fos genes in mainly invertebrate model systems: A review of commonalities and some diversities

fos genes, transcription factors with a common basic region and leucine zipper domains binding to a consensus DNA sequence (TGA{}TCA), are evolutionarily conserved in eukaryotes. Homologs can be found in many different species from yeast to vertebrates. In yeast, the homologous GCN4 gene is required to mediate "emergency" situations like nutrient deprivation and the unfolded protein response. The C. elegans homolog fos-1 is required for reproduction and vulval development, as well as in adult homeostasis. In Drosophila melanogaster, there is also a sole fos homolog: the gene kayak, with five isoforms. The kayak locus has been studied in detail. It was originally described as embryonic lethal with a "dorsal open" phenotype. Since then, kayak has been shown to be required for oocyte maturation and as a source for piRNA; for early dorsoventral specification, macrophage function, dorsal closure, endoderm differentiation, and finally during metamorphosis in wing and eye-antennal development. In mammals there are multiple fos loci, each one with alternative splicing giving rise to multiple isoforms. Overall, mammalian fos genes are required for bone, cartilage and tooth formation, and in some instances for placental angiogenesis and retinal function. We review here mainly what is known about fos function in invertebrate model systems, especially during embryogenesis. We propose that fos genes, evolutionarily conserved transcription factors, evolved early during eukaryotic development, and from its inception as part of an environmental stress response machinery, were co-opted several times during development to regulate processes that may require similar cellular responses.