Marine nitrogen-fixing cyanobacteria play a central role in the open-ocean microbial community by providing fixed nitrogen (N) to the ocean from atmospheric dinitrogen (N 2 ) gas. Once thought to be dominated by one genus of cyanobacteria, Trichodesmium, it is now clear that marine N 2 -fixing cyanobacteria in the open ocean are more diverse, include several previously unknown symbionts, and are geographically more widespread than expected. The next challenge is to understand the ecological implications of this genetic and phenotypic diversity for global oceanic N cycling. One intriguing aspect of the cyanobacterial N 2 fixers ecology is the range of cellular interactions they engage in, either with cells of their own species or with photosynthetic protists. From organelle-like integration with the host cell to a free-living existence, N 2 -fixing cyanobacteria represent the range of types of interactions that occur among microbes in the open ocean. Here, we review what is known about the cellular interactions carried out by marine N 2 -fixing cyanobacteria and where future work can help. Discoveries related to the functional roles of these specialized cells in food webs and the microbial community will improve how we interpret their distribution and abundance patterns and contributions to global N and carbon (C) cycles.Key index words: Cyanobacteria; ecology; nitrogen fixation; nitrogenase; symbioses List of Abbreviations: C, carbon; EPS, extracellular polysaccharides; N 2 , dinitrogen; N, nitrogen; PSII, photosystem II Cyanobacteria are responsible for a significant fraction of primary productivity on Earth, and undoubtedly played an important role in the oxygenation of the atmosphere since they evolved possibly up to 3.5 bya. Of particular relevance to this review, cyanobacteria were the photosynthetic predecessor of the chloroplast. Diverse cyanobacteria have the nitrogenase (nif) genes whose phylogenetic relationships suggest that N 2 fixation was either present at the time of divergence of the cyanobacteria, or inherited by lateral gene transfer early during their evolution. It should be noted that there are no N 2 -fixing eukaryotes and there is not a N 2 -fixing chloroplast. However, many terrestrial N 2 -fixing cyanobacteria form mutualistic associations with multicellular plants and fungi and intertidal N 2 -fixing cyanobacteria are associated with seagrasses (Hamisi et al. 2009) and microbial mats (Omoregie et al. 2004). In the open ocean, N 2 fixation is catalyzed by a few major lineages of cyanobacteria, including unicellular, filamentous nonheterocyst-forming, and filamentous heterocyst-forming cyanobacteria. These cyanobacteria showcase a continuum of interactions with other microbes (especially unicellular protists) and cells of the same species that range from freeliving, to loose attachment, to obligate symbiosis.Among the lineages of Bacteria that can fix N 2 in the open ocean, the N 2 -fixing cyanobacteria face a unique challenge in carrying out the disparate processes of N 2 fixation and pho...