Nitrogen (N) is a crucial macronutrient for plant growth, with nitrate as a primary inorganic N source for most plants. Beyond its role as a nutrient, nitrate also functions as a signalling molecule, influencing plant morphogenetic development. While nitrate utilization and signalling mechanisms have been extensively studied in model plants, the origin, evolution, and diversification of core components in nitrate uptake, assimilation, and signalling remain largely unexplored. In our investigation, we discovered that deep sea algae living in low nitrate conditions developed a high-affinity transport system (HATS) for nitrate uptake and a pathway of nitrate primary assimilation (NR-NiR-GS-GOGAT). In contrast, low-affinity transport systems (LATS) and the plastid GS originated from the ancestors of land and seed plants, respectively. These adaptations facilitated amino acid acquisition as plants conquered terrestrial environments. Furthermore, the intricate nitrate signalling, relying on NRT1.1 and NLP7, evolved stepwise, potentially establishing systematic regulation in bryophytes for self-regulation under complex terrestrial nitrate environments. As plants underwent terrestrialization, they underwent adaptive changes to thrive in dynamic nitrate environments, continually enhancing their nitrate uptake, assimilation, and signal transduction abilities.
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