Nitrogen-Driven Orchestration of Lateral Root Development: Molecular Mechanisms and Systemic Integration

Abstract

N, as plants' most essential nutrient, profoundly shapes root system architecture (RSA), with LRs being preferentially regulated. This review synthesizes the intricate molecular mechanisms underpinning N sensing, signaling, and its integration into developmental pathways governing LR initiation, primordium formation, emergence, and elongation. We delve deeply into the roles of specific transporters (NRT1.1, nitrate transporter 2.1 (NRT2.1)), transcription factors (Arabidopsis nitrate regulated 1 (ANR1), NLP7, TGACG motif-binding factor (TGA), squamosa promoter-binding protein-like 9 (SPL9)) and intricate hormone signaling networks (auxin, abscisic acid, cytokinins, ethylene) modulated by varying N availability (deficiency, sufficiency, excess) and chemical forms (NO₃^-, NH₄⁺, organic N). Emphasis is placed on the systemic signaling pathways, including peptide-mediated long-distance communication (CEP-C-terminally encoded peptide receptor 1 (CEPR1)) and the critical role of the shoot in modulating root responses. Furthermore, we explore the emerging significance of carbon-nitrogen (C/N) balance, post-translational modifications (ubiquitination, phosphorylation), epigenetic regulation, and the complex interplay with other nutrients (phosphorus (P), sulfur (S)) and environmental factors in shaping N-dependent LR plasticity. Recent advances utilizing single-cell transcriptomics and advanced imaging reveal unprecedented cellular heterogeneity in LR responses to N. Understanding this sophisticated regulatory network is paramount for developing strategies to enhance nitrogen use efficiency (NUE) in crops. This synthesis underscores how N acts as a master regulator, dynamically rewiring developmental programs through molecular hubs that synchronize nutrient sensing with root morphogenesis-a key adaptive strategy for resource acquisition in heterogeneous soils.

Keywords: auxin; hormone crosstalk; lateral root development; nitrate transporter; nitrogen signaling; nutrient sensing; root architecture plasticity; systemic signaling; transcription factors.