Rice Ferredoxin-Dependent Glutamate Synthase Regulates Nitrogen-Carbon Metabolomes and Is Genetically Differentiated between japonica and indica Subspecies

Plants assimilate inorganic nitrogen absorbed from soil into organic forms as Gin and Glu through the glutamine synthetase/glutamine：2-oxoglutarate amidotransferase （GS/GOGAT） cycle. Whereas GS cata- lyzes the formation of Gin from Glu and ammonia, GOGAT catalyzes the transfer of an amide group from Gin to 2-oxoglutarate to produce two molecules of Glu. However, the regulatory role of the GS/GOGAT cycle in the carbon-nitrogen balance is not well understood. Here, we report the functional characterization of rice ABNORMAL CYTOKININ RESPONSE 1 （ABC1） gene that encodes a ferredoxin-dependent （Fd）- GOGAT. The weak mutant allele abcl-1 mutant shows a typical nitrogen-deficient syndrome, whereas the T-DNA insertional mutant abcl-2 is seedling lethal. Metabolomics analysis revealed the accumulation of an excessive amount of amino acids with high N/C ratio （Gin and Asn） and several intermediates in the tricarboxylic acid cycle in abcl-1, suggesting that ABC1 plays a critical role in nitrogen assimilation and carbon-nitrogen balance. Five non-synonymous single-nucleotide polymorphisms were identified in the ABC1 coding region and characterized as three distinct haplotypes, which have been highly and specifically differentiated between japonica and indica subspecies. Collectively, these results suggest that ABC1/ OsFd-GOGAT is essential for plant growth and development by modulating nitrogen assimilation and the carbon-nitrogen balance.