# Sex-type specification and meristem development in fern gametophytes

## TL;DR

This review explores how fern gametophytes determine sex and develop meristems, focusing on molecular and environmental factors in Ceratopteris richardii.

## Contribution

The paper highlights recent discoveries on antheridiogen responses and regulatory networks in fern sex-type specification.

## Key findings

- Environmental and molecular factors influence antheridiogen responses in fern gametophytes.
- Cellular mechanisms underpin sex-type specification and meristem development in Ceratopteris.
- Spatial expression dynamics and regulatory networks are key to sexual differentiation in ferns.

## Abstract

Sex determination, involving the physiological and morphological separation of male and female gamete-producing structures, promotes outcrossing and helps maintain genetic diversity in land plants. In homosporous ferns, which lack sex chromosomes, sex determination is specified during the gametophyte stage and is regulated by intergametophytic signaling mediated by the pheromone antheridiogen. Ceratopteris richardii has long served as a model for dissecting the molecular and cellular mechanisms underlying fern sex determination and associated meristem development in fern gametophytes, and in recent years, important new progress has been achieved. In this review, we summarize recent findings that uncover key environmental and molecular factors influencing antheridiogen responses and highlight the cellular basis of sex-type specification and the initiation and maintenance of meristems in Ceratopteris gametophytes. We also discuss emerging directions in the field, emphasizing the importance of spatial expression dynamics and interacting regulatory networks in fern sexual differentiation. Future advances will likely be driven by integrating genetic tools, multi-omics platforms, and quantitative live-imaging approaches, offering a more comprehensive view of the regulatory logic underlying sex determination and meristem development in ferns.