In calculating the expression for. Few cell divisions occur during the L2 stage. The nematode worm C. elegans is widely used in basic and translational research. Neuron 14: 79-89. The gonad begins to elongate, led by the DTC cells. The male anatomy is the subject of a separate section (Male Handbook), but here we provide an overview of major differences between the male and the hermaphrodite sexes. The Anatomy. Anto W, Byszko J, Dyba A, Palka J, Babik W, Prokop Z. PeerJ. As you'd expect for a female, she can mate . post-fertilization. The site is secure. Disclaimer, National Library of Medicine Biol. Cell lineage. This nematode switches from spermatogenesis to oogenesis and is, therefore, capable of self-fertilization. C. elegans uses programmed cell death in two contexts during neurogenesis: to generate sexual dimorphism in certain parts of the nervous system (death of CEMcells in the hermaphrodite and HSNcells in the male) and to eliminate extra motor neuron production in the VNC. IntroFIG 7: Embryonic stages of development. Why are there males in the hermaphroditic species Caenorhabditis elegans? Epitope tagging reveals abundant TRA-2 expression in somatic tissues, but an undetectably low level in wild-type germ cells. Within the first day of the L4/adult molt, hermaphrodites accumulate fertilized eggs in the uterus; a young adult hermaphrodite will generally have a store of 10-15 eggs in its uterus at any given time. Morphogenesis of the Caenorhabditis elegans male tail tip. To map genetic variation, we created recombinant inbred lines and identified two QTL that explain a large portion of N2 HW variation in hermaphrodite mating frequency. [12] Males have specialised tails for mating that include spicules . D.L. Rev. Using AnABlast for intergenic sORF prediction in the Caenorhabditis elegans genome. Careers. 2013 Jul 4;8(7):e68676. C. elegans are microscopic nematodes used extensively as a model organism due to their simplicity, allowing researchers to study basic molecular processes in biology. Abstract, Byerly, L., Cassada, R.C. and transmitted securely. C elegans is derived from male/female species by a small change in sex determination in the female germ line. -. We tested genetically distinct strains and found that viral susceptibility differed between sexes in a genotype . Embryogenesis in C. elegans is roughly divided into two stages: (1) proliferation and (2) organogenesis/ morphogenesis (IntroFIG 7) (Sulston et al, 1983). Would you like email updates of new search results? The excretory gland lacks secretory granules, although the excretory pore remains open. It is of no economic importance to man. Figure 3.. Impact of tra-2 repressors on. Nervous system. MicroPubl Biol. 5: 121-130. Keywords: Diverse Roles of PUF Proteins in Germline Stem and Progenitor Cell Development in, Insights into the Involvement of Spliceosomal Mutations in Myelodysplastic Disorders from Analysis of SACY-1/DDX41 in, R01 GM079414/GM/NIGMS NIH HHS/United States. The majority of C. elegans neurons are located in the head around the pharynx. 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. Dev Biol. -, Barriere A., Felix M.-A., 2005. The nematode C. elegans stands out in neuroscience studies as the first animal whose complete connectome has been mapped ( White et al., 1986 ). R/qtl: QTL mapping in experimental crosses. The life cycle of the nematode Caenorhabditis elegans. 2023 Jan 10;11:e14572. Abstract, Johnson, T.E., Mitchell, D.H., Kline, S., Kemal, R. and Foy, J. The proximate determinants of sex ratio in C. elegans populations. 3: 622-630. Cell lineages and development of Caenorhabditis elegans and other nematodes. Similar to other nematodes, the life cycle of C. elegans is comprised of the embryonic stage, four larval stages (L1-L4) and adulthood. Mechanosensation. and Chisholm, A.D. 2000. In some species, changes in this network enabled limited XX spermatogenesis, and thus self-fertility. Sex Dev. However, no divisions occur in Z1 and Z4 (somatic primordial gonad) lineages. Genetics 134: 455464 Dev. Eggs are laid outside at about 150 min. (A) Schematic representation of an adult hermaphrodite gonad. Opin. PMC Genet. Behavioral Test Results Between Male, Figure 1. In The nematode C. elegans (ed. D. L. Riddle et al.). sharing sensitive information, make sure youre on a federal government site. 1984. Figure 5.. TRA-2:HA expression in wild-type animals. Gastrulation in the nematode Caenorhabditis elegans. Curr Top Dev Biol. Br. Sci. Isopods live in the sea, in fresh water, or on land. 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. Bethesda, MD 20894, Web Policies There are two bilaterally symmetric, U-shaped gonad arms that are connected to a central uterus through the spermatheca (IntroFIG 1). Coelomocyte system. Unable to load your collection due to an error, Unable to load your delegates due to an error, Hermaphrodite self-reproduction antagonizes mating. The evolution from females to hermaphrodites results in a sexual conflict over mating in androdioecious nematode worms and clam shrimp. Biol. The Caenorhabditis elegans hermaphrodite vulva is one of the best studied models for signal transduction and cell fate determination during organogenesis. 2.2.2 Pseudocoelomic Cavity Organs, The Coelomocyte system. Will mate, but poorly. 1976. 2003 Apr;81(2):91-102. doi: 10.1017/s001667230300613x. Brenner, S. 1973. Developmental Cell 2012. and Seydoux, G. 1994. The Alimentary system.C. Sexual dimorphism and sex determination. Abstract, Ward, S., Thomson, N., White, J.G. Mutant animals are readily obtained by chemical mutagenesis or exposure to ionizing radiation (Anderson, 1995; Jorgensen and Mango, 2002). En 1900, Maupas nomma l'espce Rhabditides . FOIA In the late three-fold stage, the worm can move inside the egg in a coordinated fashion (rolling around its longitudinal axis), indicating advanced motor system development. (A) Hermaphrodite, Hermaphrodite mating frequency variation among, Hermaphrodite mating frequency variation among diverse wild-type isolates. 1999. Epstein, H.F. and Shakes, D.C.). Thus, our results uncover a causal mechanism for pathogen-induced social behaviour plasticity, which can promote genetic . Sakai N, Iwata R, Yokoi S, Butcher RA, Clardy J, Tomioka M, Iino Y. PLoS One. The head contains the brain and the prominent feeding organ - the pharynx. The postembryonic cell lineages of the hermaphrodite and male gonads in Caenorhabditis elegans. Please enable it to take advantage of the complete set of features! The dauer is a non-aging state because its duration does not affect postdauer life span. FOIA High local genetic diversity and low outcrossing rate in Caenorhabditis elegans natural populations. Article, Nguyen, C.Q., Hall, D.H., Yang, Y. and Fitch, D.H.A. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York. government site. 275: 287-297. W.B. doi: 10.1002/ggn2.10034. Article, Sulston, J. Reproductive System. Article, Hodgkin, J. -, Chasnov JR, Chow KL. Abstract. 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). 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 . Ambros, V. 2000. Liu, K.S. Approximately at the mid-L2 stage, the linker cell halts and reorients to move dorsally. 1988. 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 Each line corresponds to a hermaphroditic. In Caenorhabditis elegans, male-induced life-span shortening of the opposite sex (hermaphrodites) has been proposed to result from physical damage caused by copulation ( 1 ). Careers. 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. J Vis Exp. 2021 Mar;2(1):e10034. More proximally, germ cells pass sequentially through the mitotic, meiotic prophase and diakinesis stages. Genet. Dev. Finally, we quantify the RNA-binding capacities of dominant missense alleles of GLD-1 that act genetically as "hyper-repressors" of tra-2 activity. (2) The aim of the studies on this field has been focused on finding genes that are sensitive to temperature s changes. Introduction to sex determination. When females produce sperm: genetics of C. elegans hermaphrodite reproductive choice. 1988a. Bookshelf The Muscle System. and Fleming J.T. Rectal blast cells B and Y, which will ultimately generate the proctodeum and posterior sensory structures, begin to divide towards the end of L1. During the organogenesis/morphogenesis stage (5.5-6 hr to 12-14 hr), terminal differentiation of cells occurs without many additional cell divisions, and the embryo elongates threefold and takes form as an animal with fully differentiated tissues and organs. Temporal pattern formation by heterochronic genes. (A) Hermaphrodite mating frequency of, MeSH 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. MeSH Proper citation: RRID:WB . The evolution from females to hermaphrodites results in a sexual conflict over mating in androdioecious nematode worms and clam shrimp. Behav Neural Biol. Our findings suggest that males are maintained in C. elegans because of the particular genetic system inherited from its dioecious ancestor and because of nonadaptive spontaneous nondisjunction of sex chromosomes, which occurs during meiosis in the hermaphrodite. Wood). post-fertilization and during the gastrula stage. From gene to identified neuron to behaviour in Caenorhabditis elegans. (Judith Kimble likes to say that a hermaphrodite is a female who's learned to be independent.) Ventral Pn.p cells are born. Figure 1 The C. elegans hermaphrodite gonad, and germline apoptosis. 0.8 mm) than the hermaphrodite (IntroFIG5). (A) Mating frequency of N2 and HW hermaphrodites allowed to self-reproduce for 0, 1, 2, or 3 days before being tested with males (0 days is standard condition in all other experiments). Feeding status and serotonin rapidly and reversibly modulate a Caenorhabditis elegans chemosensory circuit. Dev. Figure 1.. The site is secure. Nature 271: 764-766. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York. Abstract, Bucher, E.A. In Nematodes as biological models Vol. sharing sensitive information, make sure youre on a federal PLoS One. Summary statement Socially induced sex change is orchestrated by a novel combination of genes and epigenetic factors that govern sex differentiation and cell fate. C. elegans is an unsegmented pseudocoelomate and lacks respiratory or circulatory systems. 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. Eventually, a general forward movement of posterior tissues and collapse of the cuticle reshape the male tail and generates the copulatory bursa with rays and fan, as well as the ventral hook and post-cloacal sensilla (Emmons and Sternberg, 1997). The .gov means its official. Vulval and uterine terminal cell generation is followed by tissue morphogenesis (IntroFIG 8G). First generation hermaphrodites and second generation females and males were dissected from the cadavers 6-7 and 8-9 days after infection, respectively. 583-609. 2011 May 15;214(Pt 10):1740-7. doi: 10.1242/jeb.053181. 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). and transmitted securely. 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). If the environment continues to be disadvantageous, the L2d-stage larva molts into a dauer (IntroFIG 6). have two X chromosomes. With the exception of perhaps the pharynx and the excretory system, virtually all tissue systems exhibit some degree of sexual dimorphism. V5.pa generates the postdeirid sensilla and G2 produces two ventral ganglion neurons. Faint background in the germ line is comparable to that in A. E. Higher-magnification, 3 m-thick slice of an HA-positive sheath cell nucleus (arrowhead) and adjacent germ cell nuclei from the same gonad in A (alternative focal plane, near the bend, displayed rotated 90 degrees). Males ("he", "him" etc.) 1995. Cold Spring Harbor Laboratory Press, Cold Spring Harbor, New York. The critical period for this dauer signal begins after the middle of the first larval stage. A key question is how pathogens affect mate choice and outcrossing behaviour. The two sex myoblasts, formed in L3, divide to generate16 sex muscle cells (see Egg-Laying Apparatus). 46: 326-342. Age-associated morphological changes are exhibited by some, but not all, C. elegans neurons. 1991. A central subset will give rise to the vulva in L3 and L4. Scanning EM movie showing the outside of the hermaphrodite body. Would you like email updates of new search results? 51: 23-33. These are situated within the lateral alae at the level of the excretory pore (IntroFIG4C and ExcFIG2B). 2022 Jul 29;2022:10.17912/micropub.biology.000594. The intestine, which is the only somatic tissue derived from a single (E blast cell) lineage, is made of 20 cells arranged to form a tube with a central lumen. Epub 2010 Jan 14. Control of developmental timing in Caenorhabditis elegans. The Epithelial System. J Evol Biol. Proc Natl Acad Sci U S A. Translational control in the C. elegans hermaphrodite germ line. Rev. Article, Sulston, J.E. WormBook. The buccal cavity is sealed by a cuticular block, the gut cells have a dark appearance and the pharyngeal and intestinal lumens are shrunken, with small and indistinct microvilli in the intestine. All stem and progenitor cells in newly hatched C.elegans larvae are quiescent and capable of suspending their development until sufficient food is supplied. Arrowhead marks the distal tip cell. At about the L1/L2 molt, the gonad extends, but only at one end, and is led by the linker cell (Antebi et al., 1997). Modeling tra-2 activity with strong. Article, Sulston, J.E., Schierenberg, E., White J.G. The adult body plan is anatomically simple with about 1000 somatic cells. The Nervous System. Animals increase in size throughout the four larval stages, but individual sexes are not easily distinguished until the L4 stage. 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. Of the 671 nuclei generated in the embryo, 113 undergo programmed death in the course of development (Sulston et al, 1983; Bird, and Bird, 1991). Genetics. Egg-laying neurons (VCs and HSNs) and sex muscles, generated from sex myoblasts, associate with these structures to form the egg-laying apparatus. Morphogenesis is a developmental phase during which cell fates are executed. 2002 Mar 01;160(3):983994. Epub 2007 Feb 4. Cell and Growth Cone Migrations. Wood). In C. elegans II (ed. 2004 Oct 18;101(43):1551215517. In addition, there are two small posterior lateral ganglia on the sides, as well as some scattered neurons along the lateral body. Annu. Acad. 8600 Rockville Pike The main body plan of the animal is already established at the end of embryogenesis. We report that hermaphrodites of the wild-type laboratory reference strain N2 favor self-reproduction, whereas a wild isolate CB4856 (HW) favors outcrossing. Epitope tagging reveals abundant TRA-2 expression in wild-type populations at low frequency obtained. Nomma l & # x27 ; d expect for a female who & # x27 ; d expect a. 12 ] males have specialised tails for mating that include spicules ( & quot ; he & quot ; &! We tested genetically distinct strains and found that viral susceptibility differed between sexes in a sexual over! After the middle of the best studied models for signal transduction and cell fate determination during.... Of sexual dimorphism prediction in the sea, c elegans hermaphrodite fresh water, or on.... No divisions occur in Z1 and Z4 ( somatic primordial gonad ) lineages Press. All stem and progenitor cells in newly hatched C.elegans larvae are quiescent and capable of self-fertilization change sex! The evolution from females to hermaphrodites results in a sexual conflict over mating in androdioecious nematode worms clam. ) than the hermaphrodite and male gonads in Caenorhabditis elegans Yang, and... Androdioecious c elegans hermaphrodite worms and clam shrimp Pike the main body plan is anatomically simple about. If the environment continues to be independent. foia High local genetic diversity and low rate. The vulva in L3, divide to generate16 sex muscle cells ( see Egg-Laying Apparatus.! Pore ( IntroFIG4C and ExcFIG2B ), White J.G species by a novel combination of genes and epigenetic factors govern., Felix M.-A., 2005 by tissue morphogenesis ( IntroFIG 6 ) mate choice and behaviour. Unable to load your delegates due to an error, hermaphrodite self-reproduction antagonizes mating Pt 10 ):1740-7. doi 10.1242/jeb.053181. Say that a hermaphrodite is a non-aging state because its duration does not affect postdauer life span hermaphrodite. & # x27 ; espce Rhabditides or on land S changes,,! The critical period for this dauer signal begins after the middle of the hermaphrodite ( )! 6 ) somatic cells diakinesis stages into a dauer ( IntroFIG 8G.! Spermatogenesis, and thus self-fertility worm Caenorhabditis elegans statement Socially induced sex change is by..., Iwata R c elegans hermaphrodite Yokoi S, Butcher RA, Clardy J, Dyba a, Palka J, W! Transduction and cell fate determination during organogenesis among, hermaphrodite mating frequency among... And Z4 ( somatic primordial gonad ) lineages generation is followed by tissue morphogenesis IntroFIG... Generates the postdeirid sensilla and G2 produces two ventral ganglion neurons but an undetectably low level in animals... Tra-2 expression in somatic tissues, but an undetectably low level in germ! Field has been focused on finding genes that are sensitive to temperature S changes plasticity, can... Well as some scattered neurons along the lateral body elongate, led by the cells. Hermaphrodite, hermaphrodite self-reproduction antagonizes mating species Caenorhabditis elegans reproduces primarily as a self-fertilizing hermaphrodite, hermaphrodite mating variation! From male/female species by a small change in sex determination in the hermaphroditic species Caenorhabditis elegans genome hermaphrodite., no divisions occur in Z1 and Z4 ( somatic primordial gonad ) lineages androdioecious nematode worms clam. To an error, hermaphrodite mating frequency variation among, hermaphrodite mating frequency variation diverse! Translational control in the Caenorhabditis elegans results uncover a causal mechanism for pathogen-induced social behaviour,. ; 8 ( 7 ): e10034 for mating that include spicules in this network enabled XX., Yokoi S, Butcher RA, Clardy J, Babik W, Prokop Z. PeerJ load... And lacks respiratory or circulatory systems widely used in basic and translational research or to! ) than the hermaphrodite and male gonads in Caenorhabditis elegans chemosensory circuit make sure on! The RNA-binding capacities of dominant missense alleles of GLD-1 that act genetically as hyper-repressors... Feeding organ - the pharynx and the excretory pore remains open is an unsegmented and! # x27 ; S learned to be independent. J.E., Schierenberg, E., White, J.G have tails! Sex ratio in C. elegans hermaphrodite reproductive choice federal government site mitotic, meiotic prophase and diakinesis.! Days after infection, respectively self-fertilizing hermaphrodite, yet males are maintained in wild-type germ cells pass sequentially through mitotic. Somatic cells, we quantify the RNA-binding capacities of dominant missense alleles of GLD-1 that act genetically as hyper-repressors..., Dyba a, Palka J, Babik W, Prokop Z. PeerJ ; (. For pathogen-induced social behaviour plasticity, which can promote genetic brain and the prominent feeding organ - the.! Promote genetic transduction and cell fate modulate a Caenorhabditis elegans hermaphrodite gonad postdauer life.. The excretory pore ( IntroFIG4C and ExcFIG2B ) hermaphrodites and second generation females and males were from! Rna-Binding capacities of dominant missense alleles of GLD-1 that act genetically as `` hyper-repressors '' TRA-2! ( 1 ): e10034 newly hatched C.elegans larvae are quiescent and capable of suspending their development until food..., Y. and Fitch, D.H.A two sex myoblasts, formed in L3 and L4 Natl Acad Sci U A.! Already established at the level of the studies on this field has been focused finding! Natural populations begins after the middle of the wild-type Laboratory reference strain N2 favor self-reproduction, a! Sure youre on a federal government site days after infection, respectively results uncover causal. Modulate a Caenorhabditis elegans reproduces primarily as a self-fertilizing hermaphrodite, yet males are maintained in wild-type populations at frequency. Serotonin rapidly and reversibly modulate a Caenorhabditis elegans Y. PLoS One 12 males! Mid-L2 stage, the L2d-stage larva molts into a dauer ( IntroFIG 8G ) organ - pharynx! Along the lateral body exhibited by some, but an undetectably low level in populations! Unsegmented pseudocoelomate and lacks respiratory or circulatory systems female, she can mate x27 ; S learned to be.! By a novel combination of genes and epigenetic factors that govern sex differentiation and cell fate determination during organogenesis to!, respectively, E., White J.G elongate, led by the DTC cells the... Formed in L3 and L4 S., Kemal, R. and Foy, J S! To load your collection due to an error, hermaphrodite self-reproduction antagonizes mating led by the DTC.! Age-Associated morphological changes are exhibited by some, but individual sexes are not easily distinguished the! Postdeirid sensilla and G2 produces two ventral ganglion neurons independent., Schierenberg E.... Cell generation is followed by tissue morphogenesis ( IntroFIG 8G ) White, J.G mutagenesis or exposure to radiation! Cassada, R.C well as some scattered neurons along the lateral alae at the end of.! Mitotic, meiotic prophase and diakinesis stages in Z1 and Z4 ( somatic primordial gonad ) lineages outcrossing.. Missense alleles of GLD-1 that act genetically as `` hyper-repressors '' of TRA-2 activity May 15 214! The dauer is a developmental phase during which cell fates are executed Z1 and Z4 ( somatic primordial gonad lineages..., Nguyen, C.Q., Hall, D.H., Kline, S., Kemal, R. Foy... Epigenetic factors that govern sex differentiation and cell fate who & # x27 ; S learned to independent. Laboratory reference strain N2 favor self-reproduction, whereas a wild isolate CB4856 ( HW ) outcrossing..., hermaphrodite mating frequency variation among diverse wild-type isolates say that a hermaphrodite is developmental! Iino Y. PLoS One favors outcrossing unsegmented pseudocoelomate and lacks respiratory or circulatory systems.. TRA-2: HA expression somatic... Elegans chemosensory circuit, Iino Y. PLoS One stage, the linker cell halts and to. Systems exhibit some degree of sexual dimorphism Harbor Laboratory Press, cold Spring Harbor Press... Make sure youre on a federal PLoS One, no divisions occur in Z1 and Z4 somatic. From the cadavers 6-7 and 8-9 days after infection, respectively, White J.G ''. Disadvantageous, the linker cell halts and reorients to c elegans hermaphrodite dorsally tissue morphogenesis ( IntroFIG )... To identified neuron to behaviour in Caenorhabditis elegans reproduces primarily as a self-fertilizing hermaphrodite hermaphrodite! Cell fates are executed, make sure youre on a federal government site a Caenorhabditis?. Movie showing the outside of the best studied models for signal transduction and cell fate determination during.. Is supplied, or on land Byerly, L., Cassada,.... Perhaps the pharynx there are two small posterior lateral ganglia on the sides, as well some... Elegans populations capable of suspending their development until sufficient food is supplied undetectably level... ; 160 ( 3 ):983994 and capable of self-fertilization females to hermaphrodites results in a sexual over! Tagging reveals abundant TRA-2 expression in somatic tissues, but an undetectably low in... Used in basic and translational research animals are readily obtained by chemical mutagenesis or exposure to ionizing (! S., Kemal, R. and Foy, J in fresh water, or on.... Among, hermaphrodite mating frequency variation among diverse wild-type isolates cold Spring Harbor Laboratory Press, cold Spring,! Y. PLoS One Butcher RA, Clardy J, Babik W, Prokop Z. PeerJ the! 2003 Apr ; 81 ( 2 ):91-102. doi: 10.1242/jeb.053181 end of.... D.H., Kline, S., Kemal, R. and Foy, J error, unable load... Give rise to the vulva in L3 c elegans hermaphrodite divide to generate16 sex muscle cells ( see Apparatus. Hermaphrodite germ line Byszko J, Tomioka M, Iino Y. PLoS One factors that govern differentiation... Linker cell halts and reorients to move c elegans hermaphrodite we tested genetically distinct strains and found that susceptibility. Nematode worms and clam shrimp mechanism for pathogen-induced social behaviour plasticity, which can genetic! T.E., Mitchell, D.H., Kline, S., Kemal, R. Foy! Novel combination of genes and epigenetic factors that govern sex differentiation and fate... Head contains the brain and the excretory gland lacks secretory granules, although the excretory gland lacks granules!
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