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What is a C. elegans?

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<p><em><strong>Caenorhabditis elegans</strong></em> (pronounced <span title="Pronunciation in the International Phonetic Alphabet (IPA)" class="IPA"><a href="/wiki/Wikipedia:IPA_for_English" title="Wikipedia:IPA for English">/ˌsiːnoʊr&aelig;bˈdaɪt<s>ɪ</s>s ˈɛl<s>ɪ</s>g&aelig;nz/</a></span>) is a free-living, transparent <a href="/wiki/Nematode" title="Nematode">nematode</a> (roundworm), about 1&nbsp;mm in length, which lives in temperate soil environments. Research into the <a href="/wiki/Molecular_biology" title="Molecular biology">molecular</a> and <a href="/wiki/Developmental_biology" title="Developmental biology">developmental biology</a> of <em>C. elegans</em> was begun in 1974 by <a href="/wiki/Sydney_Brenner" title="Sydney Brenner">Sydney Brenner</a> and it has since been used extensively as a <a href="/wiki/Model_organism" title="Model organism">model organism</a>.<sup id="cite_ref-0" class="reference"><a href="#cite_note-0" title=""><span>[</span>1<span>]</span></a></sup></p>
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<p><a name="Biology" id="Biology"></a></p>
<h2><span class="editsection"></span><span class="mw-headline">Biology</span></h2>
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<div style="width: 182px;" class="thumbinner"><a href="/wiki/File:CrawlingCelegans.gif" title="Movement of Wild-type C. elegans" class="image"><img width="180" height="95" border="0" src="http://upload.wikimedia.org/wikipedia/commons/b/be/CrawlingCelegans.gif" alt="" class="thumbimage" /></a>
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Movement of Wild-type <em>C. elegans</em></div>
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<p><em>C. elegans</em> is <a href="/wiki/Segmentation_(biology)" title="Segmentation (biology)">unsegmented</a>, <a href="http://en.wiktionary.org/wiki/vermiform" title="wikt:vermiform" class="extiw">vermiform</a>, <a href="/wiki/Symmetry_(biology)#Bilateral_symmetry" title="Symmetry (biology)">bilaterally symmetrical</a>, with a <a href="/wiki/Cuticle" title="Cuticle">cuticle</a> integument, four main epidermal cords and a fluid-filled <a href="/wiki/Pseudocoelom" title="Pseudocoelom" class="mw-redirect">pseudocoelomate</a> cavity. Members of the species have many of the same organ systems as other animals. In the wild, they feed on bacteria that develop on decaying vegetable matter. Individuals of <em>C. elegans</em> are almost all <a href="/wiki/Hermaphrodite" title="Hermaphrodite">hermaphrodite</a>, with males comprising just 0.05% of the total population on average. The basic anatomy of <em>C. elegans</em> includes a mouth, <a href="/wiki/Pharynx" title="Pharynx">pharynx</a>, intestine, <a href="/wiki/Gonad" title="Gonad">gonad</a>, and collagenous cuticle. Males have a single-lobed gonad, <a href="/wiki/Vas_deferens" title="Vas deferens">vas deferens</a>, and a tail specialized for mating. Hermaphrodites have two ovaries, oviducts, spermatheca, and a single uterus.</p>
<p><em>C. elegans</em> eggs are laid by the hermaphrodite. After hatching, they pass through four <a href="/wiki/Larva" title="Larva">larval</a> stages (L1-L4). When crowded or in the absence of food, <em>C. elegans</em> can enter an alternative third larval stage called the <a href="/wiki/Dauer_larva" title="Dauer larva">dauer</a> state. Dauer larvae are stress-resistant and do not age. Hermaphrodites produce all their sperm in the L4 stage (150 sperm per gonadal arm) and then switch over to producing oocytes. The sperm are stored in the same area of the gonad as the oocytes until the first oocyte pushes the sperm into the spermatheca (a kind of chamber where the oocytes become fertilized by the sperm).<sup id="cite_ref-1" class="reference"><a href="#cite_note-1" title=""><span>[</span>2<span>]</span></a></sup> The male can inseminate the hermaphrodite, which will use male sperm preferentially (both types of sperm are stored in the spermatheca). When self-inseminated the wild-type worm will lay approximately 300 eggs. When inseminated by a male, the number of progeny can exceed 1,000. At 20<a href="/wiki/Celsius" title="Celsius">&deg;C</a>, the laboratory strain of <em>C. elegans</em> has an average life span of approximately 2&ndash;3 weeks and a generation time of approximately 4 days. Hermaphrodites can mate with males or self-fertilize.</p>
<p><em>C. elegans</em> has five pairs of <a href="/wiki/Autosome" title="Autosome">autosomes</a> and one pair of <a href="/wiki/Sex_chromosome" title="Sex chromosome" class="mw-redirect">sex chromosomes</a>. Sex in <em>C. elegans</em> is based on an <a href="/wiki/X0_sex-determination_system" title="X0 sex-determination system">X0 sex-determination system</a>. Hermaphrodite <em>C. elegans</em> have a matched pair of sex chromosomes (XX); the rare males have only one sex chromosome (X0).</p>
<p>It is interesting to note that <em>C. elegans</em> is one of the only forms of life not known to have a natural virus.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2" title=""><span>[</span>3<span>]</span></a></sup></p>
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<div style="width: 502px;" class="thumbinner"><a href="/wiki/File:C_elegans_anatomy.png" title="Longitudinal section through the hermaphrodite C. elegans." class="image"><img width="500" height="94" border="0" src="http://upload.wikimedia.org/wikipedia/commons/thumb/8/88/C_elegans_anatomy.png/500px-C_elegans_anatomy.png" alt="" class="thumbimage" /></a>
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Longitudinal section through the hermaphrodite <em>C. elegans.</em></div>
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<p><a name="Laboratory_uses" id="Laboratory_uses"></a></p>
<h2><span class="editsection"></span><span class="mw-headline">Laboratory uses</span></h2>
<p><em>C. elegans</em> is studied as a <a href="/wiki/Model_organism" title="Model organism">model organism</a> for a variety of reasons. <a href="/wiki/Strain_(biology)" title="Strain (biology)">Strains</a> are cheap to breed and can be frozen. When subsequently thawed they remain viable, allowing long-term storage. Because the complete cell lineage of the species has been determined, <em>C. elegans</em> has proven especially useful for studying <a href="/wiki/Cellular_differentiation" title="Cellular differentiation">cellular differentiation</a>.</p>
<p>From a research perspective, <em>C. elegans</em> has the advantage of being a <a href="/wiki/Multicellular" title="Multicellular" class="mw-redirect">multicellular</a> <a href="/wiki/Eukaryote" title="Eukaryote">eukaryotic</a> organism that is simple enough to be studied in great detail. In addition, it is transparent facilitating the study of developmental processes in the intact organism. The developmental fate of every single <a href="/wiki/Somatic_cell" title="Somatic cell">somatic cell</a> (959 in the adult hermaphrodite; 1031 in the adult male) has been mapped out. These patterns of cell lineage are largely invariant between individuals, in contrast to mammals where cell development from the embryo is more largely dependent on cellular cues. In both sexes, a large number of additional cells (131 in the hermaphrodite, most of which would otherwise become <a href="/wiki/Neuron" title="Neuron">neurons</a>), are eliminated by programmed cell death (<a href="/wiki/Apoptosis" title="Apoptosis">apoptosis</a>).</p>
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<div style="width: 252px;" class="thumbinner"><a href="/wiki/File:C_elegans_stained.jpg" title="Wild-type C. elegans hermaphrodite stained to highlight the nuclei of all cells" class="image"><img width="250" height="345" border="0" src="http://upload.wikimedia.org/wikipedia/commons/thumb/7/77/C_elegans_stained.jpg/250px-C_elegans_stained.jpg" alt="" class="thumbimage" /></a>
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Wild-type <em>C. elegans</em> hermaphrodite stained to highlight the nuclei of all cells</div>
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<p>In addition, <em>C. elegans</em> is one of the simplest organisms with a <a href="/wiki/Nervous_system" title="Nervous system">nervous system</a>. In the hermaphrodite, this comprises 302 <a href="/wiki/Neuron" title="Neuron">neurons</a> whose pattern of connectivity has been completely mapped out, and shown to be a <a href="/wiki/Small-world_network" title="Small-world network">small-world network</a>.<sup id="cite_ref-3" class="reference"><a href="#cite_note-3" title=""><span>[</span>4<span>]</span></a></sup> Research has explored the neural mechanisms responsible for several of the more interesting behaviors shown by <em>C. elegans</em>, including <a href="/wiki/Chemotaxis" title="Chemotaxis">chemotaxis</a>, <a href="/wiki/Thermotaxis" title="Thermotaxis">thermotaxis</a>, <a href="/wiki/Mechanotransduction" title="Mechanotransduction">mechanotransduction</a>, and male mating behavior.</p>
<p>A useful feature of <em>C. elegans</em> is that it is relatively straightforward to disrupt the function of specific genes by <a href="/wiki/RNA_interference" title="RNA interference">RNA interference</a> (RNAi). <a href="/wiki/Gene_silencing" title="Gene silencing">Silencing</a> the function of a gene in this way can sometimes allow a researcher to infer what the function of that gene may be. The nematode can either be soaked in (or injected with) a solution of double stranded <a href="/wiki/RNA" title="RNA">RNA</a>, the sequence of which is complementary to the sequence of the gene that the researcher wishes to disable. Alternatively, worms can be fed on genetically transformed <a href="/wiki/Bacteria" title="Bacteria">bacteria</a> which <a href="/wiki/Gene_expression" title="Gene expression">express</a> the double stranded RNA of interest.</p>
<p><em>C. elegans</em> has also been useful in the study of <a href="/wiki/Meiosis" title="Meiosis">meiosis</a>. As sperm and egg nuclei move down the length of the gonad, they undergo a temporal progression through meiotic events. This progression means that every nucleus at a given position in the gonad will be at roughly the same step in meiosis, eliminating the difficulties of heterogeneous populations of cells.</p>
<p>The organism has also been identified as a model for <a href="/wiki/Nicotine" title="Nicotine">nicotine</a> dependence as it has been found to experience the same symptoms humans experience when they <a href="/wiki/Smoking_cessation" title="Smoking cessation">quit smoking</a>.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4" title=""><span>[</span>5<span>]</span></a></sup></p>
<p>As for most model organisms, there is a dedicated online database for the species that is actively curated by scientists working in this field. The <a href="/wiki/Wormbase" title="Wormbase">WormBase</a> database attempts to collate all published information on <em>C. elegans</em> and other related nematodes. A reward of $5000 has been advertised on their website, for the finder of a new species of closely related nematode.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5" title=""><span>[</span>6<span>]</span></a></sup> Such a discovery would broaden research opportunities with the worm.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6" title=""><span>[</span>7<span>]</span></a></sup></p>
<p><a name="Genome" id="Genome"></a></p>
<h2><span class="editsection"></span><span class="mw-headline">Genome</span></h2>
<p><em>C. elegans</em> was the first multicellular organism to have its <a href="/wiki/Genome" title="Genome">genome</a> completely <a href="/wiki/Sequencing" title="Sequencing">sequenced</a>. The finished genome sequence was published in 1998,<sup id="cite_ref-7" class="reference"><a href="#cite_note-7" title=""><span>[</span>8<span>]</span></a></sup> although a number of small gaps were present (the last gap was finished by October 2002). The <em>C. elegans</em> genome sequence is approximately 100 million <a href="/wiki/Base_pair" title="Base pair">base pairs</a> long and contains approximately 20,000 <a href="/wiki/Gene" title="Gene">genes</a>. The vast majority of these genes encode for <a href="/wiki/Protein" title="Protein">proteins</a> but there are likely to be as many as 1,000 <a href="/wiki/RNA_gene" title="RNA gene" class="mw-redirect">RNA genes</a>. Scientific curators continue to appraise the set of known genes, such that new gene predictions continue to be added and incorrect ones modified or removed.</p>
<p>In 2003, the genome sequence of the related nematode <em><a href="/wiki/Caenorhabditis_briggsae" title="Caenorhabditis briggsae">C. briggsae</a></em> was also determined, allowing researchers to study the comparative genomics of these two organisms.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8" title=""><span>[</span>9<span>]</span></a></sup> Work is now ongoing to determine the genome sequences of more nematodes from the same <a href="/wiki/Genus" title="Genus">genus</a> such as <em>C. remanei</em>,<sup id="cite_ref-9" class="reference"><a href="#cite_note-9" title=""><span>[</span>10<span>]</span></a></sup> <em>C. japonica</em><sup id="cite_ref-10" class="reference"><a href="#cite_note-10" title=""><span>[</span>11<span>]</span></a></sup> and <em>C. brenneri</em>.<sup id="cite_ref-11" class="reference"><a href="#cite_note-11" title=""><span>[</span>12<span>]</span></a></sup> These newer genome sequences are being determined by using the <a href="/wiki/Whole_genome_shotgun" title="Whole genome shotgun" class="mw-redirect">whole genome shotgun</a> technique which means that the resulting genome sequences are likely to not be as complete or accurate as <em>C. elegans</em> (which was sequenced using the 'hierarchical' or <a href="/wiki/Clone_(genetics)" title="Clone (genetics)" class="mw-redirect">clone</a>-by-clone approach).</p>
<p>The official version of the <em>C. elegans</em> genome sequence continues to change as and when new evidence reveals errors in the original sequencing (<a href="/wiki/DNA_sequencing" title="DNA sequencing">DNA sequencing</a> is not an error-free process). Most changes are minor, adding or removing only a few <a href="/wiki/Base_pair" title="Base pair">base pairs</a> (bp) of DNA. E.g. the WS169 release of <a rel="nofollow" href="http://www.wormbase.org/" title="http://www.wormbase.org" class="external text">WormBase</a> (December 2006) lists a net gain of 6 bp to the genome sequence.<sup id="cite_ref-WS169_12-0" class="reference"><a href="#cite_note-WS169-12" title=""><span>[</span>13<span>]</span></a></sup> Occasionally more extensive changes are made, e.g. the WS159 release of May 2006 added over 300 bp to the sequence.<sup id="cite_ref-WS159_13-0" class="reference"><a href="#cite_note-WS159-13" title=""><span>[</span>14<span>]</span></a></sup></p>
<p><a name="Evolution" id="Evolution"></a></p>
<h2><span class="editsection"></span><span class="mw-headline">Evolution</span></h2>
<p>It has been shown that a small number of conserved <a href="/wiki/Protein" title="Protein">protein</a> sequences from <a href="/wiki/Sea_sponge" title="Sea sponge" class="mw-redirect">sponges</a> are more similar to humans than to <em>C. elegans</em>.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14" title=""><span>[</span>15<span>]</span></a></sup> This suggests that there has been an accelerated rate of evolution in the <em>C. elegans</em> <a href="/wiki/Lineage_(evolution)" title="Lineage (evolution)">lineage</a>. The same study found that several <a href="/wiki/Phylogenetically" title="Phylogenetically" class="mw-redirect">phylogenetically</a> ancient genes are not present in <em>C. elegans</em>.</p>
<p><a name="Scientific_community" id="Scientific_community"></a></p>
<h2><span class="editsection"></span><span class="mw-headline">Scientific community</span></h2>
<p>In 2002, the <a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a> was awarded to <a href="/wiki/Sydney_Brenner" title="Sydney Brenner">Sydney Brenner</a>, <a href="/wiki/H._Robert_Horvitz" title="H. Robert Horvitz">H. Robert Horvitz</a> and <a href="/wiki/John_Sulston" title="John Sulston" class="mw-redirect">John Sulston</a> for their work on the genetics of organ development and <a href="/wiki/Apoptosis" title="Apoptosis">programmed cell death</a> (PCD) in <em>C. elegans</em>. The 2006 <a href="/wiki/Nobel_Prize_in_Physiology_or_Medicine" title="Nobel Prize in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a> was awarded to <a href="/wiki/Andrew_Fire" title="Andrew Fire">Andrew Fire</a> and <a href="/wiki/Craig_C._Mello" title="Craig C. Mello" class="mw-redirect">Craig C. Mello</a>, for their discovery of <a href="/wiki/RNA_interference" title="RNA interference">RNA interference</a> in <em>C. elegans</em>.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15" title=""><span>[</span>16<span>]</span></a></sup>. In 2008 <a href="/wiki/Martin_Chalfie" title="Martin Chalfie">Martin Chalfie</a> shared a <a href="/wiki/Nobel_Prize_in_Chemistry" title="Nobel Prize in Chemistry">Nobel Prize in Chemistry</a> for his work on <a href="/wiki/Green_fluorescent_protein" title="Green fluorescent protein">green fluorescent protein</a> (GFP) in <em>C. elegans</em>.</p>
<p>Because all research into <em>C. elegans</em> essentially started with <a href="/wiki/Sydney_Brenner" title="Sydney Brenner">Sydney Brenner</a> in the 1970s, many scientists working in this field share a close connection to Brenner (they either worked as a <a href="/wiki/Post-doctoral" title="Post-doctoral" class="mw-redirect">post-doctoral</a> or <a href="/wiki/Post-graduate" title="Post-graduate" class="mw-redirect">post-graduate</a> researcher in Brenner's lab or in the lab of someone who previously worked with Brenner). Because most people who worked in his lab went on to establish their own worm research labs, there is now a fairly well documented 'lineage' of <em>C. elegans</em> scientists. This lineage was recorded in some detail at the 2003 International Worm Meeting and the results were stored in the <a href="/wiki/Wormbase" title="Wormbase">Wormbase</a> database.</p>
<p><a name="In_the_media" id="In_the_media"></a></p>
<h2><span class="editsection"></span><span class="mw-headline">In the media</span></h2>
<p><em>C. elegans</em> made news when it was discovered that specimens had survived the <a href="/wiki/Space_Shuttle_Columbia_disaster" title="Space Shuttle Columbia disaster">Space Shuttle <em>Columbia</em> disaster</a> in February 2003.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16" title=""><span>[</span>17<span>]</span></a></sup> Later, in January 2009, it was announced that live samples of C.elegans from the <a href="/wiki/University_of_Nottingham" title="University of Nottingham">University of Nottingham</a> will spend two weeks on the <a href="/wiki/International_Space_Station" title="International Space Station">International Space Station</a> as part of a project to explore the effects of <a href="/wiki/Zero_gravity" title="Zero gravity" class="mw-redirect">zero gravity</a> on muscle development and its physiology. The emphasis of the research will be on the genetic basis of <a href="/wiki/Muscle_atrophy" title="Muscle atrophy">muscle atrophy</a>. This has relevance to space travel, but also to individuals who are bed-ridden, <a href="/wiki/Geriatric" title="Geriatric" class="mw-redirect">geriatric</a> or <a href="/wiki/Diabetic" title="Diabetic" class="mw-redirect">diabetic</a>.<sup id="cite_ref-17" class="reference"><a href="#cite_note-17" title=""><span>[</span>18<span>]</span></a></sup></p>
<p><a name="See_also" id="See_also"></a></p>
<h2><span class="editsection"></span><span class="mw-headline">See also</span></h2>
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<li><a href="/wiki/Animal_testing_on_invertebrates" title="Animal testing on invertebrates">Animal testing on invertebrates</a> </li>
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<p><a name="References" id="References"></a></p>
<h2><span class="editsection"></span><span class="mw-headline">References</span></h2>
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<li id="cite_note-0"><strong><a href="#cite_ref-0" title="">^</a></strong> <cite style="font-style: normal;" id="CITEREFBrenner.2C_S.1974" class="Journal">Brenner, S. (May 1974). &quot;<a rel="nofollow" href="http://dev.wormbase.org/papers/31_Brenner74.pdf" title="http://dev.wormbase.org/papers/31_Brenner74.pdf" class="external text">The Genetics of <em>Caenorhabditis elegans</em></a>&quot; (PDF). <em><a href="/wiki/Genetics_(journal)" title="Genetics (journal)">Genetics</a></em> <strong>77</strong>: 71&ndash;94<span class="printonly">. <a rel="nofollow" href="http://dev.wormbase.org/papers/31_Brenner74.pdf" title="http://dev.wormbase.org/papers/31_Brenner74.pdf" class="external free">http://dev.wormbase.org/papers/31_Brenner74.pdf</a></span>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.atitle=The+Genetics+of+%27%27Caenorhabditis+elegans%27%27&amp;rft.jtitle=%5B%5BGenetics+%28journal%29%7CGenetics%5D%5D&amp;rft.aulast=Brenner%2C+S.&amp;rft.au=Brenner%2C+S.&amp;rft.date=May+1974&amp;rft.volume=77&amp;rft.pages=71%E2%80%9394&amp;rft_id=http%3A%2F%2Fdev.wormbase.org%2Fpapers%2F31_Brenner74.pdf&amp;rfr_id=info:sid/en.wikipedia.org:Caenorhabditis_elegans" class="Z3988"><span style="display: none;">&nbsp;</span></span> </li>
<li id="cite_note-1"><strong><a href="#cite_ref-1" title="">^</a></strong> <cite style="font-style: normal;" id="CITEREFNayak.2C_S..2C_J._Goree_.26_T._Schedl2004" class="Journal">Nayak, S., J. Goree &amp; T. Schedl (2004). &quot;<a rel="nofollow" href="http://biology.plosjournals.org/perlserv/?request=get-document&amp;doi=10.1371/journal.pbio.0030006" title="http://biology.plosjournals.org/perlserv/?request=get-document&amp;doi=10.1371/journal.pbio.0030006" class="external text"><em>fog-2</em> and the Evolution of Self-Fertile Hermaphroditism in <em>Caenorhabditis</em></a>&quot;. <em><a href="/wiki/PLoS_Biology" title="PLoS Biology">PLoS Biology</a></em> <strong>3</strong> (1): e6. <a href="/wiki/Digital_object_identifier" title="Digital object identifier">doi</a>:<span class="neverexpand"><a rel="nofollow" href="http://dx.doi.org/10.1371%2Fjournal.pbio.0030006" title="http://dx.doi.org/10.1371%2Fjournal.pbio.0030006" class="external text">10.1371/journal.pbio.0030006</a></span><span class="printonly">. <a rel="nofollow" href="http://biology.plosjournals.org/perlserv/?request=get-document&amp;doi=10.1371/journal.pbio.0030006" title="http://biology.plosjournals.org/perlserv/?request=get-document&amp;doi=10.1371/journal.pbio.0030006" class="external free">http://biology.plosjournals.org/perlserv/?request=get-document&amp;doi=10.1371/journal.pbio.0030006</a></span>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.atitle=%27%27fog-2%27%27+and+the+Evolution+of+Self-Fertile+Hermaphroditism+in+%27%27Caenorhabditis%27%27&amp;rft.jtitle=%5B%5BPLoS+Biology%5D%5D&amp;rft.aulast=Nayak%2C+S.%2C+J.+Goree+%26+T.+Schedl&amp;rft.au=Nayak%2C+S.%2C+J.+Goree+%26+T.+Schedl&amp;rft.date=2004&amp;rft.volume=3&amp;rft.issue=1&amp;rft.pages=e6&amp;rft_id=info:doi/10.1371%2Fjournal.pbio.0030006&amp;rft_id=http%3A%2F%2Fbiology.plosjournals.org%2Fperlserv%2F%3Frequest%3Dget-document%26doi%3D10.1371%2Fjournal.pbio.0030006&amp;rfr_id=info:sid/en.wikipedia.org:Caenorhabditis_elegans" class="Z3988"><span style="display: none;">&nbsp;</span></span> </li>
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<li id="cite_note-14"><strong><a href="#cite_ref-14" title="">^</a></strong> <cite style="font-style: normal;" id="CITEREFGamulin2000" class="Journal">Gamulin, V (December 2000). &quot;Sponge proteins are more similar to those of <em>Homo sapiens</em> than to <em>Caenorhabditis elegans</em>&quot;. <em>Biological Journal of the Linnean Society</em> (Academic Press) <strong>71</strong> (4): 821&ndash;828.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.atitle=Sponge+proteins+are+more+similar+to+those+of+%27%27Homo+sapiens%27%27+than+to+%27%27Caenorhabditis+elegans%27%27&amp;rft.jtitle=Biological+Journal+of+the+Linnean+Society&amp;rft.aulast=Gamulin&amp;rft.aufirst=V&amp;rft.au=Gamulin%2C+V&amp;rft.date=December+2000&amp;rft.volume=71&amp;rft.issue=4&amp;rft.pages=821%E2%80%93828&amp;rft.pub=Academic+Press&amp;rfr_id=info:sid/en.wikipedia.org:Caenorhabditis_elegans" class="Z3988"><span style="display: none;">&nbsp;</span></span> </li>
<li id="cite_note-15"><strong><a href="#cite_ref-15" title="">^</a></strong> <cite style="font-style: normal;" id="CITEREFFire_A.2C_Xu_S.2C_Montgomery_MK.2C_Kostas_SA.2C_Driver_SE.2C_Mello_CC1998" class="Journal">Fire A, Xu S, Montgomery MK, Kostas SA, Driver SE, Mello CC (February 1998). &quot;Potent and specific genetic interference by double-stranded RNA in Caenorhabditis elegans&quot;. <em>Nature</em> <strong>391</strong> (6669): 806&ndash;11. <a href="/wiki/Digital_object_identifier" title="Digital object identifier">doi</a>:<span class="neverexpand"><a rel="nofollow" href="http://dx.doi.org/10.1038%2F35888" title="http://dx.doi.org/10.1038%2F35888" class="external text">10.1038/35888</a></span>. <a href="http://www.ncbi.nlm.nih.gov/pubmed/9486653" title="http://www.ncbi.nlm.nih.gov/pubmed/9486653" class="external">PMID 9486653</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&amp;rft.genre=article&amp;rft.atitle=Potent+and+specific+genetic+interference+by+double-stranded+RNA+in+Caenorhabditis+elegans&amp;rft.jtitle=Nature&amp;rft.aulast=Fire+A%2C+Xu+S%2C+Montgomery+MK%2C+Kostas+SA%2C+Driver+SE%2C+Mello+CC&amp;rft.au=Fire+A%2C+Xu+S%2C+Montgomery+MK%2C+Kostas+SA%2C+Driver+SE%2C+Mello+CC&amp;rft.date=February+1998&amp;rft.volume=391&amp;rft.issue=6669&amp;rft.pages=806%E2%80%9311&amp;rft_id=info:doi/10.1038%2F35888&amp;rft_id=info:pmid/9486653&amp;rfr_id=info:sid/en.wikipedia.org:Caenorhabditis_elegans" class="Z3988"><span style="display: none;">&nbsp;</span></span> </li>
<li id="cite_note-16"><strong><a href="#cite_ref-16" title="">^</a></strong> &quot;<a rel="nofollow" href="http://news.bbc.co.uk/1/hi/sci/tech/2992123.stm" title="http://news.bbc.co.uk/1/hi/sci/tech/2992123.stm" class="external text">Worms survived Columbia disaster</a>&quot;, <a href="/wiki/BBC_News" title="BBC News">BBC News</a> (2003-05-01)<span class="reference-accessdate">. Retrieved on 11 July 2008</span>.<span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Adc&amp;rft.type=newspaperArticle&amp;rft.subject=News&amp;rft.title=Worms+survived+Columbia+disaster&amp;rft.identifier=http%3A%2F%2Fnews.bbc.co.uk%2F1%2Fhi%2Fsci%2Ftech%2F2992123.stm&amp;rft.publisher=%5B%5BBBC+News%5D%5D&amp;rft.date=2003-05-01" class="Z3988"><span style="display: none;">&nbsp;</span></span> </li>
<li id="cite_note-17"><strong><a href="#cite_ref-17" title="">^</a></strong> <a rel="nofollow" href="http://news.bbc.co.uk/1/hi/england/nottinghamshire/7835020.stm" title="http://news.bbc.co.uk/1/hi/england/nottinghamshire/7835020.stm" class="external free">http://news.bbc.co.uk/1/hi/england/nottinghamshire/7835020.stm</a> </li>
</ol>
</div>
<p><a name="Publications" id="Publications"></a></p>
<h2><span class="editsection"></span><span class="mw-headline">Publications</span></h2>
<ul>
<li><cite style="font-style: normal;" id="CITEREFBird.2C_Jean.3B_Bird.2C_Alan_C.1991" class="book">Bird, Jean; Bird, Alan C. (1991). <em>The structure of nematodes</em>. Boston: <a href="/wiki/Academic_Press" title="Academic Press">Academic Press</a>. pp.&nbsp;1, 69&ndash;70, 152&ndash;153, 165, 224&ndash;225. <a href="/wiki/Special:BookSources/0120996510" class="internal">ISBN 0-12-099651-0</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=The+structure+of+nematodes&amp;rft.aulast=Bird%2C+Jean%3B+Bird%2C+Alan+C.&amp;rft.au=Bird%2C+Jean%3B+Bird%2C+Alan+C.&amp;rft.date=1991&amp;rft.pages=pp.%26nbsp%3B1%2C+69%E2%80%9370%2C+152%E2%80%93153%2C+165%2C+224%E2%80%93225&amp;rft.place=Boston&amp;rft.pub=%5B%5BAcademic+Press%5D%5D&amp;rft.isbn=0-12-099651-0&amp;rfr_id=info:sid/en.wikipedia.org:Caenorhabditis_elegans" class="Z3988"><span style="display: none;">&nbsp;</span></span> </li>
</ul>
<ul>
<li><cite style="font-style: normal;" id="CITEREFHope.2C_Ian_A.1999" class="book">Hope, Ian A. (1999). <em>C. elegans: a practical approach</em>. Oxford [Oxfordshire]: <a href="/wiki/Oxford_University_Press" title="Oxford University Press">Oxford University Press</a>. pp.&nbsp;1&ndash;6. <a href="/wiki/Special:BookSources/0199637385" class="internal">ISBN 0-19-963738-5</a>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=C.+elegans%3A+a+practical+approach&amp;rft.aulast=Hope%2C+Ian+A.&amp;rft.au=Hope%2C+Ian+A.&amp;rft.date=1999&amp;rft.pages=pp.%26nbsp%3B1%E2%80%936&amp;rft.place=Oxford+%5BOxfordshire%5D&amp;rft.pub=%5B%5BOxford+University+Press%5D%5D&amp;rft.isbn=0-19-963738-5&amp;rfr_id=info:sid/en.wikipedia.org:Caenorhabditis_elegans" class="Z3988"><span style="display: none;">&nbsp;</span></span> </li>
</ul>
<ul>
<li><cite style="font-style: normal;" id="CITEREFRiddle.2C_D.L..2C_T._Blumenthal.2C_R._J._Meyer_.26_J._R._Priess1997" class="book">Riddle, D.L., T. Blumenthal, R. J. Meyer &amp; J. R. Priess (1997). <em><a rel="nofollow" href="http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=ce2" title="http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=ce2" class="external text">C. elegans <em>II</em></a></em>. Cold Spring Harbor Laboratory Press, New York. pp.&nbsp;1&ndash;4, 679&ndash;683. <a href="/wiki/Special:BookSources/0879695323" class="internal">ISBN 0-87969-532-3</a><span class="printonly">. <a rel="nofollow" href="http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=ce2" title="http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=ce2" class="external free">http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=ce2</a></span>.</cite><span title="ctx_ver=Z39.88-2004&amp;rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&amp;rft.genre=book&amp;rft.btitle=%27%27C.+elegans%27%27+II&amp;rft.aulast=Riddle%2C+D.L.%2C+T.+Blumenthal%2C+R.+J.+Meyer+%26+J.+R.+Priess&amp;rft.au=Riddle%2C+D.L.%2C+T.+Blumenthal%2C+R.+J.+Meyer+%26+J.+R.+Priess&amp;rft.date=1997&amp;rft.pages=pp.%26nbsp%3B1%E2%80%934%2C+679%E2%80%93683&amp;rft.pub=Cold+Spring+Harbor+Laboratory+Press%2C+New+York&amp;rft.isbn=0-87969-532-3&amp;rft_id=http%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fbooks%2Fbv.fcgi%3Frid%3Dce2&amp;rfr_id=info:sid/en.wikipedia.org:Caenorhabditis_elegans" class="Z3988"><span style="display: none;">&nbsp;</span></span> </li>
</ul>
<p><a name="Online_resources" id="Online_resources"></a></p>
<h2><span class="editsection"></span><span class="mw-headline">Online resources</span></h2>
<ul>
<li><a rel="nofollow" href="http://www.wormbase.org/" title="http://www.wormbase.org" class="external text">WormBase</a> - an extensive online database covering the biology and genomics of <em>C. elegans</em> and other nematodes </li>
<li><a rel="nofollow" href="http://www.wormbook.org/" title="http://www.wormbook.org/" class="external text">WormBook</a> - a free online compendium of all aspects of <em>C. elegans</em> biology, including laboratory protocols </li>
<li><a rel="nofollow" href="http://www.wormatlas.org/" title="http://www.wormatlas.org" class="external text">Wormatlas</a> - an online database for behavioral and structural anatomy of <em>C. elegans</em> </li>
<li><a rel="nofollow" href="http://www.sanger.ac.uk/Projects/C_elegans" title="http://www.sanger.ac.uk/Projects/C_elegans" class="external text">Wellcome Trust Sanger Institute <em>C. elegans</em> page</a> - half of the genome sequence is still maintained by this institute </li>
<li><a rel="nofollow" href="http://genome.wustl.edu/genome.cgi?GENOME=Caenorhabditis%20elegans" title="http://genome.wustl.edu/genome.cgi?GENOME=Caenorhabditis%20elegans" class="external text">WashU Genome Sequencing Center <em>C. elegans</em> page</a> - the institute maintaining the other half of the genome </li>
<li><a rel="nofollow" href="http://www.ncbi.nlm.nih.gov/IEB/Research/Acembly/index.html?worm" title="http://www.ncbi.nlm.nih.gov/IEB/Research/Acembly/index.html?worm" class="external text">AceView WormGenes</a> - another genome database for <em>C. elegans</em>, maintained at the NCBI </li>
<li><a rel="nofollow" href="http://www.tcnj.edu/~nayaklab" title="http://www.tcnj.edu/~nayaklab" class="external text">TCNJ Worm Lab</a> - Easy to follow protocols and pictures for <em>C. elegans</em> research. Made by undergrads for undergrads. </li>
<li><a rel="nofollow" href="http://www.wormclassroom.org/" title="http://www.wormclassroom.org" class="external text">Worm Classroom</a> - An education portal for <em>C. elegans</em> </li>
<li><a rel="nofollow" href="http://www.textpresso.org/" title="http://www.textpresso.org/" class="external text">Textpresso</a> - WormBase search engine </li>
<li><a rel="nofollow" href="http://www.bio.unc.edu/faculty/goldstein/lab/movies.html" title="http://www.bio.unc.edu/faculty/goldstein/lab/movies.html" class="external text"><em>C. elegans</em> movies</a> - Timelapse films made by <em>C. elegans</em> researchers worldwide </li>
<li><a rel="nofollow" href="http://www.ncbi.nlm.nih.gov/books/bv.fcgi?call=bv.View..ShowTOC&amp;rid=ce2.TOC" title="http://www.ncbi.nlm.nih.gov/books/bv.fcgi?call=bv.View..ShowTOC&amp;rid=ce2.TOC" class="external text"><em>C. elegans</em> II</a> - a free online textbook. </li>
<li><a rel="nofollow" href="http://www.silencinggenomes.org/" title="http://www.silencinggenomes.org" class="external text">Silencing Genomes</a> RNA interference (RNAi) experiments and bioinformatics in <em>C. elegans</em> for education. From the Dolan DNA Learning Center of Cold Spring Harbor Laboratory. </li>
<li><a rel="nofollow" href="http://www.ciml.univ-mrs.fr/EWBANK_jonathan/3D/mainpage.html" title="http://www.ciml.univ-mrs.fr/EWBANK_jonathan/3D/mainpage.html" class="external text">C.elegans 3D model by the Ewbank Lab</a> - Videos and photos that explain the basic anatomy of <em>C. elegans</em> </li>
<li><a rel="nofollow" href="http://www.wormtracker.de/" title="http://www.wormtracker.de/" class="external text">WormTracker</a> </li>
</ul>
<p><a name="Nobel_lectures" id="Nobel_lectures"></a></p>
<h2><span class="editsection"></span><span class="mw-headline">Nobel lectures</span></h2>
<ul>
<li>Brenner S (2002) Nature's Gift to Science. In. <a rel="nofollow" href="http://nobelprize.org/nobel_prizes/medicine/laureates/2002/brenner-lecture.pdf" title="http://nobelprize.org/nobel_prizes/medicine/laureates/2002/brenner-lecture.pdf" class="external free">http://nobelprize.org/nobel_prizes/medicine/laureates/2002/brenner-lecture.pdf</a> </li>
<li>Horvitz HR (2002) Worms, Life and Death. In. <a rel="nofollow" href="http://nobelprize.org/nobel_prizes/medicine/laureates/2002/horvitz-lecture.pdf" title="http://nobelprize.org/nobel_prizes/medicine/laureates/2002/horvitz-lecture.pdf" class="external free">http://nobelprize.org/nobel_prizes/medicine/laureates/2002/horvitz-lecture.pdf</a> </li>
<li>Sulston JE (2002) The Cell Lineage and Beyond. In. <a rel="nofollow" href="http://nobelprize.org/nobel_prizes/medicine/laureates/2002/sulston-lecture.pdf" title="http://nobelprize.org/nobel_prizes/medicine/laureates/2002/sulston-lecture.pdf" class="external free">http://nobelprize.org/nobel_prizes/medicine/laureates/2002/sulston-lecture.pdf</a> </li>
</ul>
<p><a name="External_links" id="External_links"></a></p>
<h2><span class="editsection"></span><span class="mw-headline">External links</span></h2>
<ul>
<li><a rel="nofollow" href="http://www.nytimes.com/2006/11/21/science/21find.html" title="http://www.nytimes.com/2006/11/21/science/21find.html" class="external text">Nematodes With a Craving for Nicotine</a> </li>
</ul>
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