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Most actin-related proteins (Arps) are highly conserved and carry out well-defined cellular functions in eukaryotes. However, many lineages like  Drosophila  and mammals encode divergent non-canonical Arps whose roles remain unknown. To elucidate the function of non-canonical Arps, we focus on  Arp53D , which is highly expressed in testes and retained throughout  Drosophila  evolution. We show that Arp53D localizes to fusomes and actin cones, two germline-specific actin structures critical for sperm maturation, via a unique N-terminal tail. Surprisingly, we find that male fertility is not impaired upon  Arp53D  loss, yet population cage experiments reveal that  Arp53D  is required for optimal fitness in  Drosophila melanogaster . To reconcile these findings, we focus on  Arp53D  function in ovaries and embryos where it is only weakly expressed. We find that under heat stress  Arp53D -knockout (KO) females lay embryos with reduced nuclear integrity and lower viability; these defects are further exacerbated in  Arp53D -KO embryos. Thus, despite its relatively recent evolution and primarily testis-specific expression, non-canonical  Arp53D  is required for optimal embryonic development in  Drosophila .

An actin-related protein that is most highly expressed in Drosophila testes is critical for embryonic development