Systemic Optimization of Legume Nodulation: A Shoot-Derived Regulator, miR2111

Long-distance signaling between the shoot and roots of land plants plays a crucial role in ensuring their growth and development in a fluctuating environment, such as with soil nutrient deficiencies. MicroRNAs (miRNAs) are considered to contribute to such environmental adaptation via long-distance signaling since several miRNAs are transported between the shoot and roots in response to various soil nutrient changes. Leguminous plants adopt a shoot-mediated long-distance signaling system to maintain their mutualism with symbiotic nitrogen-fixing rhizobia by optimizing the number of symbiotic organs and root nodules. Recently, the involvement and importance of shoot-derived miR2111 in regulating nodule numbers have become evident. Shoot-derived miR2111 can systemically enhance rhizobial infection, and its accumulation is quickly suppressed in response to rhizobial inoculation and high-concentration nitrate application. In this mini-review, we briefly summarize the recent progress on the systemic optimization of nodulation in response to external environments, with a focus on systemic regulation via miR2111.

Rhizobia use a pathogenic-like effector to hijack leguminous nodulation signalling

Legume plants form a root-nodule symbiosis with rhizobia. This symbiosis establishment generally relies on rhizobium-produced Nod factors (NFs) and their perception by leguminous receptors (NFRs) that trigger nodulation. However, certain rhizobia hijack leguminous nodulation signalling via their type III secretion system, which functions in pathogenic bacteria to deliver effector proteins into host cells. Here, we report that rhizobia use pathogenic-like effectors to hijack legume nodulation signalling. The rhizobial effector Bel2-5 resembles the XopD effector of the plant pathogen Xanthomonas campestris and could induce nitrogen-fixing nodules on soybean nfr mutant. The soybean root transcriptome revealed that Bel2-5 induces expression of cytokinin-related genes, which are important for nodule organogenesis and represses ethylene- and defense-related genes that are deleterious to nodulation. Remarkably, Bel2-5 introduction into a strain unable to nodulate soybean mutant affected in NF perception conferred nodulation ability. Our findings show that rhizobia employ and have customized pathogenic effectors to promote leguminous nodulation signalling.

Soil acidity and nutrient deficiency cause poor legume nodulation in the permanent pasture and mixed farming zones of south-eastern Australia

Pasture legumes must be adequately and effectively nodulated in order to reach nitrogen-fixation targets. Of 225 pasture paddocks sampled across the Central Tablelands, Central West, Monaro and Riverina regions of New South Wales, 93% had inadequate legume nodulation. Legume content was significantly higher in the mixed faming zone (>50%, Central West and Riverina) than the permanent pasture zone (26%, Central Tablelands; 28% Monaro). Available phosphorus (P) was below critical levels in 40% of paddocks sampled and sulfur (S) in 73% of paddocks; >35% of all paddocks had soil pHCa <5.0. Deficiency of P was more prevalent in the Central Tablelands (63% of paddocks), whereas S deficiency occurred more frequently in the Central West (95% of paddocks). Legume nodule scores were associated with host legume species, soil pH, available P and/or S, and cation exchange capacity, which collectively accounted for 73% of variation. For Trifolium spp., at soil pHCa >5.55, nodulation was predicted to be near adequate (score 3.95, where adequate = 4). At pHCa <5.55, higher available S resulted in a higher nodulation score (2.42) than in paddocks where S was deficient (score 0–1.97). These results suggest that improving the capacity of legumes to supply nitrogen should focus on addressing soil acidity and plant nutrition, specifically P and S.