Cellular anatomy of arbuscular mycorrhizal fungi

Arbuscular mycorrhizal (AM) fungi are ancient plant mutualists that are ubiquitous across terrestrial ecosystems. These fungi are unique among most eukaryotes because they form multinucleate, open-pipe mycelial networks, where nutrients, organelles, and chemical signals move bidirectionally across a continuous cytoplasm. AM fungi play a crucial role in ecosystem functioning by supporting plant growth, mediating ecosystem diversity, and contributing to carbon cycling. It is estimated that plant communities allocate ∼3.93 Gt CO₂e to AM fungi every year, much of which is stored as lipids inside the fungal network. Despite their ecological significance, the cellular biology of AM fungi remains underexplored. Here, we synthesise the current knowledge on AM fungal cellular structure and organisation. We examine AM fungal development at different biological levels - the hypha and its content, hyphal networks and AM fungal spores - and explore key cellular dynamics. This includes cell wall composition, cytoplasmic contents, nuclear and lipid organisation and dynamics, network architecture, and connectivity. We highlight how their unique cellular arrangement enables complex cytoplasmic flow and nutrient exchange processes across their open-pipe mycelial networks. We discuss how both established and novel techniques, including microscopy, culturing, and high-throughput image analysis, are helping to resolve previously unknown aspects of AM fungal biology. By comparing these insights with established knowledge in other, well-studied filamentous fungi, we identify critical knowledge gaps and propose questions for future research to further our understanding of fundamental AM fungal cell biology and its contributions to ecosystem health.