Relationship Between the Vascular Bundle Structure of Panicle Branches and the Filling of Inferior Spikelets in Large-Panicle Japonica Rice

The vascular bundles of rice panicles serve to connect the source and the sink, as well as serving as a channel for the transportation of materials. In this study, two homozygous japonica rice strains were used as materials. The vascular bundle structures of the branches in different positions within a rice panicle were observed, and their cross-sectional areas were calculated. In addition, the ultrastructure of the central large vascular bundle (LVB) phloem in the rachillae of superior spikelets (SS) and inferior spikelets (IS) was observed during the grain filling period. Moreover, the soluble sugar and protein contents of the SS and IS rachillae were also measured to study whether the differences in the structure of vascular bundles of the branches were related to the plumpness of grain at different positions. The results showed that vascular bundle cross-sectional areas of the basal primary branches were greater than those in the upper primary branches. Moreover, there was little difference in the areas of vascular bundles between the basal secondary branches and upper secondary branches. However, the vascular bundle areas of the IS rachillae were lower than those in the SS rachillae. Therefore, we believe that the poor vascular tissue channel of the IS rachillae could be the limiting factor in IS plumpness. The results also showed that a similar time course in the degradation pattern of some organelles of the sieve elements and companion cells in central LVB was observed in the SS rachillae and IS rachillae during the grain filling period. Compared with the IS rachillae, more abundant mitochondria and plasmodesmata were found in the companion cells of SS rachillae at the beginning of the filling stage, while no significant differences between SS and IS rachillae were identified at the middle and late filling stages, which implies that the SS rachillae were relatively more effective at transportation compared with the IS rachillae at the initial filling stage. Therefore, the undeveloped vascular bundles of the IS rachillae and their poor physiology and lack of ability to transport at the initial filling stages could be the limiting factor in IS plumpness.

GC-MS-based metabolite profiling of key differential metabolites between superior and inferior spikelets of rice during the grain filling stage

Background The asynchronous filling between superior spikelets (SS) and inferior spikelets (IS) in rice has become a research hotspot. The stagnant development and poor grain filling of IS limit yields and the formation of good quality rice. A large number of studies on this phenomenon have been carried out from the genome, transcriptome and proteome level, indicating that asynchronous filling of SS and IS filling is a complex, but orderly physiological and biochemical process involving changes of a large number of genes, protein expression and modification. However, the analysis of metabolomics differences between SS and IS is rarely reported currently. Results This study utilized untargeted metabolomics and identified 162 metabolites in rice spikelets. Among them, 17 differential metabolites associated with unsynchronized grain filling between SS and IS, 27 metabolites were related to the stagnant development of IS and 35 metabolites related to the lower maximum grain-filling rate of IS compared with the SS. We found that soluble sugars were an important metabolite during grain filling for SS and IS. Absolute quantification was used to further analyze the dynamic changes of 4 types of soluble sugars (sucrose, fructose, glucose, and trehalose) between SS and IS. The results showed that sucrose and trehalose were closely associated with the dynamic characteristics of grain filling between SS and IS. The application of exogenous sugar showed that trehalose functioned as a key sugar signal during grain filling of IS. Trehalose regulated the expression of genes related to sucrose conversion and starch synthesis, thereby promoting the conversion of sucrose to starch. The difference in the spatiotemporal expression of TPS-2 and TPP-1 between SS and IS was an important reason that led to the asynchronous change in the trehalose content between SS and IS. Conclusions The results from this study are helpful for understanding the difference in grain filling between SS and IS at the metabolite level. In addition, the present results can also provide a theoretical basis for the next step of using metabolites to regulate the filling of IS.

Efficiency of Sucrose to Starch Metabolism Is Related to the Initiation of Inferior Grain Filling in Large Panicle Rice

The poor grain-filling initiation often causes the poor development of inferior spikelets (IS) which limits the yield potential of large panicle rice (Oryza sativa L.). However, it remains unclear why IS often has poor grain-filling initiation. In addressing this problem, this study conducted a field experiment involving two large panicle rice varieties, namely CJ03 and W1844, in way of removing the superior spikelets (SS) during flowering to force enough photosynthate transport to the IS. The results of this study showed that the grain-filling initiation of SS was much earlier than the IS in CJ03 and W1844, whereas the grain-filling initiation of IS in W1844 was evidently more promoted compared with the IS of CJ03 by removing spikelets. The poor sucrose-unloading ability, i.e., carbohydrates contents, the expression patterns of OsSUTs, and activity of CWI, were highly improved in IS of CJ03 and W1844 by removing spikelets. However, there was a significantly higher rise in the efficiency of sucrose to starch metabolism, i.e., the expression patterns of OsSUS4 and OsAGPL1 and activities of SuSase and AGPase, for IS of W1844 than that of CJ03. Removing spikelets also led to the changes in sugar signaling of T6P and SnRK1 level. These changes might be related to the regulation of sucrose to starch metabolism. The findings of this study suggested that poor sucrose-unloading ability delays the grain-filling initiation of IS. Nonetheless, the efficiency of sucrose to starch metabolism is also strongly linked with the grain-filling initiation of IS.

GC-MS-Based Metabolite Profiling of Key Differential Metabolites Between Superior and Inferior Spikelets of Rice During the Grain Filling Stage

Background:The asynchronous filling between superior spikelets (SS) and inferior spikelets (IS) in rice has become a research hotspot. The stagnant development and poor grain filling of IS limit yields and the formation of good quality rice. A large number of studies on this phenomenon have been carried out from the genome, transcriptome and proteome level, indicating that asynchronous filling of SS and IS filling is a complex, but orderly physiological and biochemical process involving changes of a large number of genes, protein expression and modification. However, the analysis of metabolomics differences between SS and IS is rarely reported currently.Results:This study utilized untargeted metabolomics and identified 162 metabolites in rice spikelets. Among them, 17 differential metabolites associated with unsynchronized grain filling between superior spikelets (SS) and IS, 27 metabolites were related to the stagnant development of IS and 35 metabolites related to the lower maximum grain-filling rate of IS compared with the SS. We found that soluble sugars were an important metabolite during grain filling for SS and IS. Absolute quantification was used to further analyze the dynamic changes of 4 types of soluble sugars (sucrose, fructose, glucose, and trehalose) between SS and IS. The results showed that sucrose and trehalose were closely associated with the dynamic characteristics of grain filling between SS and IS. The application of exogenous sugar showed that trehalose functioned as a key sugar signal during grain filling. Trehalose regulated the expression of genes related to sucrose conversion and starch synthesis, thereby promoting the conversion of sucrose to starch. The difference in the spatiotemporal expression of TPS-2 and TPP-1 between SS and IS was an important reason that led to the difference in trehalose contents between SS and IS.Conclusions:The results from this study are helpful for understanding the difference in grain filling between SS and IS at the metabolite level. In addition, the present results can also provide a theoretical basis for the next step of using metabolites to regulate the filling of IS.

Seed filling determines seed vigour of superior and inferior spikelets during hybrid rice (Oryza sativa) seed production

Growth and development of spikelets are asynchronous during hybrid rice seed production. However, knowledge is limited about the variation in seed vigour between superior and inferior spikelets. The main objective of this study was to evaluate the differences in seed vigour between superior and inferior spikelets of hybrid rice, and to explore the mechanisms associated with such variations with regards to seed filling characteristics and starch accumulation. Field experiments were conducted in 2017 and 2018, and seed vigour parameters, seed filling characteristics and starch accumulation were determined. We found that significant differences were recorded in seed vigour parameters between superior and inferior spikelets. Germination percentage (GP) and vigour index (VI) of superior spikelets were higher than that of inferior spikelets, irrespective of year or variety. Moreover, significant differences were observed in seed filling characteristics and starch accumulation between superior and inferior spikelets. Lower total starch content, amylose content and amylopectin content were recorded in inferior spikelets with low seed filling rate and long seed filling duration. Superior spikelets with high seed filling rate and short seed filling duration increased final seed weight by 23.9 and 22.3% for each variety, respectively. Therefore, these results raise the possibility that seed filling influences the starch accumulation and seed weight, and is closely associated with differences in the vigour of seeds from superior and inferior spikelets during hybrid rice seed production.