
|
|
<svg xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink" version="1.1" x="0px" y="0px" viewBox="0 0 249 249" style="enable-background:new 0 0 249 249;" xml:space="preserve">
|
|
<style type="text/css">
|
|
.st10 {
|
|
fill: #FFFFFF;
|
|
}
|
|
|
|
.st110 {
|
|
fill: none;
|
|
stroke: #FFFFFF;
|
|
stroke-width: 8;
|
|
stroke-miterlimit: 10;
|
|
}
|
|
</style>
|
|
<title>Facebook</title>
|
|
<g>
|
|
<g>
|
|
<path class="st10" d="M159,99.1h-24V88.4c0-5,3.3-6.2,5.7-6.2h16.8V60l-24.4-0.1c-22.1,0-26.2,16.5-26.2,27.1v12.1H90v22.5h16.9 v67.5H135v-67.5h21.7L159,99.1z"></path>
|
|
</g>
|
|
</g>
|
|
<circle class="st110" cx="123.6" cy="123.2" r="108.2"></circle>
|
|
</svg>
|
|
</a></li>
|
|
<li class="list-inline-item"><a href="https://www.youtube.com/user/NLMNIH" aria-label="Youtube" target="_blank" rel="noopener noreferrer"><svg xmlns="http://www.w3.org/2000/svg" xmlns:xlink="http://www.w3.org/1999/xlink" version="1.1" x="0px" y="0px" viewBox="0 0 249 249" style="enable-background:new 0 0 249 249;" xml:space="preserve">
|
|
<title>Youtube</title>
|
|
<style type="text/css">
|
|
.st4 {
|
|
fill: none;
|
|
stroke: #FFFFFF;
|
|
stroke-width: 8;
|
|
stroke-miterlimit: 10;
|
|
}
|
|
|
|
.st5 {
|
|
fill: #FFFFFF;
|
|
}
|
|
</style>
|
|
<circle class="st4" cx="124.2" cy="123.4" r="108.2"></circle>
|
|
<g transform="translate(0,-952.36218)">
|
|
<path class="st5" d="M88.4,1037.4c-10.4,0-18.7,8.3-18.7,18.7v40.1c0,10.4,8.3,18.7,18.7,18.7h72.1c10.4,0,18.7-8.3,18.7-18.7 v-40.1c0-10.4-8.3-18.7-18.7-18.7H88.4z M115.2,1058.8l29.4,17.4l-29.4,17.4V1058.8z"></path>
|
|
</g>
|
|
</svg></a></li>
|
|
</ul>
|
|
</div>
|
|
<div class="col-lg-3 col-12">
|
|
<p class="address_footer text-white">National Library of Medicine<br />
|
|
<a href="https://www.google.com/maps/place/8600+Rockville+Pike,+Bethesda,+MD+20894/@38.9959508,-77.101021,17z/data=!3m1!4b1!4m5!3m4!1s0x89b7c95e25765ddb:0x19156f88b27635b8!8m2!3d38.9959508!4d-77.0988323" class="text-white" target="_blank" rel="noopener noreferrer">8600 Rockville Pike<br />
|
|
Bethesda, MD 20894</a></p>
|
|
</div>
|
|
<div class="col-lg-3 col-12 centered-lg">
|
|
<p><a href="https://www.nlm.nih.gov/web_policies.html" class="text-white">Web Policies</a><br />
|
|
<a href="https://www.nih.gov/institutes-nih/nih-office-director/office-communications-public-liaison/freedom-information-act-office" class="text-white">FOIA</a><br />
|
|
<a href="https://www.hhs.gov/vulnerability-disclosure-policy/index.html" class="text-white" id="vdp">HHS Vulnerability Disclosure</a></p>
|
|
</div>
|
|
<div class="col-lg-3 col-12 centered-lg">
|
|
<p><a class="supportLink text-white" href="https://support.nlm.nih.gov/">Help</a><br />
|
|
<a href="https://www.nlm.nih.gov/accessibility.html" class="text-white">Accessibility</a><br />
|
|
<a href="https://www.nlm.nih.gov/careers/careers.html" class="text-white">Careers</a></p>
|
|
</div>
|
|
</div>
|
|
<div class="row">
|
|
<div class="col-lg-12 centered-lg">
|
|
<nav class="bottom-links">
|
|
<ul class="mt-3">
|
|
<li>
|
|
<a class="text-white" href="//www.nlm.nih.gov/">NLM</a>
|
|
</li>
|
|
<li>
|
|
<a class="text-white" href="https://www.nih.gov/">NIH</a>
|
|
</li>
|
|
<li>
|
|
<a class="text-white" href="https://www.hhs.gov/">HHS</a>
|
|
</li>
|
|
<li>
|
|
<a class="text-white" href="https://www.usa.gov/">USA.gov</a>
|
|
</li>
|
|
</ul>
|
|
</nav>
|
|
</div>
|
|
</div>
|
|
</div>
|
|
</section>
|
|
<script type="text/javascript" src="/portal/portal3rc.fcgi/rlib/js/InstrumentOmnitureBaseJS/InstrumentNCBIConfigJS/InstrumentNCBIBaseJS/InstrumentPageStarterJS.js?v=1"> </script>
|
|
<script type="text/javascript" src="/portal/portal3rc.fcgi/static/js/hfjs2.js"> </script>
|
|
</div>
|
|
</div>
|
|
</div>
|
|
<!--/.page-->
|
|
</div>
|
|
<!--/.wrap-->
|
|
</div><!-- /.twelve_col -->
|
|
</div>
|
|
<!-- /.grid -->
|
|
|
|
<span class="PAFAppResources"></span>
|
|
|
|
<!-- BESelector tab -->
|
|
|
|
|
|
|
|
<noscript><img alt="statistics" src="/stat?jsdisabled=true&ncbi_db=books&ncbi_pdid=book-part&ncbi_acc=NBK11536&ncbi_domain=webvision&ncbi_report=record&ncbi_type=fulltext&ncbi_objectid=&ncbi_pcid=/NBK11536/&ncbi_pagename=Inner Plexiform Layer - Webvision - NCBI Bookshelf&ncbi_bookparttype=chapter&ncbi_app=bookshelf" /></noscript>
|
|
|
|
|
|
<!-- usually for JS scripts at page bottom -->
|
|
<!--<component id="PageFixtures" label="styles"></component>-->
|
|
|
|
|
|
<!-- CE8B5AF87C7FFCB1_0191SID /projects/books/PBooks@9.11 portal105 v4.1.r689238 Tue, Oct 22 2024 16:10:51 -->
|
|
<span id="portal-csrf-token" style="display:none" data-token="CE8B5AF87C7FFCB1_0191SID"></span>
|
|
|
|
<script type="text/javascript" src="//static.pubmed.gov/portal/portal3rc.fcgi/4216699/js/3879255/4121861/3501987/4008961/3893018/3821238/4062932/4209313/4212053/4076480/3921943/3400083/3426610.js" snapshot="books"></script></body>
|
|
</html> |