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<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="#__NBK11516_dtls__">Show details</a><div style="display:none" class="ui-widget" id="__NBK11516_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/ch08Amelan/" title="Previous page in this title">< Prev</a><a class="active page_link next" href="/books/n/webvision/Archetecture_Fovea/" title="Next page in this title">Next ></a></div></div></div></div></div>
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<div class="main-content lit-style" itemscope="itemscope" itemtype="http://schema.org/CreativeWork"><div class="meta-content fm-sec"><h1 id="_NBK11516_"><span class="title" itemprop="name">Glial cells of the Retina by Helga Kolb</span></h1><p class="contrib-group"><span itemprop="author">Helga Kolb</span>, PhD.</p><a data-jig="ncbitoggler" href="#__NBK11516_ai__" style="border:0;text-decoration:none">Author Information and Affiliations</a><div style="display:none" class="ui-widget" id="__NBK11516_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></div><div class="jig-ncbiinpagenav body-content whole_rhythm" data-jigconfig="allHeadingLevels: ['h2'],smoothScroll: false" itemprop="text"><p>Three basic types of glial cell are found in the human retina, Muller cells, astroglia and microglia. All were described for the retina by Cajal more than one hundred years ago (1892).</p><div id="ch09glia.sec1"><h2 id="_ch09glia_sec1_">1. Muller cells.</h2><p>Muller cells are the principal glial cell of the retina. They form architectural support structures stretching radially across the thickness of the retina and are the limits of the retina at the outer and inner limiting membrane respectively. A complete understanding of the shape of a Muller cell is best seen after Golgi staining as shown originally by Cajal (1892) below.</p><p>Muller cell bodies sit in the inner nuclear layer and project irregularly thick and thin processes in either direction to the outer limiting membrane and to the inner limiting membrane. Muller cell processes insinuate themselves between cell bodies of the neurons in the nuclear layers and envelope groups of neural processes in the plexiform layers (<a class="figpopup" href="/books/NBK11516/figure/ch09glia.F1/?report=objectonly" target="object" rid-figpopup="figch09gliaF1" rid-ob="figobch09gliaF1">Fig. 1</a>). In fact retinal neural processes are only allowed direct contact, without enveloping Muller cell processes, at their synapses.</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch09gliaF1" co-legend-rid="figlgndch09gliaF1"><a href="/books/NBK11516/figure/ch09glia.F1/?report=objectonly" target="object" title="Fig. 1. " class="img_link icnblk_img figpopup" rid-figpopup="figch09gliaF1" rid-ob="figobch09gliaF1"><img class="small-thumb" src="/books/NBK11516/bin/ch09glia-Image004.gif" src-large="/books/NBK11516/bin/ch09glia-Image004.jpg" alt="Fig. 1. " /></a><div class="icnblk_cntnt" id="figlgndch09gliaF1"><h4 id="ch09glia.F1"><a href="/books/NBK11516/figure/ch09glia.F1/?report=objectonly" target="object" rid-ob="figobch09gliaF1">Fig. 1. </a></h4><p class="float-caption no_bottom_margin">Golgi stained Muller cells </p></div></div><p>A single progenitor cell gives rise to both Muller cells and retinal neurons(<a class="bk_pop" href="#ch09glia.REF.turner.1987.131">Turner and Cepko, 1987</a>) although apparently in two phases. The earliest phase neurons born at the apical margin of the neuroepithelium adjacent to the pigment epithelium produces primary neurons consisting of cone cells, horizontal cells and ganglion cells (<a class="figpopup" href="/books/NBK11516/figure/ch09glia.F2/?report=objectonly" target="object" rid-figpopup="figch09gliaF2" rid-ob="figobch09gliaF2">Fig. 2</a>, right). The second phase of cells also born at the apical margins produces Muller cells and rod photoreceptors, bipolar cells and amacrine cells (<a class="bk_pop" href="#ch09glia.REF.reichenbach.1995">Reichenbach and Robinson, 1995</a>) (<a class="figpopup" href="/books/NBK11516/figure/ch09glia.F2/?report=objectonly" target="object" rid-figpopup="figch09gliaF2" rid-ob="figobch09gliaF2">Fig. 2</a>, left). All the developing neurons and the Muller cells have to migrate inward to their final position and it is thought that the Muller cell processes and trunks guide much of the neuron migrations and direct the neurite differentiations.</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch09gliaF2" co-legend-rid="figlgndch09gliaF2"><a href="/books/NBK11516/figure/ch09glia.F2/?report=objectonly" target="object" title="Fig. 2. " class="img_link icnblk_img figpopup" rid-figpopup="figch09gliaF2" rid-ob="figobch09gliaF2"><img class="small-thumb" src="/books/NBK11516/bin/ch09glia-Image005.gif" src-large="/books/NBK11516/bin/ch09glia-Image005.jpg" alt="Fig. 2. " /></a><div class="icnblk_cntnt" id="figlgndch09gliaF2"><h4 id="ch09glia.F2"><a href="/books/NBK11516/figure/ch09glia.F2/?report=objectonly" target="object" rid-ob="figobch09gliaF2">Fig. 2. </a></h4><p class="float-caption no_bottom_margin">3-D schematic drawing of the relationship between Muller cell and other retinal neurons </p></div></div><p>The junctions forming the outer limiting membrane are between Muller cells and other Muller cells and photoreceptor cells as sturdy desmosomes or zonula adherens. In some species gap junctions (specialized membrane associations and channels that allow passage of small molecules and ions) or tight junctions are part of these Muller cell junctions (<a class="bk_pop" href="#ch09glia.REF.miller.1970.323">Miller and Dowling, 1970</a>) but not so in mammalian species where no dye coupling has ever been observed (<a class="bk_pop" href="#ch09glia.REF.robinson.1993.167">Robinson et al., 1993</a>; <a class="bk_pop" href="#ch09glia.REF.reichenbach.1995">Reichenbach and Robinson, 1995</a>). The surface of the Muller cell facing the pigment epithelium and subretinal space is expanded by many projections of the Muller cell membrane known as apical villi. The inner limiting membrane, on the other hand, is formed by the conical endfeet of the Muller cell but no specialized junctions are seen here. Muller cells also form endfeet on the large retinal blood vessels at the inner surface of the retina. The surface of the Muller cell membrane facing the vitreous is covered with a mucopolysaccharide material and thus forms a true basement membrane.</p><p>Muller cells contain glycogen, mitochondria and intermediate filaments which are immunoreative for vimentin and to some extent to glial fibrillary acidicprotein (GFAP). The latter filaments are normally in the inner half of the retinal Muller cells and their endfeet (<a class="figpopup" href="/books/NBK11516/figure/ch09glia.F3/?report=objectonly" target="object" rid-figpopup="figch09gliaF3" rid-ob="figobch09gliaF3">Fig. 3</a>, left), but following trauma to the retina such as retinal detachment, both vimentin and GFAP are massively upregulated and found throughout the cell (<a class="figpopup" href="/books/NBK11516/figure/ch09glia.F3/?report=objectonly" target="object" rid-figpopup="figch09gliaF3" rid-ob="figobch09gliaF3">Fig. 3</a>, right) (<a class="bk_pop" href="#ch09glia.REF.guerin.1990">Guerin et al., 1990</a>; <a class="bk_pop" href="#ch09glia.REF.fisher.1995">Fisher and Lewis, 1995</a>).</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch09gliaF3" co-legend-rid="figlgndch09gliaF3"><a href="/books/NBK11516/figure/ch09glia.F3/?report=objectonly" target="object" title="Fig. 3. " class="img_link icnblk_img figpopup" rid-figpopup="figch09gliaF3" rid-ob="figobch09gliaF3"><img class="small-thumb" src="/books/NBK11516/bin/ch09glia-Image006.gif" src-large="/books/NBK11516/bin/ch09glia-Image006.jpg" alt="Fig. 3. " /></a><div class="icnblk_cntnt" id="figlgndch09gliaF3"><h4 id="ch09glia.F3"><a href="/books/NBK11516/figure/ch09glia.F3/?report=objectonly" target="object" rid-ob="figobch09gliaF3">Fig. 3. </a></h4><p class="float-caption no_bottom_margin">GFAP immunoreactivity in Muller cells </p></div></div><p>Muller cells have a range of functions all of which are vital to the health of the retinal neurons. Muller cells function in a symbiotic relationship with the neurons (for an excellent review see <a class="bk_pop" href="#ch09glia.REF.reichenbach.1995">Reichenbach and Robinson, 1995</a>). Thus Muller cell functions include:</p><dl class="temp-labeled-list"><dt>1.</dt><dd><p class="no_top_margin">Supplying endproducts of anaerobic metabolism (breakdown of glycogen) to fuel aerobic metabolism in the nerve cells.</p></dd><dt>2.</dt><dd><p class="no_top_margin">They mop up neural waste products such as carbon dioxide and ammonia and recycle spent amino acid transmitters.</p></dd><dt>3.</dt><dd><p class="no_top_margin">They protect neurons from exposure to excess neurotransmitters such as glutamate using well developed uptake mechanisms to recycle this transmitter. They are particularly characterized by the presence of high concentrations of glutamine synthase.</p></dd><dt>4.</dt><dd><p class="no_top_margin">They may be involved in both phagocytosis of neuronal debris and release of neuroactive substances such as GABA, taurine and dopamine.</p></dd><dt>5.</dt><dd><p class="no_top_margin">They are thought to synthesize retinoic acid from retinol (retinoic acid is known to be important in in the development of the eye and the nervous system) (<a class="bk_pop" href="#ch09glia.REF.edwards.1994.231">Edwards, 1994</a>)</p></dd><dt>6.</dt><dd><p class="no_top_margin">They control homeostasis and protect neurons from deleterious changes in their ionic environment by taking up extracellular K+ and redistributing it.</p></dd><dt>7.</dt><dd><p class="no_top_margin">They contribute to the generation of the electroretinogram (ERG) b-wave (<a class="bk_pop" href="#ch09glia.REF.miller.1970.323">Miller and Dowling, 1970</a>; <a class="bk_pop" href="#ch09glia.REF.newman.1984.164">Newman and Odette, 1984</a>), the slow P3 component of the ERG (<a class="bk_pop" href="#ch09glia.REF.karwoski.1977.244">Karwoski and Proenza, 1977</a>) and the scotopic threshold response (STR) (<a class="bk_pop" href="#ch09glia.REF.frishman.1989.1233">Frishman and Steinberg, 1989</a>). They do so by regulation of K+ distribution across the retinal vitreous border, across the whole retina and locally in the inner plexiform layer of the retina (<a class="figpopup" href="/books/NBK11516/figure/ch09glia.F4/?report=objectonly" target="object" rid-figpopup="figch09gliaF4" rid-ob="figobch09gliaF4">Fig. 4</a>, from <a class="bk_pop" href="#ch09glia.REF.reichenbach.1995">Reichenbach and Robinson, 1995</a>, adapted from <a class="bk_pop" href="#ch09glia.REF.newman.1989.153">Newman, 1989</a>).</p></dd></dl><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch09gliaF4" co-legend-rid="figlgndch09gliaF4"><a href="/books/NBK11516/figure/ch09glia.F4/?report=objectonly" target="object" title="Fig. 4. " class="img_link icnblk_img figpopup" rid-figpopup="figch09gliaF4" rid-ob="figobch09gliaF4"><img class="small-thumb" src="/books/NBK11516/bin/ch09glia-Image007.gif" src-large="/books/NBK11516/bin/ch09glia-Image007.jpg" alt="Fig. 4. " /></a><div class="icnblk_cntnt" id="figlgndch09gliaF4"><h4 id="ch09glia.F4"><a href="/books/NBK11516/figure/ch09glia.F4/?report=objectonly" target="object" rid-ob="figobch09gliaF4">Fig. 4. </a></h4><p class="float-caption no_bottom_margin">Regulation of K+ by Muller cells in Muller cells </p></div></div></div><div id="ch09glia.sec2"><h2 id="_ch09glia_sec2_">2. Astrocytes.</h2><p>Astrocytes are not glial cells of the retinal neuroepithelium but enter the developing retina from the brain along the developing optic nerve (<a class="bk_pop" href="#ch09glia.REF.stone.1987.35">Stone and Dreher, 1987</a>; <a class="bk_pop" href="#ch09glia.REF.chanling.1994.357">Chan-Ling 1994</a>). They have a characterisic morphology of a flattened cell body and a fibrous series of radiating processes. Intermediate filaments fill their processes and thus they stain dramatically with antibodies against GFAP (<a class="bk_pop" href="#ch09glia.REF.schnitzer.1988.209">Schnitzer, 1988</a>). Astrocyte cell bodies and processes are almost entirely restricted to the nerve fiber layer of the retina. Their morphology changes from the periphery to the optic nerve head: from a symmetrical stellate form in peripheral retina (<a class="figpopup" href="/books/NBK11516/figure/ch09glia.F5a/?report=objectonly" target="object" rid-figpopup="figch09gliaF5a" rid-ob="figobch09gliaF5a">Figs. 5a and b</a>) (Schitzer, 1988) to extremely elongated near the optic nerve (<a class="figpopup" href="/books/NBK11516/figure/ch09glia.F6/?report=objectonly" target="object" rid-figpopup="figch09gliaF6" rid-ob="figobch09gliaF6">Fig. 6</a> and <a class="figpopup" href="/books/NBK11516/figure/ch09glia.F7/?report=objectonly" target="object" rid-figpopup="figch09gliaF7" rid-ob="figobch09gliaF7">7</a>).</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch09gliaF5a" co-legend-rid="figlgndch09gliaF5a"><a href="/books/NBK11516/figure/ch09glia.F5a/?report=objectonly" target="object" title="Fig. 5a. " class="img_link icnblk_img figpopup" rid-figpopup="figch09gliaF5a" rid-ob="figobch09gliaF5a"><img class="small-thumb" src="/books/NBK11516/bin/ch09glia-Image008.gif" src-large="/books/NBK11516/bin/ch09glia-Image008.jpg" alt="Fig. 5a. . Astrocytes in peripheral retina." /></a><div class="icnblk_cntnt" id="figlgndch09gliaF5a"><h4 id="ch09glia.F5a"><a href="/books/NBK11516/figure/ch09glia.F5a/?report=objectonly" target="object" rid-ob="figobch09gliaF5a">Fig. 5a. </a></h4><p class="float-caption no_bottom_margin">Astrocytes in peripheral retina. Schnitzer, 1988 </p></div></div><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch09gliaF6" co-legend-rid="figlgndch09gliaF6"><a href="/books/NBK11516/figure/ch09glia.F6/?report=objectonly" target="object" title="Fig. 6. " class="img_link icnblk_img figpopup" rid-figpopup="figch09gliaF6" rid-ob="figobch09gliaF6"><img class="small-thumb" src="/books/NBK11516/bin/ch09glia-Image010.gif" src-large="/books/NBK11516/bin/ch09glia-Image010.jpg" alt="Fig. 6. . Astrocytes in central retina." /></a><div class="icnblk_cntnt" id="figlgndch09gliaF6"><h4 id="ch09glia.F6"><a href="/books/NBK11516/figure/ch09glia.F6/?report=objectonly" target="object" rid-ob="figobch09gliaF6">Fig. 6. </a></h4><p class="float-caption no_bottom_margin">Astrocytes in central retina. Schnitzer, 1988 </p></div></div><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch09gliaF7" co-legend-rid="figlgndch09gliaF7"><a href="/books/NBK11516/figure/ch09glia.F7/?report=objectonly" target="object" title="Fig. 7. " class="img_link icnblk_img figpopup" rid-figpopup="figch09gliaF7" rid-ob="figobch09gliaF7"><img class="small-thumb" src="/books/NBK11516/bin/ch09glia-Image011.gif" src-large="/books/NBK11516/bin/ch09glia-Image011.jpg" alt="Fig. 7. . Astrocytes in cat retina stained by intracellular injection of HRP." /></a><div class="icnblk_cntnt" id="figlgndch09gliaF7"><h4 id="ch09glia.F7"><a href="/books/NBK11516/figure/ch09glia.F7/?report=objectonly" target="object" rid-ob="figobch09gliaF7">Fig. 7. </a></h4><p class="float-caption no_bottom_margin">Astrocytes in cat retina stained by intracellular injection of HRP. Courtesy of Ralph Nelson. </p></div></div><p>In immunocytochemical staining (<a class="figpopup" href="/books/NBK11516/figure/ch09glia.F5b/?report=objectonly" target="object" rid-figpopup="figch09gliaF5b" rid-ob="figobch09gliaF5b">Fig. 5b</a>) and in HRP intracellular injections (<a class="figpopup" href="/books/NBK11516/figure/ch09glia.F7/?report=objectonly" target="object" rid-figpopup="figch09gliaF7" rid-ob="figobch09gliaF7">Fig. 7</a>) stained astrocytes clearly exhibit processes aligned along the ganglion cell axons coursing through the nerve fibre layer. In distribution, astrocytes reach their peak on the optic nerve head and have a fairly uniform decline in density in radiating rings from the nerve head. They are not present in the avascular fovea or ora serrata.</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch09gliaF5b" co-legend-rid="figlgndch09gliaF5b"><a href="/books/NBK11516/figure/ch09glia.F5b/?report=objectonly" target="object" title="Fig. 5b. " class="img_link icnblk_img figpopup" rid-figpopup="figch09gliaF5b" rid-ob="figobch09gliaF5b"><img class="small-thumb" src="/books/NBK11516/bin/ch09glia-Image009.gif" src-large="/books/NBK11516/bin/ch09glia-Image009.jpg" alt="Fig. 5b. . Astrocytes (red) in rat retina among ganglion cells (blue) and associated with blood vessels (green)." /></a><div class="icnblk_cntnt" id="figlgndch09gliaF5b"><h4 id="ch09glia.F5b"><a href="/books/NBK11516/figure/ch09glia.F5b/?report=objectonly" target="object" rid-ob="figobch09gliaF5b">Fig. 5b. </a></h4><p class="float-caption no_bottom_margin">Astrocytes (red) in rat retina among ganglion cells (blue) and associated with blood vessels (green). Courtesy of Laura Fernandez-Sanchez and Nicolas Cuenca </p></div></div><p>Thick and thin astrocytes have been distinguished on the basis of staining with antibodies to GFAP (Trevino et al., 1996). Thus astrocytes are arranged over the surface of the ganglion cell axon bundles as they course into the optic nerve head forming a tube through which the axons run (<a class="figpopup" href="/books/NBK11516/figure/ch09glia.F8/?report=objectonly" target="object" rid-figpopup="figch09gliaF8" rid-ob="figobch09gliaF8">Fig. 8</a>). Gap junctions and zonula adherens junctions have been described between astrocytic processes in cat retina (Höllander et al., 1991).</p><div class="iconblock whole_rhythm clearfix ten_col fig" id="figch09gliaF8" co-legend-rid="figlgndch09gliaF8"><a href="/books/NBK11516/figure/ch09glia.F8/?report=objectonly" target="object" title="Fig. 8. " class="img_link icnblk_img figpopup" rid-figpopup="figch09gliaF8" rid-ob="figobch09gliaF8"><img class="small-thumb" src="/books/NBK11516/bin/ch09glia-Image012.gif" src-large="/books/NBK11516/bin/ch09glia-Image012.jpg" alt="Fig. 8. " /></a><div class="icnblk_cntnt" id="figlgndch09gliaF8"><h4 id="ch09glia.F8"><a href="/books/NBK11516/figure/ch09glia.F8/?report=objectonly" target="object" rid-ob="figobch09gliaF8">Fig. 8. </a></h4><p class="float-caption no_bottom_margin">3-D block of astrocytes arranged over the surface of ganglion cell axon bundles </p></div></div><p>The blood vessels running in and among the ganglion cell bundles are also covered by by both processes and even an occasional cell body of an astrocyte. The function of astrocytes enveloping ganglion cell axons and the relationship to blood vessels of the nerve fibre layer suggests they are axonal and vascular glial sheaths and part of a blood-brain barrier. Similar to Muller cells, they are known to contain abundant glycogen and they may form a nutritive service in providing glucose to the neurons. In addition they probably serve a role in ionic homeostasis in regulating extracellular potassium levels and metabolism of neurotransmitters like GABA.</p></div><div id="ch09glia.sec3"><h2 id="_ch09glia_sec3_">3. Microglial cells.</h2><p>The third glial cell type is supposedly of mesodermal origin and thus, strictly speaking are not neuroglial as are the astrocytes and Muller cells. They enter the retina coincident with the mesenchymal precursors of retinal blood vessels in development (<a class="bk_pop" href="#ch09glia.REF.chanling.1994.357">Chan-Ling, 1994</a>). Microglial cells are ubiquitous in the human retina being found in every layer of the retina.</p><p>In Golgi-stained retina they look like strange, multipolar forms with small cell bodies and irregular short processes. In fact, in Golgi preparations they have sometimes been mistaken for nerve cells particularly when they lie in a nuclear layer with a single orientation of their processes in the plexiform layer.</p><div id="ch09glia.F9" class="figure bk_fig"><div class="graphic"><img src="/books/NBK11516/bin/ch09glia-Image001.jpg" alt="Fig. 9. " /></div><h3><span class="label">Fig. 9. </span></h3><div class="caption"><p>Golgi staining of microglial cells</p></div></div><p>Microglial cells may be of two types. One form is thought to enter the retina at early stages of development from the optic nerve mesenchyme and lie dormant in the retinal layers for much of the life of the retina. The other form of microglia appear to be blood-borne cells, possible originating from vessel pericytes (<a class="bk_pop" href="#ch09glia.REF.boycott.1981.679">Boycott and Hopkins, 1981</a>; <a class="bk_pop" href="#ch09glia.REF.gallego.1986.165">Gallego, 1986</a>). Both types can be stimulated into a macrophagic function after trauma to the retina, and then they engage in phagocytosis of degenerating retinal neurons.</p><div id="ch09glia.F10a" class="figure bk_fig"><div class="graphic"><img src="/books/NBK11516/bin/ch09glia-Image002.jpg" alt="Fig. 10a. . Lectin-stained microglial cell from Chan-Lin, 1994." /></div><h3><span class="label">Fig. 10a. </span></h3><div class="caption"><p>Lectin-stained microglial cell from Chan-Lin, 1994.</p></div></div><div id="ch09glia.F10b" class="figure bk_fig"><div class="graphic"><img src="/books/NBK11516/bin/ch09glia-Image003.jpg" alt="Fig. 10b. . Human retina with Astrocytes and Microglia." /></div><h3><span class="label">Fig. 10b. </span></h3><div class="caption"><p>Human retina with Astrocytes and Microglia. This image from Scott McLeod from Jerry Lutty’s lab, is a wholemount human retina preparation triple laneled with fluorescent antibodies that stain blood vessels (blue), astrocytes (red) and microglia (green).</p></div></div></div><div id="ch09glia.sec4"><h2 id="_ch09glia_sec4_">4. References.</h2><ol><li><div class="bk_ref" id="ch09glia.REF.boycott.1981.679">Boycott
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BB, Hopkins
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JM. Microglia in the retina of monkey and other mammals; its distinction from other types of glia and horizontal cells. Neuroscience.1981;6:679–688. []
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[<a href="https://pubmed.ncbi.nlm.nih.gov/6165924" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 6165924</span></a>]</div></li><li><div class="bk_ref" id="ch09glia.REF.cajal.1972">Cajal SR. In: Thorpe SA, Glickstein M, translators. 1892. The structure of the retina. Springfield (IL): Thomas; 1972.</div></li><li><div class="bk_ref" id="ch09glia.REF.chanling.1994.357">Chan-Ling
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Glial, neuronal and vascular interactions in the mammalian retina. Prog. Ret. Eye Res. 1994;13:357–389.</div></li><li><div class="bk_ref" id="ch09glia.REF.edwards.1994.231">Edwards
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RB. Biosynthesis of retinoic acid by Müller glial cells: a model for the central nervous system? Prog. Ret. Eye Res. 1994;13:231–242.</div></li><li><div class="bk_ref" id="ch09glia.REF.fisher.1995">Fisher SK, Lewis GP. Photoreceptors and beyond: cellular and molecular effects of retinal detachment.2nd Great Basin Visual Science Symposium, II, University of Utah Press. 1995</div></li><li><div class="bk_ref" id="ch09glia.REF.frishman.1989.1233">Frishman
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LJ, Steinberg
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RH. Light-evoked increases in [K+]o in proximal portion of the dark-adapted cat retina. J Neurophysiol. 1989;61:1233–1243. []
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Comparative studies on horizontal cells and a note on microglial cells. Prog. Ret. Res. 1986;5:165–206.</div></li><li><div class="bk_ref" id="ch09glia.REF.guerin.1990">Guerin CJ, Anderson DH, Fisher SK. Changes in intermediate filament immunolabeling occur in response to retinal detachment and reattachment in primates.Invest. Ophthal. Vis. Sci. 1990;31:1474–1482. [] [<a href="https://pubmed.ncbi.nlm.nih.gov/2387680" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 2387680</span></a>]</div></li><li><div class="bk_ref" id="ch09glia.REF.karwoski.1977.244">Karwoski
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LM. Relationship between Muller cell responses, a local transretinal potential, and potassium flux. J Neurophysiol. 1977;40:244–259.[]
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RF, Dowling
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JE. Intracellular responses of the Muller (glial) cells of the mudpuppy retina: their relation to b-wave of the electroretinogram. J Neurophysiol.1970;33:323–341. []
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LL. Model of electroretinogram b-wave generation: a test of the K+ hypothesis. J Neurophysiol. 1984;51:164–182. []
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EA. Electrophysiology of retinal glial cells. Prog. Ret. Res. 1989;8:153–172.</div></li><li><div class="bk_ref" id="ch09glia.REF.reichenbach.1995">Reichenbach A, Robinson SR. The involvement of Müller cells in the outer retina. In: Djamgoz MBA, Archer SN, Vallerga S, editors. Neurobiology and clinical aspects of the outer retina. London: Chapman & Hall; 1995. p. 395-416.</div></li><li><div class="bk_ref" id="ch09glia.REF.robinson.1993.167">Robinson
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DI. Unidirectional coupling of gap junctions between neuroglia. Proc. Austr. Neurosci. Soc. 1993;3:167.
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[<a href="https://pubmed.ncbi.nlm.nih.gov/8093125" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 8093125</span></a>]</div></li><li><div class="bk_ref" id="ch09glia.REF.schnitzer.1988.209">Schnitzer
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Astrocytes in mammalian retina. Prog. Ret. Res. 1988;7:209–232.</div></li><li><div class="bk_ref" id="ch09glia.REF.stone">Stone J, Makarov F, Hollander H. The glial ensheathment of the soma and axon hillock of retinal ganglion cells. Vis Neurosci. 12:273–279. [] [<a href="https://pubmed.ncbi.nlm.nih.gov/7786848" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 7786848</span></a>]</div></li><li><div class="bk_ref" id="ch09glia.REF.stone.1987.35">Stone
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J, Dreher
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Z. Relationship between astrocytes, ganglion cells and vasculature of the retina. J Comp Neurol. 1987;255:35–49. []
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[<a href="https://pubmed.ncbi.nlm.nih.gov/3819008" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 3819008</span></a>]</div></li><li><div class="bk_ref" id="ch09glia.REF.trivino.1996.2015">Trivino
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[<a href="https://pubmed.ncbi.nlm.nih.gov/8776468" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 8776468</span></a>]</div></li><li><div class="bk_ref" id="ch09glia.REF.turner.1987.131">Turner
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DL, Cepko
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CL. A common progenitor for neurons and glia persists in rat retina late in development. Nature. 1987;328:131–136. []
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[<a href="https://pubmed.ncbi.nlm.nih.gov/3600789" ref="pagearea=cite-ref&targetsite=entrez&targetcat=link&targettype=pubmed">PubMed<span class="bk_prnt">: 3600789</span></a>]</div></li></ol></div><div id="bk_toc_contnr"></div></div></div>
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<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: NBK11516</span><span class="label">PMID: <a href="https://pubmed.ncbi.nlm.nih.gov/21413380" title="PubMed record of this page" ref="pagearea=meta&targetsite=entrez&targetcat=link&targettype=pubmed">21413380</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/ch08Amelan/" title="Previous page in this title">< Prev</a><a class="active page_link next" href="/books/n/webvision/Archetecture_Fovea/" title="Next page in this title">Next ></a></div></div></div></div>
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<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/NBK11516/?report=reader">PubReader</a></li><li><a href="/books/NBK11516/?report=printable">Print View</a></li><li><a data-jig="ncbidialog" href="#_ncbi_dlg_citbx_NBK11516" data-jigconfig="width:400,modal:true">Cite this Page</a><div id="_ncbi_dlg_citbx_NBK11516" style="display:none" title="Cite this Page"><div class="bk_tt">Kolb H. Glial cells of the Retina by Helga Kolb. 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/NBK11516/pdf/Bookshelf_NBK11516.pdf">PDF version of this page</a> (3.3M)</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="#ch09glia.sec1" ref="log$=inpage&link_id=inpage">Muller cells.</a></li><li><a href="#ch09glia.sec2" ref="log$=inpage&link_id=inpage">Astrocytes.</a></li><li><a href="#ch09glia.sec3" ref="log$=inpage&link_id=inpage">Microglial cells.</a></li><li><a href="#ch09glia.sec4" ref="log$=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 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ref="ordinalpos=1&linkpos=1&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> Neurotransmitters in the Retina by Helga Kolb.</a><span class="source">[Webvision: The Organization of...]</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> Neurotransmitters in the Retina by Helga Kolb.<div class="brieflinkpopdesc"><em xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="author">Kolb H. </em><em xmlns:np="http://ncbi.gov/portal/XSLT/namespace" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" class="cit">Webvision: The Organization of the Retina and Visual System. 1995</em></div></div></li><li class="brieflinkpopper two_line"><a class="brieflinkpopperctrl" 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