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Deoxycytidine monophosphate deaminase (DCD) is a key enzyme in the  de novo  dTTP biosynthesis pathway. Previous studies have indicated that DCD plays key roles in the maintenance of the balance of dNTP pools, cell cycle progression, and plant development. However, few studies have elucidated the functions of the DCD gene in Panicoideae plants.  Setaria  has been proposed as an ideal model of Panicoideae grasses, especially for C 4  photosynthesis research. Here, a  Setaria italica  stripe leaf mutant ( sistl2 ) was isolated from EMS-induced lines of “Yugu1,” the wild-type parent. The  sistl2  mutant exhibited semi-dwarf, striped leaves, abnormal chloroplast ultrastructure, and delayed cell cycle progression compared with Yugu1. High-throughput sequencing and map-based cloning identified the causal gene  SiSTL2 , which encodes a DCD protein. The occurrence of a single-base G to A substitution in the fifth intron introduced alternative splicing, which led to the early termination of translation. Further physiological and transcriptomic investigation indicated that  SiSTL2  plays an essential role in the regulation of chloroplast biogenesis, cell cycle, and DNA replication, which suggested that the gene has conserved functions in both foxtail millet and rice. Remarkably, in contrast to DCD mutants in C 3  rice,  sistl2  showed a significant reduction in leaf cell size and affected C 4  photosynthetic capacity in foxtail millet. qPCR showed that  SiSTL2  had a similar expression pattern to typical C 4  genes in response to a low CO 2  environment. Moreover, the loss of function of  SiSTL2  resulted in a reduction of leaf  13 C content and the enrichment of DEGs in photosynthetic carbon fixation. Our research provides in-depth knowledge of the role of DCD in the C 4  photosynthesis model  S. italica  and proposed new directions for further study of the function of DCD.

frontiers in plant science

SiSTL2 Is Required for Cell Cycle, Leaf Organ Development, Chloroplast Biogenesis, and Has Effects on C4 Photosynthesis in Setaria italica (L.) P. Beauv.