Review: Emerging roles of brassinosteroid in nutrient foraging

Brassinosteroids (BRs) are well-characterized growth hormones that are critical for plant growth, development, and productivity. Genetic and molecular studies have revealed the key components of BR biosynthesis and signaling pathways. The membrane-localized BR signaling receptor, BRASSINOSTEROID INSENSITIVE1 (BRI1) binds directly to its ligand and initiates series of signaling events that led to the activation of BR transcriptional regulators, BRASSINAZOLE RESISTANT1 (BZR1) and BRI1-ETHYL METHANESULFONATE-SUPPRESSOR1 (BES1/BZR2) to regulate the cellular processes. Insights from Arabidopsis research revealed tissue and cell typespecific roles of BR in controlling cell elongation and maintenance of stem cell niche in roots. More recently, BRs have gained much attention in regulating the root growth during nutrient deficiency such as nitrogen, phosphorus, and boron. Differential distribution of nutrients in the rhizosphere alters BR hormone levels and signaling to reprogram spatial distribution of root system architecture (RSA) such as a change in primary root growth, lateral root numbers, length, and angle, root hair formation and elongation. These morpho-physiological changes in RSA are also known as an adaptive root trait or foraging response of the plant. In this review, we highlight the role of BRs in regulating RSA to increase root foraging response during fluctuating nutrient availability.