Efficient sucrose transport and metabolism are vital for seed and pollen development in plants. Cell wall invertases (CINs) hydrolyze sucrose into glucose and fructose, maintaining a sucrose gradient in the apoplast of sink tissues. In rice, two CIN isoforms, OsCIN1 and OsCIN2, were identified as being specifically expressed in the anthers but not in pollen. Functional analyses through genetic crosses and mutant characterization showed that oscin1/2 double mutants exhibit a sporophytic male-sterile phenotype and produce shrunken seeds. This suggests that CIN activity is essential for proper pollen development and seed formation in rice. Observation of the progeny genotypes and phenotypes from various genetic crosses revealed that the phenotype of oscin1/2 seeds is determined by the genotype of the maternal tissue, indicating the critical role of CIN function in the apoplast between maternal and filial tissues for sucrose transport and metabolism. The CIN activity in the anthers and seeds of wild-type rice was found to be significantly higher—over 500-fold in the anthers and 5-fold in the seeds—than in the leaves, highlighting the importance of CIN in facilitating the efficient unloading of sucrose. These findings suggest that the fine-tuning of CIN activity in the apoplast, achieved through tissue-specific expression and CIN isoform regulation, plays a key role in determining the carbohydrate distribution across different tissues. Understanding this regulatory mechanism could provide opportunities to manipulate carbohydrate allocation to sink organs, potentially enhancing crop yields.

Cell wall invertases (CINs) establish sucrose gradients between source and sink tissues, essential for the allocation of photoassimilates. Rice possesses nine CIN genes, among which OsCIN1 and OsCIN2 have been reported as key regulators of sink strength. To test whether increasing CIN activity enhances grain yield, we generated OsCIN1 overexpression lines in rice driven by the CaMV 35S promoter. Subcellular localization analysis of OsCIN1–GFP confirmed its apoplastic localization. OsCIN1 promoter::GUS analyses verified expression in vascular tissues and revealed predominant signals in the ovular vascular and lateral stylar vascular traces during seed development. Although CIN activity was markedly elevated throughout the plant, the resulting phenotypes were unexpected. Sugar profiling of flag leaves at the flowering stage showed almost complete sucrose depletion in the overexpression (OX) lines, accompanied by increased hexose and starch accumulation. Under field conditions, OsCIN1 OX plants exhibited ~50% fewer tillers and a lower 1000-grain weight relative to wild type (WT), resulting in reduced productivity. Ectopic expression of OsCIN1 disrupted the sucrose concentration gradient, weakened carbon partitioning to sink tissues, and impaired key agronomic traits. Collectively, sugar flux is governed by the spatiotemporal patterning of CINs, highlighting that precise spatial and temporal control of CIN activity is required to increase yield.