Finding a nitrogen niche: a systems integration of local and systemic nitrogen signalling in plants.

The ability of plants to sense their nitrogen (N) microenvironment in the soil and deploy strategic root growth in N-rich patches requires exquisite systems integration. Remarkably, this new paradigm for systems biology research has intrigued plant biologists for more than a century, when a split-root framework was first used to study how plants sense and respond to heterogeneous soil nutrient environments. This systemic N-signalling mechanism, allowing plants to sense and forage for mineral nutrients in resource-rich patches, has important implications for agriculture. In this review, we will focus on how advances in the post-genomic era have uncovered the gene regulatory networks underlying systemic N-signalling. After defining how local and systemic N-signalling can be experimentally distinguished for molecular study using a split-root system, the genetic factors that have been shown to mediate local and/or systemic N-signalling are reviewed. Second, the genetic mechanism of this regulatory system is broadened to the whole genome level. To do this, publicly available N-related transcriptomic datasets are compared with genes that have previously been identified as local and systemic N responders in a split-root transcriptome dataset. Specifically, (i) it was found that transcriptional reprogramming triggered by homogeneous N-treatments is composed of both local and systemic responses, (ii) the spatio-temporal signature of local versus systemic responsive genes is defined, and (iii) the conservation of systemic N-signalling between Arabidopsis and Medicago is assessed. Finally, the potential mediators, i.e. metabolites and phytohormones, of the N-related long-distance signals, are discussed.