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Contrary to dogma, evolutionarily young and dynamic genes can encode essential functions. We find that evolutionarily dynamic  ZAD-ZNF  genes, which encode the most abundant class of insect transcription factors, are more likely to encode essential functions in  Drosophila melanogaster  than ancient, conserved  ZAD-ZNF  genes. We focus on the  Nicknack ZAD-ZNF  gene, which is evolutionarily young, poorly retained in  Drosophila  species, and evolves under strong positive selection. Yet we find that it is necessary for larval development in  D. melanogaster.  We show that  Nicknack  encodes a heterochromatin-localizing protein like its paralog  Oddjob , also an evolutionarily dynamic yet essential  ZAD-ZNF  gene. We find that the divergent  D. simulans  Nicknack protein can still localize to  D. melanogaster  heterochromatin and rescue viability of female but not male  Nicknack- null  D. melanogaster . Our findings suggest that innovation for rapidly changing heterochromatin functions might generally explain the essentiality of many evolutionarily dynamic  ZAD-ZNF  genes in insects.

Innovation of heterochromatin functions drives rapid evolution of essential ZAD-ZNF genes in Drosophila