Cold adaptation in drylands: transcriptomic insights into cold-stressedNostoc flagelliformeand characterization of a hypothetical gene with cold and nitrogen stress tolerance

Environmental stressors, especially low temperature, are very common on the earth's dryland systems. Terrestrial cyanobacteria have evolved with cold adaptability in addition to extreme dryness and high irradiation resistance. The dryland soil surface-dwelling species,Nostoc flagelliforme, serves as a potential model organism to gain insights into cyanobacterial cold adaptation. In this study, we performed transcriptomic analysis ofN.flagelliformesamples in response to low temperature. The results revealed that the biological processes, such as terpenoid biosynthetic process, oxidoreductase activity, carbohydrate metabolism, biosynthesis of secondary metabolites, lipid and nitrogen metabolism, were significantly and dynamically changed during the cold stress. It was noteworthy that the transcription of the denitrification pathway for ammonia accumulation was enhanced, implying an importance for nitrogen utilization in stress resistance. In addition, characterization of a cold-responsive hypothetical genecsrnf1found that it could greatly improve the cold-resistant performance of cells when it was heterologously expressed in transgenicNostocsp. PCC 7120. It was also found thatcsrnf1transgenic strain exhibited resistance to nitrogen-deficient environmental stress. Considering that dryland cyanobacteria have to cope with low temperature on infertile soils, this study would enrich our understanding on the importance of multifunction of the genes for environmental cold adaptation in drylands.