Germ Cell Fate Determination in C. elegans


Germ cell development is essential for the sexual reproduction of animals. Germ cells are immortal in that they contribute to successive generations, whereas somatic cells perish with the individual. The nematode Caenorhabditis elegans is a leading system for elucidating the developmental mechanisms controlling the germ cell fate determination because this organism is amenable to molecular genetic and mechanistic analysis. This review discusses the molecular mechanisms specifying the fate of germ cells from their journey in the embryo to fertilization.

Key Concepts

  • Germ cell development is essential for the sexual reproduction of animals.
  • Primordial germ cells are formed early in embryogenesis.
  • Germ cell specification happens in the absence of transcription.
  • Germ cell proliferation in the adult is regulated by the somatic niche.
  • Oocyte meiotic maturation is under hormonal control.
  • Gene expression in germline is regulated by a combination of epigenetic and posttranscriptional mechanisms.

Keywords: germ cell; stem cell; germline; cell fate; germ granules; primordial germ cells; oocyte maturation; totipotency

Figure 1. Embryonic origin of the germline. Embryonic germline lineage from the one‐cell stage to the ∼100‐cell stage and corresponding embryo schematics are shown in the lineage tree. Germ plasm is shown in purple, and P granules as red dots. Orange marks transcriptionally quiescent nuclei.
Figure 2. Adult C. elegans gonad. (a) One of the two hermaphrodite gonads arms. The somatic distal tip cell (DTC) caps the gonad. The colours of the germline nuclei indicate developmental stages: light green, mitotic; green crescent, entry into meiosis; green, pachytene and blue, diplotene and diakinesis. P granules (red) localise at the nuclear periphery until diakinesis. Sperm are brown. (b) Male gonad. The single arm of the gonad is capped with two distal tip cells. The colours of the nuclei indicate mitotic, transition and pachytene stages analogous to that of the hermaphrodite shown in (a); dark purple nuclei: primary spermatocytes; brown: sperm.
Figure 3. Signalling in the stem cell niche. LAG‐2 and APX‐1 ligands expressed in the DTC activate GLP‐1 Notch signalling in the germline. This signalling activates transcription of LST‐1 and SYGL‐1, which maintain germline stem cells. GLD‐1 and GLD‐2 function downstream and regulate the initiation of meiosis. Another critical component of the germ cell niche are gap junction channels composed of the innexin proteins, INX‐8 and ‐9 in the soma and INX‐14 and ‐21 in the germline.


Ahringer J and Kimble J (1991) Control of the sperm‐oocyte switch in Caenorhabditis elegans hermaphrodites by the fem‐3 3′ untranslated region. Nature 349: 346–348.

Arur S, Ohmachi M, Nayak S, et al. (2009) Multiple ERK substrates execute single biological processes in Caenorhabditis elegans germ‐line development. Proceedings of the National Academy of Sciences of the United States of America 106: 4776–4781.

Billi AC, Fischer SE and Kim JK (2014) Endogenous RNAi pathways in C. elegans. In: WormBook (ed.) The C. elegans Research Community. WormBook. DOI: 10.1895/wormbook.1.170.1,

Bowerman B, Draper BW, Mello CC and Priess JR (1993) The maternal gene skn‐1 encodes a protein that is distributed unequally in early C. elegans embryos. Cell 74: 443–452.

Brangwynne CP, Eckmann CR, Courson DS, et al. (2009) Germline P granules are liquid droplets that localize by controlled dissolution/condensation. Science 324: 1729–1732.

Burrows AE, Sceurman BK, Kosinski ME, et al. (2006) The C. elegans Myt1 ortholog is required for the proper timing of oocyte maturation. Development 133: 697–709.

Claycomb JM, Batista PJ, Pang KM, et al. (2009) The Argonaute CSR‐1 and its 22G‐RNA cofactors are required for holocentric chromosome segregation. Cell 139: 123–134.

Detwiler MR, Reuben M, Li X, Rogers E and Lin R (2001) Two zinc finger proteins, OMA‐1 and OMA‐2, are redundantly required for oocyte maturation in C. elegans. Developmental Cell 1: 187–199.

Drake M, Furuta T, Suen KM, et al. (2014) A requirement for ERK‐dependent Dicer phosphorylation in coordinating oocyte‐to‐embryo transition in C. elegans. Developmental Cell 31: 614–628.

Eddy EM (2006) Germ plasm and the molecular determinants of germ cell fate. In: eLS. DOI: 10.1038/npg.els.0005960.

Espiritu EB and Rose LS (2013) Caenorhabditis elegans embryo: establishment of asymmetry. In: eLS. DOI: 10.1002/9780470015902.a0001506.pub3.

Farboud B, Nix P, Jow MM, Gladden JM and Meyer BJ (2013) Molecular antagonism between X‐chromosome and autosome signals determines nematode sex. Genes & Development 27: 1159–1178.

Furuhashi H, Takasaki T, Rechtsteiner A, et al. (2010) Trans‐generational epigenetic regulation of C. elegans primordial germ cells. Epigenetics Chromatin 3: 15.

Han T, Manoharan AP, Harkins TT, et al. (2009) 26G endo‐siRNAs regulate spermatogenic and zygotic gene expression in Caenorhabditis elegans. Proceedings of the National Academy of Sciences of the United States of America 106: 18674–18679.

Hansen D and Schedl T (2013) Stem cell proliferation versus meiotic fate decision in Caenorhabditis elegans. Advances in Experimental Medicine & Biology 757: 71–99.

Hubbard EJ, Korta DZ and Dalfó D (2013) Physiological control of germline development. Advances in Experimental Medicine & Biology 757: 101–131.

Johnson CL and Spence AM (2011) Epigenetic licensing of germline gene expression by maternal RNA in C. elegans. Science 332: 1311–1314.

Kershner AM, Shin H, Hansen TJ and Kimble J (2014) Discovery of two GLP‐1/Notch target genes that account for the role of GLP‐1/Notch signaling in stem cell maintenance. Proceedings of the National Academy of Sciences of the United States of America 111: 3739–3744.

Kim S, Spike C and Greenstein D (2013) Control of oocyte growth and meiotic maturation in Caenorhabditis elegans. Advances in Experimental Medicine & Biology 757: 277–320.

Kimble JE and White JG (1981) On the control of germ cell development in Caenorhabditis elegans. Developmental Biology 81: 208–219.

Knight SW and Bass BL (2001) A role for the RNase III enzyme DCR‐1 in RNA interference and germ line development in Caenorhabditis elegans. Science 293: 2269–2271.

Kosinski M, McDonald K, Schwartz J, Yamamoto I and Greenstein D (2005) C. elegans sperm bud vesicles to deliver a meiotic maturation signal to distant oocytes. Development 132: 3357–3369.

Lee MH, Ohmachi M, Arur S, et al. (2007) Multiple functions and dynamic activation of MPK‐1 extracellular signal‐regulated kinase signaling in Caenorhabditis elegans germline development. Genetics 177: 2039–2062.

Liang CG, Su YQ, Fan HY, Schatten H and Sun QY (2007) Mechanisms regulating oocyte meiotic resumption: roles of mitogen‐activated protein kinase. Molecular Endocrinology 21: 2037–2055.

Madl JE and Herman RK (1979) Polyploids and sex determination in Caenorhabditis elegans. Genetics 93: 393–402.

McJunkin K and Ambros V (2014) The embryonic mir‐35 family of microRNAs promotes multiple aspects of fecundity in Caenorhabditis elegans. G3 (Bethesda) 4: 1747–1754.

Mello CC, Draper BW, Krause M, Weintraub H and Priess JR (1992) The pie‐1 and mex‐1 genes and maternal control of blastomere identity in early C. elegans embryos. Cell 70: 163–176.

Nousch M and Eckmann CR (2013) Translational control in the Caenorhabditis elegans germ line. Advances in Experimental Medicine & Biology 757: 205–247.

Pazdernik N and Schedl T (2013) Introduction to germ cell development in Caenorhabditis elegans. Advances in Experimental Medicine & Biology 757: 1–16.

Robertson S and Lin R (2013) The oocyte‐to‐embryo transition. Advances in Experimental Medicine & Biology 757: 351–372.

Rose L and Gönczy P (2014) Polarity establishment, asymmetric division and segregation of fate determinants in early C. elegans embryos. WormBook: 1–43.

Schaner CE, Deshpande G, Schedl PD and Kelly WG (2003) A conserved chromatin architecture marks and maintains the restricted germ cell lineage in worms and flies. Developmental Cell 5: 747–757.

Schisa J (2012) New Insights into the regulation of RNP granule assembly in oocytes. International Review of Cell and Molecular Biology 295: 233–289.

Seth M, Shirayama M, Gu W, et al. (2013) The C. elegans CSR‐1 argonaute pathway counteracts epigenetic silencing to promote germline gene expression. Developmental Cell 27: 656–663.

Seydoux G and Fire A (1994) Soma‐germline asymmetry in the distributions of embryonic RNAs in Caenorhabditis elegans. Development 120: 2823–2834.

Seydoux G, Mello CC, Pettitt J, et al. (1996) Repression of gene expression in the embryonic germ lineage of C. elegans. Nature 382: 713–716.

Spike CA, Coetzee D, Eichten C, et al. (2014a) The TRIM‐NHL protein LIN‐41 and the OMA RNA‐binding proteins antagonistically control the prophase‐to‐metaphase transition and growth of Caenorhabditis elegans oocytes. Genetics 198: 1535–1558.

Spike CA, Coetzee D, Nishi Y, et al. (2014b) Translational control of the oogenic program by components of OMA ribonucleoprotein particles in Caenorhabditis elegans. Genetics 198: 1513–1533.

Starich TA, Hall DH and Greenstein D (2014) Two classes of gap junction channels mediate soma‐germline interactions essential for germline proliferation and gametogenesis in Caenorhabditis elegans. Genetics 198: 1127–1153.

Strome S and Updike D (2015) Specifying and protecting germ cell fate. Nature Reviews Molecular Cell Biology 16: 406–416.

Updike D and Strome S (2010) P granule assembly and function in Caenorhabditis elegans germ cells. Journal of Andrology 31: 53–60.

Van Wynsberghe PM and Maine EM (2013) Epigenetic control of germline development. Advances in Experimental Medicine & Biology 757: 373–403.

Vella MC and Slack FJ (2005) C. elegans microRNAs. In: WormBook (ed.) The C. elegans Research Community. WormBook. DOI: 10.1895/wormbook.1.26.1,

Voronina E (2012) The diverse functions of germline P‐granules in Caenorhabditis elegans. Molecular Reproduction & Development 80: 624–631.

Voronina E, Paix A and Seydoux G (2012) The P granule component PGL‐1 promotes the localization and silencing activity of the PUF protein FBF‐2 in germline stem cells. Development 139: 3732–3740.

Wedeles CJ, Wu MZ and Claycomb JM (2013) Protection of germline gene expression by the C. elegans Argonaute CSR‐1. Developmental Cell 27: 664–671.

Yang H, Vallandingham J, Shiu P, et al. (2014) The DEAD box helicase RDE‐12 promotes amplification of RNAi in cytoplasmic foci in C. elegans. Current Biology 24: 832–838.

Youngman EM and Claycomb JM (2014) From early lessons to new frontiers: the worm as a treasure trove of small RNA biology. Frontiers in Genetics 5: 416.

Zanetti S and Puoti A (2013) Sex determination in the Caenorhabditis elegans germline. Advances in Experimental Medicine & Biology 757: 41–69.

Zarkower S (2006) Somatic sex determination. In: WormBook (ed.) The C. elegans Research Community. WormBook. DOI: 10.1895/wormbook.1.84.1,

Zhang B, Gallegos M, Puoti A, et al. (1997) A conserved RNA‐binding protein that regulates sexual fates in the C. elegans hermaphrodite germ line. Nature 390: 477–484.

Further Reading

Austin J and Kimble J (1987) glp‐1 is required in the germ line for regulation of the decision between mitosis and meiosis in C. elegans. Cell 51: 589–599.

Ferrell JE (1999) Xenopus oocyte maturation: new lessons from a good egg. Bioessays 21: 833–842.

Miller MA, Nguyen VQ, Lee MH, et al. (2001) A sperm cytoskeletal protein that signals oocyte meiotic maturation and ovulation. Science 291: 2144–2147.

Schedl T (2013) Germ cell development in C. elegans. Advances in Experimental Medicine and Biology 757: 1–425.

Contact Editor close
Submit a note to the editor about this article by filling in the form below.

* Required Field

How to Cite close
Voronina, Ekaterina, and Greenstein, David(Apr 2016) Germ Cell Fate Determination in C. elegans. In: eLS. John Wiley & Sons Ltd, Chichester. [doi: 10.1002/9780470015902.a0001501.pub2]