image_width = 300px
image_caption = An adult hermaphrodite "C. elegans" worm
phylum = Nematoda
genus = "
species = "C. elegans"
binomial = "Caenorhabditis elegans"
binomial_authority = Maupas, 1900
"Caenorhabditis elegans" (pronEng|ˌsiːnoʊræbˈdaɪt
ɪs ˈɛl ɪgænz) is a free-living nematode(roundworm), about 1 mm in length, which lives in temperate soil environments. Research into the molecular and developmental biologyof "C. elegans" was begun in 1974 by Sydney Brennerand it has since been used extensively as a model organism. [cite journal | quotes=no |author=Brenner, S. |year=1974 |month=May |url=http://dev.wormbase.org/papers/31_Brenner74.pdf |title=The Genetics of "Caenorhabditis elegans" |journal=Genetics |volume=77 |pages=71–94|format=PDF]
"C. elegans" is unsegmented, , bilaterally symmetrical, with a
cuticleintegument, four main epidermal cords and a fluid-filled pseudocoelomate 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 "C. elegans" are almost all hermaphrodite, with males comprising just 0.05% of the total population on average. The basic anatomy of "C. elegans" includes a mouth, pharynx, intestine, gonad, and collagenous cuticle. Males have a single-lobed gonad, vas deferens, and a tail specialized for mating. Hermaphrodites have two ovaries, oviducts, spermatheca, and a single uterus.
"C. elegans" eggs are laid by the hermaphrodite. After hatching, they pass through four
larval stages (L1-L4). When crowded or in the absence of food, "C. elegans" can enter an alternative third larval stage called the dauer 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). [cite journal | quotes=no |url=http://biology.plosjournals.org/perlserv/?request=get-document&doi=10.1371/journal.pbio.0030006 |title="fog-2" and the Evolution of Self-Fertile Hermaphroditism in "Caenorhabditis"|journal= PLoS Biology|volume=3 |issue=1 |pages=e6 |author=Nayak, S., J. Goree & T. Schedl |year=2004 |doi=10.1371/journal.pbio.0030006] 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°C, the laboratory strain of "C. elegans" has an average life span of approximately 2–3 weeks and a generation time of approximately 4 days. Hermaphrodites can mate with males or self-fertilize.
"C. elegans" has five pairs of
autosomes and one pair of sex chromosomes. Sex in "C. elegans" is based on an X0 sex-determination system. Hermaphrodite "C. elegans" have a matched pair of sex chromosomes (XX); the rare males have only one sex chromosome (X0).
"C. elegans" is studied as a
model organismfor a variety of reasons. Strains 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, "C. elegans" has proven especially useful for studying cellular differentiation.
From a research perspective, "C. elegans" has the advantage of being a
multicellulareukaryotic organism that is simple enough to be studied in great detail. The developmental fate of every single somatic cell(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 neurons), are eliminated by programmed cell death ( apoptosis).
In addition, "C. elegans" is one of the simplest organisms with a
nervous system. In the hermaphrodite, this comprises 302 neurons whose pattern of connectivity has been completely mapped out, and shown to be a small-world network. [cite journal |quotes=no |author=Watts D. J. & S. H. Strogatz |url=http://www.nature.com/nature/journal/v393/n6684/abs/393440a0.html |title=Collective dynamics of 'small-world' networks |journal=Nature |year=1998 |month=June |volume=393 |issue=6684 |pages=440–442 |doi=10.1038/30918] Research has explored the neural mechanisms responsible for several of the more interesting behaviors shown by "C. elegans", including chemotaxis, thermotaxis, mechanotransduction, and male mating behavior. Unusually, the neurons fire no action potentials.
A useful feature of "C. elegans" is that it is relatively straightforward to disrupt the function of specific genes by
RNA interference(RNAi). Silencing 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 RNA, 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 bacteriawhich express the double stranded RNA of interest.
"C. elegans" has also been useful in the study of
meiosis. 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.
The organism has also been identified as a model for
nicotinedependence as it has been found to experience the same symptoms humans experience when they quit smoking. [cite journal | quotes=no |author=Feng et al. |url=http://www.cell.com/content/article/abstract?uid=PIIS0092867406012955 |title=A C. elegans Model of Nicotine-Dependent Behavior: Regulation by TRP-Family Channels |journal=Cell |year=2006 |month=November |volume=127 |pages=621–633 |doi=10.1016/j.cell.2006.09.035 |pmid=17081982]
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 WormBase database attempts to collate all published information on "C. elegans" 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. [cite web |title="Caenorhabditis" isolation guide |url=http://wormbase.org/external/2007/nematode_isolation_guide/nematode_isolation_guide.html |publisher=WormBase |accessdate=2007-08-30] Such a discovery would broaden research opportunities with the worm. [cite journal |title=Slime for a dime |journal=Science |month=August |year=2007 |volume=317 |issue=5842 |pages=1157 |doi=10.1126/science.317.5842.1157b |last=Dolgin |first=Elie]
"C. elegans" was the first multicellular organism to have its
genomecompletely sequenced. The finished genome sequence was published in 1998, [cite journal |quotes=no |author=The "C. elegans" Sequencing Consortium |year=1998 |url=http://www.sciencemag.org/cgi/content/abstract/282/5396/2012 |title=Genome sequence of the nematode "C. elegans": a platform for investigating biology |journal=Science |volume=282 |pages=2012–2018 |doi=10.1126/science.282.5396.2012 |pmid=9851916] although a number of small gaps were present (the last gap was finished by October 2002). The "C. elegans" genome sequence is approximately 100 million base pairs long and contains approximately 20,000 genes. The vast majority of these genes encode for proteins but there are likely to be as many as 1,000 RNA genes. 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.
In 2003, the genome sequence of the related nematode "C. briggsae" was also determined, allowing researchers to study the comparative genomics of these two organisms. [cite journal | quotes=no |author=Stein, L. D. "et al." |year=2003 |doi= 10.1371/journal.pbio.0000045 |title=The Genome Sequence of "Caenorhabditis briggsae": A Platform for Comparative Genomics |journal=
PLoS Biology|volume=1 |pages=166–192] Work is now ongoing to determine the genome sequences of more nematodes from the same genussuch as "C. remanei", [cite web |url=http://genome.wustl.edu/genome.cgi?GENOME=Caenorhabditis%20remanei |author=Genome Sequencing Center |title="Caenorhabditis remanei": Background |accessdate=2008-07-11 |publisher= Washington University School of Medicine] "C. japonica" [cite web |url=http://genome.wustl.edu/genome.cgi?GENOME=Caenorhabditis%20japonica |author=Genome Sequencing Center |title="Caenorhabditis japonica": Background |accessdate=2008-07-11 |publisher= Washington University School of Medicine] and "C. brenneri". [cite web |url=http://genome.wustl.edu/genome.cgi?GENOME=Caenorhabditis%20brenneri |author=Genome Sequencing Center |title="Caenorhabditis brenneri": Background |accessdate=2008-07-11 |publisher= Washington University School of Medicine] These newer genome sequences are being determined by using the whole genome shotguntechnique which means that the resulting genome sequences are likely to not be as complete or accurate as "C. elegans" (which was sequenced using the 'hierarchical' or clone-by-clone approach).
The official version of the "C. elegans" genome sequence continues to change as and when new evidence reveals errors in the original sequencing (
DNA sequencingis not an error free process). Most changes are usually minor, adding or removing only a few base pairs (bp) of DNA. E.g. the WS169 release of [http://www.wormbase.org WormBase] (December 2006) lists a net gain of 6 bp to the genome sequence.cite web | title = WormBaseWiki WS169 release notes | url=http://www.wormbase.org/wiki/index.php/WS169 | accessdate = 2007-02-21 | publisher = Wormbase] Occasionally more extensive changes are made, e.g. the WS159 release of May 2006 added over 300 bp to the sequence.cite web | title = WormBaseWiki WS159 release notes | url=http://www.wormbase.org/wiki/index.php/WS159 | accessdate = 2007-01-21 | publisher = Wormbase]
It has been shown that a small number of conserved
proteinsequences from sponges are more similar to humans than to "C. elegans". [cite journal |last=Gamulin |first=V |title=Sponge proteins are more similar to those of "Homo sapiens" than to "Caenorhabditis elegans" |journal=Biological Journal of the Linnean Society |volume=71 |issue=4 |pages=821–828 |publisher=Academic Press |month=December |year=2000] This suggests that there has been an accelerated rate of evolution in the "C. elegans" lineage. The same study found that several phylogeneticallyancient genes are not present in "C. elegans".
In 2002, the
Nobel Prize in Physiology or Medicinewas awarded to Sydney Brenner, H. Robert Horvitzand John Sulstonfor their work on the genetics of organ development and programmed cell death (PCD) in "C. elegans". The 2006 Nobel Prize in Physiology or Medicinewas awarded to Andrew Fireand Craig C. Mello, for their discovery of RNA interferencein "C. elegans". [cite journal |author=Fire A, Xu S, Montgomery MK, Kostas SA, Driver SE, Mello CC |title=Potent and specific genetic interference by double-stranded RNA in Caenorhabditis elegans |journal=Nature |volume=391 |issue=6669 |pages=806–11 |year=1998 |month=February |pmid=9486653 |doi=10.1038/35888 |accessdate=2008-07-08] . In 2008 Martin Chalfieshared a Nobel Prize in Chemistryfor his work on green fluorescent protein(GFP) in "C. elegans".
Because all research into "C. elegans" essentially started with
Sydney Brennerin the 1970s, many scientists working in this field share a close connection to Brenner (they either worked as a post-doctoralor post-graduateresearcher 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 "C. elegans" scientists. This lineage was recorded in some detail at the 2003 International Worm Meeting and the results were stored in the Wormbasedatabase.
In the media
"C. elegans" made news when it was discovered that specimens had survived the Space Shuttle "Columbia" disaster in February 2003. [cite news |title=Worms survived Columbia disaster |url=http://news.bbc.co.uk/1/hi/sci/tech/2992123.stm |publisher=
BBC News|date= 2003-05-01|accessdate=2008-07-11]
Animal testing on invertebrates
*cite book |author=Bird, Jean; Bird, Alan C. |title=The structure of nematodes |publisher=
Academic Press|location=Boston |year=1991 |isbn=0-12-099651-0 |pages=pp 1, 69–70, 152–153, 165, 224–225
*cite book |author=Hope, Ian A. |title=C. elegans: a practical approach |publisher=
Oxford University Press|location=Oxford [Oxfordshire] |year=1999 |pages=pp 1–6 |isbn=0-19-963738-5
*cite book |author=Riddle, D.L., T. Blumenthal, R. J. Meyer & J. R. Priess |year=1997 |url=http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=ce2 |title="C. elegans" II |publisher=Cold Spring Harbor Laboratory Press, New York |pages=pp 1-4, 679–683 |isbn=0-87969-532-3
* [http://www.wormbase.org WormBase] - an extensive online database covering the biology and genomics of "C. elegans" and other nematodes
* [http://www.wormbook.org/ WormBook] - a free online compendium of all aspects of "C. elegans" biology, including laboratory protocols
* [http://www.wormatlas.org Wormatlas] - an online database for behavioral and structural anatomy of "C. elegans"
* [http://www.sanger.ac.uk/Projects/C_elegans Wellcome Trust Sanger Institute "C. elegans" page] - half of the genome sequence is still maintained by this institute
* [http://genome.wustl.edu/genome.cgi?GENOME=Caenorhabditis%20elegans WashU Genome Sequencing Center "C. elegans" page] - the institute maintaining the other half of the genome
* [http://www.ncbi.nlm.nih.gov/IEB/Research/Acembly/index.html?worm AceView WormGenes] - another genome database for "C. elegans", maintained at the NCBI
* [http://www.tcnj.edu/~nayaklab TCNJ Worm Lab] - Easy to follow protocols and pictures for "C. elegans" research. Made by undergrads for undergrads.
* [http://www.wormclassroom.org Worm Classroom] - An education portal for "C. elegans"
* [http://www.textpresso.org/ Textpresso] - WormBase search engine
* [http://www.bio.unc.edu/faculty/goldstein/lab/movies.html "C. elegans" movies] - Timelapse films made by "C. elegans" researchers worldwide
* [http://www.ncbi.nlm.nih.gov/books/bv.fcgi?call=bv.View..ShowTOC&rid=ce2.TOC "C. elegans" II] - a free online textbook.
* [http://www.silencinggenomes.org Silencing Genomes] RNA interference (RNAi) experiments and bioinformatics in "C. elegans" for education. From the Dolan DNA Learning Center of Cold Spring Harbor Laboratory.
* [http://www.ciml.univ-mrs.fr/EWBANK_jonathan/3D/mainpage.html C.elegans 3D model by the Ewbank Lab] - Videos and photos that explain the basic anatomy of "C. elegans"
* [http://www.wormtracker.de/ WormTracker]
*Brenner S (2002) Nature's Gift to Science. In. http://nobelprize.org/nobel_prizes/medicine/laureates/2002/brenner-lecture.pdf
*Horvitz HR (2002) Worms, Life and Death. In. http://nobelprize.org/nobel_prizes/medicine/laureates/2002/horvitz-lecture.pdf
*Sulston JE (2002) The Cell Lineage and Beyond. In. http://nobelprize.org/nobel_prizes/medicine/laureates/2002/sulston-lecture.pdf
* [http://www.nytimes.com/2006/11/21/science/21find.html Nematodes With a Craving for Nicotine]
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