The C. elegans genome encodes a single Hand bHLH transcription factor. Either hnd-1(RNAi) or a hnd-1 deletion causes partially penetrant defects in viability and gonadogenesis. Dead embryos and young larvae are often misshapen at the posterior end. Our primary focus has been the role of hnd-1 in gonadogenesis. Wild-type C. elegans has two somatic gonadal precursors and two primordial germ cells in stereotyped positions within its four-celled gonadal primordium. The hnd-1 gene affects the presence and position of both the somatic gonadal precursors and primordial germ cells within the primordium, but does not appear to have any role in later gonadogenesis. hnd-1 probably acts within the somatic gonadal precursors or their mesodermal predecessors; defects in primordial germ cells and germ line appear to be secondary. In hnd-1 mutants, somatic gonadal precursors are generated normally, but are not maintained properly and sometimes die. A similar role in controlling the maintenance of precursor fates has been described for other genes governing early organogenesis, including the zebrafish Hand gene hands off. We also report the discovery of two genes, ehn-1 and ehn-3, that have overlapping functions with hnd-1 in embryogenesis and gonadogenesis.

The Caenorhabditis elegans somatic gonadal precursors (SGPs) are multipotent progenitors that give rise to all somatic tissues of the adult reproductive system. The hunchback and Ikaros-like gene ehn-3 is expressed specifically in SGPs and is required for their development into differentiated tissues of the somatic gonad. To find novel genes involved in SGP development, we used a weak allele of ehn-3 as the basis for a reverse genetic screen. Feeding RNAi was used to screen ∼2400 clones consisting of transcription factors, signaling components, and chromatin factors. The screen identified five members of the C. elegans SWI/SNF chromatin remodeling complex as genetic enhancers of ehn-3. We characterized alleles of 10 SWI/SNF genes and found that SWI/SNF subunits are required for viability and gonadogenesis. Two conserved SWI/SNF complexes, PBAF and BAF, are defined by their unique array of accessory subunits around a common enzymatic core that includes a catalytic Swi2/Snf2–type ATPase. Tissue-specific RNAi experiments suggest that C. elegans PBAF and BAF complexes control different processes during somatic gonadal development: PBRM-1, a signature subunit of PBAF, is important for normal SGP development, whereas LET-526, the distinguishing subunit of BAF, is required for development of a differentiated cell type, the distal tip cell (DTC). We found that the SWSN-4 ATPase subunit is required for SGP and DTC development. Finally, we provide evidence that C. elegans PBAF subunits and hnd-1/dHand are important for the cell fate decision between SGPs and their differentiated sisters, the head mesodermal cells.

During animal development, a complex of Par3, Par6 and atypical protein kinase C (aPKC) plays a central role in cell polarisation. The small G protein Cdc42 also functions in cell polarity and has been shown in some cases to act by regulating the Par3 complex. However, it is not yet known whether Cdc42 and the Par3 complex widely function together in development or whether they have independent functions. For example, many studies have implicated Cdc42 in cell migrations, but the Par3 complex has only been little studied, with conflicting results. Here we examine the requirements for CDC-42 and the PAR-3/PAR-6/PKC-3 complex in a range of different developmental events. We found similar requirements in all tissues examined, including polarised growth of vulval precursors and seam cells, migrations of neuroblasts and axons, and the development of the somatic gonad. We also propose a novel role for primordial germ cells in mediating coalescence of the Caenorhabditis elegans gonad. These results indicate that CDC-42 and the PAR-3/PAR-6/aPKC complex function together in diverse cell types.