Diversity of nitrogen fixation strategies in Mediterranean legumes.

Symbiotic N2 fixation (SNF) brings nitrogen into ecosystems, fuelling much of the world's agriculture(1) and sustaining carbon storage(2,3). However, it can also cause nitrogen saturation, exacerbating eutrophication and greenhouse warming(4-7). The balance of these effects depends on the degree to which N2-fixing plants adjust how much N2 they fix based on their needs (their SNF 'strategies')(5,6). Genetic, biochemical and physiological details of SNF are well known for certain economically important species(8,9), but the diversity of N2-fixing plants(10) and bacteria(11) is enormous, and little is known about most N2-fixing symbioses in natural ecosystems(12). Here, we show that co-occurring, closely related herbs exhibit diverse SNF strategies. In response to a nitrogen supply gradient, four species fixed less N2 than they needed (over-regulation), two fixed what they needed (facultative) and two did not downregulate SNF (obligate). No species downregulated but fixed more N2 than it needed (under-regulation or incomplete downregulation), but some species under-regulated or incompletely downregulated structural allocation to SNF. In fact, most species maintained nodules (the root structures that house symbionts) when they did not fix N2, suggesting decoupling of SNF activity and structure. Simulations showed that over-regulation of SNF activity is more adaptive than under-regulation or incomplete downregulation, and that different strategies have wildly different effects on ecosystem-level nitrogen cycling.