Functions and Regulation of HAM Family Genes in Meristems During Gametophyte and Sporophyte Generations.

A fascinating feature of land plants is their ability to continually initiate new tissues and organs throughout their lifespan, driven by a pool of pluripotent stem cells located in meristems. In seed plants, various types of meristems are initiated and maintained during the sporophyte generation, while their gametophytes lack meristems and rely on sporophyte tissues for growth. In contrast, seed-free vascular plants, such as ferns, develop meristems during both the sporophyte and gametophyte generations, allowing for the independent growth of both generations. Recent findings have highlighted both conserved and lineage-specific roles of the HAIRY MERISTEM (HAM) family of GRAS-domain transcriptional regulators in various meristems throughout the land plant lifecycle. Here, we review and discuss how HAM genes maintain meristem indeterminacy in both sporophytes and gametophytes, with a focus on studies performed in two model species: the flowering plant Arabidopsis thaliana and the fern Ceratopteris richardii. Additionally, we summarize the crucial and tightly regulated functions of the microRNA171 (miR171)-HAM regulatory modules, which define HAM spatial patterns and activities during meristem development across various meristem identities in land plants.