and Russell, R.L. On the lateral sides, the hypodermis is interrupted by the syncytial row of seam cells which form alae on the cuticle surface during certain developmental stages (IntroFIG 3C) (see Seam Cells). doi: 10.7717/peerj.14572. Mechanical forces shaping individual cells play a key role during tissue morphogenesis. Age. Males can successfully mate with a hermaphrodite for 6 days after their last larval molt and can father approximately 3000 progeny (Hodgkin, 1988). Various tissues open to the outside through this cuticle (IntroFIG 3). Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York. eCollection 2021. C. elegans is an unsegmented pseudocoelomate and lacks respiratory or circulatory systems. 2014 Jun 2;14:116. doi: 10.1186/1471-2148-14-116. C. elegans all have 5 autosomes. doi: 10.17912/micropub.biology.000594. Figure 4.. PLoS One. Tail tip hypodermal cells remodel, generating the rounded tail of the adult (IntroFIG 5E) (Nguyen et al., 1999). The gonad of the dauer is arrested at the L2 stage (IntroFIG 8E) (Cassada and Russell, 1975; Riddle, 1988; Sulston, 1988). This general body plan does not change during postembryonic development. Figure 1.. 2002 Feb 5;12(3):R111-20. The dauer larva. 2019 Jun 6;(148):10.3791/59783. Before Please enable it to take advantage of the complete set of features! The C elegans hermaphrodite is basically a female, except that she makes some sperm of her own. Genet. B. Anti-HA antibodies detect TRA-2B:HA in lysates from XX, Each line corresponds to a hermaphroditic (solid) or feminized (dashed) genotype in Table 1. 2006 Jan 01;:114. A conserved RNA-binding protein that regulates sexual fates in the C. elegans hermaphrodite germ line Beilin Zhang Download Free PDF View PDF Biochemical and Biophysical Research Communications Characterization of RAB-like4, the first identified RAB-like protein from Trypanosoma cruzi with GTPase activity 2005 Rosane Silva Download Free PDF 16: R726R735 The molecular underpinnings of fertility: Genetic approaches in. Together, our findings demonstrate that C. elegans hermaphrodites actively regulate the choice between selfing and crossing, highlight the existence of natural variation in hermaphrodite choice, and lay the groundwork for molecular dissection of this evolutionarily important trait. 1988. PLoS Biol. 2005 Jan;3(1):e6. In the male germline, which produces only sperm, meiosis begins during L3 stage. The metazoan roundworm has a long history as post-fertilization. Bookshelf -, Broman K. W., Wu H., Sen S., Churchill G. A., 2003. Abstract. Curr. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York. Sensory regulation of male mating behavior in Caenorhabditis elegans. In Nematodes as biological models Vol. Molecular approaches to aggregation behavior and social attachment. C. elegans are microscopic nematodes used extensively as a model organism due to their simplicity, allowing researchers to study basic molecular processes in biology.Most C. elegans are hermaphrodites, possessing two X chromosomes and the ability to reproduce asexually, but approximately 0.1% are males, arising due to a spontaneous loss of an X chromosome. Each line corresponds to a hermaphroditic. 2018 Sep;10(3):e43. We normalized the results from IP samples as rpm total reads matched to C. elegans genome. . IntroFIG 4C: Paired sensilla of the anterior deirid, posterior deirid, and phasmid, left lateral side. Selection and maintenance of androdioecy in Caenorhabditis elegans. Tanner D, Carigo D, Sevilla C, Lewis M, Harris G. MicroPubl Biol. Article, Sulston, J. History of research on C. elegans and other free-living nematodes as model organisms. The molecular basis for this astonishing transformation remains broadly unknown. Two much narrower openings on the lateral sides of the tail whip exist for the phasmid sensilla at the junction of the seam cells and the tail hypodermis (IntroFIG 4C). Chapter 5. pp 123-155. The nematode worm C. elegans is widely used in basic and translational research. In order to evaluate the behavioral sex differences in C. elegans, we expanded upon existing literature and compared spontaneous movement, sensitivity to mechanosensation, and sensitivity to chemosensation between males and hermaphrodites. It is an important model system for biological research in many fields including genomics, cell biology, neuroscience and aging (http://www.wormbook.org/). 2004 Oct 18;101(43):1551215517. C. elegans commonly exist as self-fertiliz-ing hermaphrodites, although a small percentage (<1%) of the microscopic worms are males due to altered X chromosome-au-tosomal ratios. The Epithelial System. The Caenorhabditis elegans adult hermaphrodite germline is surrounded by a thin tube formed by somatic sheath cells that support germ cells as they mature from the stem-like mitotic state through meiosis, gametogenesis, and ovulation. 1995 Mar;12(2):346-58 Abstract, Johnson, T.E., Mitchell, D.H., Kline, S., Kemal, R. and Foy, J. FOIA When females produce sperm: genetics of C. elegans hermaphrodite reproductive choice. B Biol. 15492-15495, December 1998 Genetics Two functionally dependent acetylcholine subunits are encoded in a single Caenorhabditis elegans operon MILLET TREININ*,BOAZ GILLO,LILACH LIEBMAN, AND MARTIN CHALFIE* *Department of Biological Sciences, Columbia University, New York, NY 10027; and Department of Physiology and The Kuhne Minerva . The length of the animal at each stage is marked next to the stage name in micrometers (m). official website and that any information you provide is encrypted Sex differences in decision-making: Identifying multisensory behavioral differences in males and hermaphrodites. 1988. Before Would you like email updates of new search results? Among its many advantages for study are its short life cycle, compact genome, stereotypical development, ease of propagation and small size. Figure 5. An increasing concentration of purified, recombinant MBP-tagged GLD-1 or mutant variants was incubated with a fluorescein labeled RNA oligonucleotide comprising a fragment from the. Figure 1 The C. elegans hermaphrodite gonad, and germline apoptosis. Cell and Growth Cone Migrations. Biol. Sexual dimorphism in Caenorhabditis elegans stress resistance. Disclaimer, National Library of Medicine 1992 Nov;58(3):211-21. doi: 10.1016/0163-1047(92)90496-q. Similar to other nematodes, C. elegans has an unsegmented, cylindrical body shape that is tapered at the ends (IntroFIG 1; IntroMOVIE 1). The nematode C. elegans stands out in neuroscience studies as the first animal whose complete connectome has been mapped ( White et al., 1986 ). These variants bind RNA more weakly in vitro than does wild-type GLD-1. The site is secure. Spatiotemporal restriction of signaling plays a critical role in animal development and tissue homeostasis. Genetic regulation of embryonic sex differentiation is well established in vertebrates but . Disclaimer, National Library of Medicine For sperm motility imaging, male worms carrying . Bull. For detailed review of anatomic structures in male, refer to Male Handbook. Six male sex myoblasts are generated. Article, Rankin, C.H. Careers. -, Chao MY, Komatsu H, Fukuto HS, Dionne HM, Hart AC. The arrow denotes where delamination of the cuticle surface near the vulva has occurred. The aim of this study was to investigate the spatial and temporal distribution of Exhippolysmata oplophoroides and relate its abundance to various environmental variables (bottom water temperature and salinity, texture of sediment and percentage of dissolved organic matter). The end of each larval stage is marked with a molt, during which a new, stage-specific cuticle is synthesized and the old one is shed. Using AnABlast for intergenic sORF prediction in the Caenorhabditis elegans genome. The cells of the nervous system are organized into ganglia in the head and tail. Nat. The pharynx is separated from the outer tube of tissues and pseudocoelom by its own basal lamina (IntroFIG 2B-D). elegans ( hermaphrodite)(male), 5XX,5 X, . Riddle et al.). The .gov means its official. Academic Press, California. First generation hermaphrodites and second generation females and males were dissected from the cadavers 6-7 and 8-9 days after infection, respectively. The nematode C. elegans is one of the most important model organisms for understanding neurobiology. eCollection 2020. Eggs are laid outside at about 150 min. Single-Protein-Specific Redox Targeting in Live Mammalian Cells and C. elegans. Out of 1090 somatic cells generated during hermaphrodite development, 131 undergo programmed cell death at characteristic times (Driscoll, 1995). doi: 10.1073/pnas.0403369101. 10: 428-433. D.L. The outer tube (body wall) consists of cuticle, hypodermis, excretory system, neurons, and muscles, and the inner tube comprises the pharynx, intestine, and, in the adult, gonad. The Reproductive system. 2010 Mar;23(3):539-56. doi: 10.1111/j.1420-9101.2009.01919.x. 2021 Oct 19;2021:10.17912/micropub.biology.000475. The .gov means its official. 207: 86-106. The role of hermaphrodites in the experimental evolution of increased outcrossing rates in Caenorhabditis elegans. The oocytes are fertilized by the sperm in spermatheca. The Caenorhabditis elegans hermaphrodite vulva is one of the best studied models for signal transduction and cell fate determination during organogenesis. Lond. Genet. C. elegans is a powerful genetic model whose optical transparency enables the observation of biological processes in real-time and the determination of the subcellular localization of fluorescently tagged proteins of interest during any stage of the C. elegans life cycle. The reproductive system consists of a single-armed gonad (IntroFIG 5C; Male Somatic Gonad and Germline) that opens to the exterior at the cloaca (anus) via a modified rectal epithelial chamber called the proctodeum (IntroFIG 5D) (see Proctodeum). The gonad continues to grow posteriorly along the ventral midline, and the vas deferens and the seminal vesicle differentiate. 2010 Feb;53(2):83-102. doi: 10.1139/g09-090. In C. elegans, one of these selfing species, the global sex-determining gene tra-2 is regulated in germ cells by a conserved RBP, GLD-1, via the 3' untranslated region (3'UTR) of its transcript. These two strains were isolatedfrom the wild in different places and so have undergone extensive genetic drift since they lastshared a common ancestor. Modeling tra-2 activity with strong. The hermaphrodite is a modified female that in the fourth larval stage makes and stores sperm to be used later to fertilize oocytes produced within the gonad of the same animal after spermatogenesis is finished. Figure 6.. TRA-2 expression in fog-2 and gld-1 null mutants. 3.2.6 Adult. During the second half of L1, somatic gonad precursors Z1 and Z4 produce 12 cells in the hermaphrodite (IntroFIG 8C). 2022 May 19;2022:10.17912/micropub.biology.000569. Before Federal government websites often end in .gov or .mil. (2) The aim of the studies on this field has been focused on finding genes that are sensitive to temperature s changes. Unable to load your collection due to an error, Unable to load your delegates due to an error, Hermaphrodite self-reproduction antagonizes mating. Gastrulation in the nematode Caenorhabditis elegans. doi: 10.17912/micropub.biology.000475. The genetics of behavior. Meiosis in the germ line begins at L3/L4 molt in the proximal arms of the gonad, and the germ cells differentiate into mature sperm. Its completely mapped neural connectome of 302 neurons and fully characterized and stereotyped development have made it a prototype for understanding nervous system structure, development, and function. Sakai N, Iwata R, Yokoi S, Butcher RA, Clardy J, Tomioka M, Iino Y. PLoS One. The resulting diploid zygotes are stored in the uterus and laid outside thorough the vulva, which protrudes at the ventral midline (see Reproductive System: Somatic Gonad, Germ Line and Egg-laying Apparatus). Wood). Chapter 4. pp 81-122. All observations and measurements were performed within one week of harvest. Abstract, Ward, S., Thomson, N., White, J.G. The postembryonic cell lineages of the hermaphrodite and male gonads in Caenorhabditis elegans. 2002. Br. Wood). To determine the role of males in C. elegans, we develop a mathematical model for the genetic system of hermaphrodites that can either self-fertilize or be fertilized by males and we perform laboratory . This paper describes the fabrication and use of a minimalist microfluidic chip for performing microinjections. Sem. Post-embryonic development in the ventral cord of Caenorhabditis elegans. 1977. FOG-2, a novel F-box containing protein, associates with the GLD-1 RNA binding protein and directs male sex determination in the C. elegans hermaphrodite germline. The .gov means its official. 54: 78-92. Anderson, P. 1995. Male. Abstract, Slack, F. and Ruvkun, G. 1997. The distal gonad arms continue their migration centripetally along the dorsal body wall muscles, and by the L4/adult molt, they complete their trajectory close to midline (Antebi et al, 1997). The approximate level of each cross section is labeled in, IntroFIG 3: Scanning electron microscopy (SEM) images of adult, Scanning electron micrograph (SEM) of adult, Transmission electron micrograph (TEM), transverse section of the nubbin of a posterior deirid sensillum. Kreuzinger-Janik B, Gansfort B, Ptatscheck C. Sci Rep. 2022 Apr 2;12(1):5578. doi: 10.1038/s41598-022-09631-w. Diversity in mating behavior of hermaphroditic and male-female Caenorhabditis nematodes. Reproductive System. An official website of the United States government. 2022 Jun 29;23(13):7202. doi: 10.3390/ijms23137202. Article, Sulston, J.E. The excretory gland lacks secretory granules, although the excretory pore remains open. 2017 May 1;117(5):1911-1934. doi: 10.1152/jn.00555.2016. Abstract, Cassada, R.C. Post-embryonic development is triggered by feeding after hatching. -, Goodman MB. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York. 2020 Dec 8;36(19):4827-4832. doi: 10.1093/bioinformatics/btaa608. The Excretory System. 2.2.2 Pseudocoelomic Cavity Organs, The Coelomocyte system. Brenner, S. 1973. -, Barriere A., Felix M.-A., 2005. 56: 110-156. The processes from most neurons travel in either the ventral or dorsal nerve cord and project to the nerve ring (NR) in the head which constitutes the major neuropil in the animal (IntroFIG 2C)(see Nervous System). See this image and copyright information in PMC. The lips also contain papillae for 6 inner labial (IL) sensilla and small bumps for 6 outer labial (OL) sensilla, as well as 4 cephalic (CEP) sensilla (IntroFIG 4A and IntroTABLE 1). 29:269-271. Dev. Von Ehrenstein, G. and Schierenberg, E. 1980. C. elegans are microscopic nematodes used extensively as a model organism due to their simplicity, allowing researchers to study basic molecular processes in biology. 2007 Nov 6;17(21):1847-57. doi: 10.1016/j.cub.2007.09.011. (C-D) Unmated QG2288 hermaphrodites with no apparent physical damage. and Brenner, S. 1975. doi: 10.1371/journal.pbio.0030006. At hatching, males are already distinguishable from hermaphrodites because of the more posterior location of one ventral coelomocyte, the larger size of the nuclei of two rectal cells (B and Y), the absence of hermaphrodite specific neurons (HSNs) that undergo programmed cell death during embryogenesis, and the presence of CEM neurons. Proper citation: RRID:WB . Wood). With the exception of perhaps the pharynx and the excretory system, virtually all tissue systems exhibit some degree of sexual dimorphism. A C. elegans-specific GLD-1 cofactor, FOG-2, is also required for hermaphrodite sperm fate, but how it modifies GLD-1 function is unknown. If the environment continues to be disadvantageous, the L2d-stage larva molts into a dauer (IntroFIG 6). The L2d larva retains the potential to form either a dauer larva or an L3 larva, depending on the persistence of the dauer inducing environmental parameters (Riddle, 1988). and Anness, M.A.R. The intestinal contents are excreted to the outside via a rectal valve that connects the gut to the rectum and anus. Scale bar: 100 m. Embryology. 2010 Mar;23(3):539-56. doi: 10.1111/j.1420-9101.2009.01919.x. To characterize underlying neural mechanisms, we show that N2 hermaphrodites deficient in mechanosensation or chemosensation (e.g., mec-3 and osm-6 mutants) exhibit high mating frequency, implicating hermaphrodite perception of males as a requirement for low mating frequency. The site is secure. Similar to other nematodes, the life cycle of C. elegans is comprised of the embryonic stage, four larval stages (L1-L4) and adulthood. The germ line precursors Z2 and Z3 also start to divide. Abstract, Kimble, J. and Hirsh, D. 1979. Vulval precursor fates are specified, and committed cells divide to generate vulval terminal cells by early L4. 95, pp. Summary statement Socially induced sex change is orchestrated by a novel combination of genes and epigenetic factors that govern sex differentiation and cell fate. 5: 121-130. Evolutionary history of gld-1 and. First cleavage occurs at about 40 min. 21. pp. The dauer larva, a postembryonic developmental variant of the nematode Caenorhabditis elegans. HHS Vulnerability Disclosure, Help 51: 23-33. Dev. Here we investigated the effect of Orsay virus on the mating dynamics of the androdiecious (male and hermaphrodite) nematode Caenorhabditis elegans. Evolution of plant breeding systems. Unable to load your collection due to an error, Unable to load your delegates due to an error. and Bird J. Molecular lesion: deletion of entire unc-10 coding region. The nervous system has 89 additional neurons that include several classes of tail sensilla: the rays, which extend from the tail and lie in a cuticular fan (see Rays); the hook (see Hook); and the post-cloacal sensilla, which are located on the ventral exterior of the tail (see PCS). 1997. Article. The combined data suggest that NDPK enzymes control the availability of surface receptors to regulate cell-sensing cues during cell migration. It is an important model system for biological research in many fields including genomics, cell biology, neuroscience and aging (http://www.wormbook.org/). These Z2-Z3 divisions occur continuously from L1 through adulthood (Kimble and Hirsh, 1979). Developmental Cell 2012. (A) Hermaphrodite mating frequency of 38 wild-type isolates representing haplotype diversity of. The most profound differences are seen in tissues of the posterior, which bears the male copulatory apparatus. Most C. elegans are hermaphrodites, possessing two X chromosomes and the ability to reproduce asexually, but approximately 0.1% are males, arising due to a spontaneous loss of an X chromosome. Bookshelf 46: 323-353. 28: 23-40. We report that hermaphrodites of the wild-type laboratory reference strain N2 favor self-reproduction, whereas a wild isolate CB4856 (HW) favors outcrossing. Figure 1. Epub 2011 Nov 21. R. Soc. Careers. Abstract, Sulston, J.E. -, Mol Biol Evol. The first muscle twitches are observed at 430 min after the first cell cleavage (between 1.5- and 2-fold stages) (IntroFIG 7). C. elegans can also endure harsh environmental conditions by switching to a facultative diapause stage called the dauer larva which can survive four to eight times the normal 3-week life span (Cassada and Russell, 1975). IntroFIG 7: Embryonic stages of development. In the presence of food, cell divisions resume and the post-embryonic developmental program begins 3 hours after hatching (Ambros, 2000). Connectivity changes in a class of motoneurone during the development of a nematode. eCollection 2023. 1991. Tsukamoto T, Gearhart MD, Kim S, Mekonnen G, Spike CA, Greenstein D. Genetics. In this stage, the germ cell (Z2 and Z3 daughters) divisions continue, approximately quadrupling in number (IntroFIG 8D). See this image and copyright information in PMC. We postulate that a genetic mutation occurred during the evolution of C. elegans hermaphrodites, resulting in the loss of an attracting sex pheromone present in the ancestor of both C. elegans and C. remanei. Caenorhabditis elegans is a small, free-living soil nematode (roundworm) that lives in many parts of the world and survives by feeding on microbes, primarily bacteria (IntroFIG 1). 1976. This suggests the GLD-1-FOG-2 complex regulates uncharacterized sites within tra-2, or other mRNA targets. In sexually reproducing organisms, cells of the germ line form gametes and establish a physical link, an unbroken chain, from generation to generation. Sex Determination in Nematode Germ Cells. In our paradigms, we found that males and hermaphrodites exhibit similar spontaneous movement as well as similar slow and sustained behaviors such as chemotaxis, but differ in quick-response to mechanical and chemosensory stimuli. 2014 May 13;3(4):239262. Sex determination in the Caenorhabditis elegans germ line. After 3 days of incubation in the liquid assay a biofilm was clearly visible by light microscopy on both the head and vulva region of the worms. Three pairs of coelomocytes located in the pseudocoelomic cavity function as scavenger cells that endocytose fluid from the pseudocoelom and are suggested to comprise a primitive immune system in C. elegans (see Coelomocyte System). En 1900, Maupas nomma l'espce Rhabditides . Clipboard, Search History, and several other advanced features are temporarily unavailable. Chapter 12. pp.295-334. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York. Chapter 12. pp 393-412. Biol. This rather small. Male. The strains can be kept as frozen stocks for long periods of time. A sexually conditioned switch of chemosensory behavior in C. elegans. The free-living nematode worm Caenorhabditis elegans reproduces primarily as a self-fertilizing hermaphrodite, yet males are maintained in wild-type populations at low frequency. 2022 Aug 11;17(8):e0272452. Abstract, Wood, W.B. Dev. Male C. elegans larvae initially display the same simple cylindrical body plan as hermaphrodites, but from the L2 stage onward, the shape of their posterior half changes as their sexual organs begin to develop (IntroFIG 5) (Sulston and Horvitz, 1977; Sulston et al., 1980; Nguyen et al., 1999). USA Vol. 1976. 8600 Rockville Pike Also, during the L1 stage, one class of ventral cord motor neurons (DDn) go through complete synaptic reorganization without any cell-shape change. Epub 2004 Dec 28. Biol. Abstract, Jorgensen, E.M. and Mango, S.E. and Chisholm, A.D. 2000. The germ line within the distal gonad arms (ovaries) is syncytial with germline nuclei surrounding a central cytoplasmic core. Behav Neural Biol. At the end of the L2 stage, the animal may enter an arrested state called the dauer larva if the environmental conditions are not favorable for further growth. eCollection 2022. Hermaphrodites (commonly referred to with female pronouns - "she", "her" etc.) A hermaphrodite that self-fertilizes can produce about 300 progeny because of the limited number of sperm, but if mating with a male occurs, the progeny number can increase to 1200-1400. The animal can be maintained in the laboratory where it is grown on agar plates or liquid cultures with E. coli as the food source. From this time onward, the embryonic substages can be defined by specific cell migrations, gain in cell number, and periods of synchronous stem-cell divisions. 24: 1589-1599. nr. All stem and progenitor cells in newly hatched C.elegans larvae are quiescent and capable of suspending their development until sufficient food is supplied. Sci. Morphogenesis is a developmental phase during which cell fates are executed. Despite its simple anatomy, the animal displays a large repertoire of behavior including locomotion; foraging; feeding; defecation; egg laying; dauer larva formation; sensory responses to touch, smell, taste, and temperature; and some complex behaviors like male mating, social behavior, and learning and memory (Rankin, 2002; de Bono, 2003). Of the eight classes of motor neurons (DAn, DBn, VAn, VBn, VCn, ASn, VDn, and DDn) in the adult hermaphrodite ventral cord, five (VAn, VBn, VCn, ASn and VDn) are generated at the end of the L1 stage from 13 precursors (W and P1-P12) (IntroFIG 8A) (Sulston, 1976; Sulston and Horvitz, 1977; Chalfie and White 1988). Clipboard, Search History, and several other advanced features are temporarily unavailable. sharing sensitive information, make sure youre on a federal Isopoda is an order of crustaceans that includes woodlice and their relatives. -, Genetics. At the L4 stage, hermaphrodites have a tapered tail and the developing vulva (white arrowhead) can be seen as a clear half circle in the center of the ventral side. 1988b. In the body, a continuous row of neuron cell bodies lies at the midline, adjacent to the ventral hypodermis. Numbers in blue along the arrows indicate the length of time the animal spends at a certain stage. 2013 Jul 4;8(7):e68676. As in the hermaphrodite, Z1 and Z4 divide, producing 10 somatic gonad precursor cells. Experimental evolution of sperm count in protandrous self-fertilizing hermaphrodites. doi: 10.1002/wdev.136. *. Together, our findings demonstrate that C. elegans hermaphrodites actively regulate the choice between selfing and crossing, highlight the existence of natural variation in hermaphrodite choice, and lay the groundwork for molecular dissection of this evolutionarily important trait. The majority of C. elegans neurons are located in the head around the pharynx. The vulva is another large opening on the ventral side at the midbody (IntroFIG 3D), and the anus forms another ventral opening, just before the tail whip (IntroFIG 3B). (A) Hermaphrodite, Hermaphrodite mating frequency variation among, Hermaphrodite mating frequency variation among diverse wild-type isolates. Mutations in two independent pathways are sufficient to create hermaphroditic nematodes. [12] Males have specialised tails for mating that include spicules . Using serial electron microscopy synapse-level neural maps have been constructed both for adult male and hermaphrodite ( White et al., 1986; Cook et al., 2019 ). WikiZero zgr Ansiklopedi - Wikipedia Okumann En Kolay Yolu Z3 daughters ) divisions continue, approximately quadrupling in number ( IntroFIG 6 ) Isopoda an... As model organisms dauer larva, a postembryonic developmental variant of the best models..., Gearhart MD, Kim S, Mekonnen G, Spike CA, Greenstein D. Genetics New. 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