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Iron is an essential element for plants growth and development. Biofortification has become an efficient way for iron accumulation in grains to enhance the nutritional quality of crops. Both iron and sucrose are transported via phloem during grain filling in rice. However, the relationship between iron and sucrose transport remains unclear. In this study, we aimed to elucidate the relationship between sucrose transport and iron accumulation in rice. We first found that the efficient iron accumulated stage and sucrose unloading stage overlapped at 5–10 DAF during grain filling. The close relationship was further validated by transgenic lines (Y2 and Y6) overexpressing yeast (Y) invertase (INV). The sucrose efflux rate in phloem decreased by 40.1% and 30.6%, and the iron efflux rate in phloem was also reduced by 73.7% and 51.0% in Y2 and Y6, respectively. Thus, the iron content in grains was decreased. The transcriptome analysis on grains of WT and Y2 at 10 DAF was used to compare the difference in carbohydrate and iron metabolism. Seven upregulated DEGs and twenty-four downregulated DEGs related to carbohydrate metabolism were identified in Y2. The expression levels of starch synthesis and sugar transport genes in Y2 were lower than those in WT. The expression levels of iron-related genes, including OsTOM1, OsNAS1, OsNAS2, OsIRT1, OsYSL9, and OsOPT7, were significantly downregulated in Y2. These results indicated that blocking photoassimilates allocation between source and sink reduces iron accumulation in grains. [ABSTRACT FROM AUTHOR]

The adverse effect of future global warming on rice quality is a fact that cannot be ignored. Evidence from our previous field warming experiments confirmed that warming had a significant negative impact on rice appearance and processing quality during the grain-filling stage. Therefore, clarifying the regulation mechanism of cultivation measures on the formation of quality under elevated temperature and formulating countermeasures to ensure the production of high-quality rice are of great significance in the future. As a cultivation measure that is easy to operate in the field, nitrogen has been shown to be effective in mitigating the quality deterioration induced by field warming in our previous study. In this study, results of actual field warming and nitrogen treatments showed that nitrogen mainly dominated grain nitrogen metabolism under elevated temperature, further affecting the composition and proportion of grain storage proteins. Furthermore, nitrogen significantly up-regulated the expression of GluA-1, pro14 and pro17 in protein synthesis and PDIL1-1 in protein processing pathway under elevated temperature, which further alleviated the decrease of prolamin content and enhanced the glutelin content. The transcriptional regulation of storage protein synthesis genes by nitrogen further induced the changes of storage substances, which will help us to further analyze the formation mechanism of rice quality under elevated temperature and provide a basis for the establishment of rational nitrogen regulation pathways for high-quality rice production under global warming in the future. [ABSTRACT FROM AUTHOR]

The remobilization of non-structural carbohydrates (NSCs) in the stem is essential for rice grain filling so as to improve grain yield. We conducted a two-year field experiment to deeply investigate their relationship. Two large-panicle rice varieties with similar spikelet size, CJ03 and W1844, were used to conduct two treatments (removing-spikelet group and control group). Compared to CJ03, W1844 had higher 1000-grain weight, especially for the grain growth of inferior spikelets (IS) after removing the spikelet. These results were mainly ascribed to the stronger sink strength of W1844 than that of CJ03 contrasting in the same group. The remobilization efficiency of NSC in the stem decreased significantly after removing the spikelet for both CJ03 and W1844, and the level of sugar signaling in the T6P-SnRK1 pathway was also significantly changed. However, W1844 outperformed CJ03 in terms of the efficiency of carbon reserve remobilization under the same treatments. More precisely, there was a significant difference during the early grain-filling stage in terms of the conversion of sucrose and starch. Interestingly, the sugar signaling of the T6P and SnRK1 pathways also represented an obvious change. Hence, sugar signaling may be promoted by sink strength to remobilize the NSCs of the rice stem during grain filling to further advance crop yield. [ABSTRACT FROM AUTHOR]

Grain chalkiness is a key quality trait of the rice grain, whereas its underlying mechanism is still not thoroughly understood because of the complex genetic and environmental interactions. We identified a notched-belly (NB) mutant that has a notched-line on the belly of grains. The line dissects the endosperm into two distinct parts, the upper translucent part, and the bottom chalky part in the vicinity of the embryo. Using this mutant, our previous studies clued the negative influence of embryo on the biochemical makeup of the endosperm, suggesting the need for the in-depth study of the embryo effect on the metabolome of developing endosperm. This study continued to use the NB mutant to evolve a novel comparison method to clarify the role of embryo in the formation of a chalky endosperm. Grain samples of the wild-type (WT) and NB were harvested at 10, 20, and 30 days after fertilization (DAF), and then divided into subsamples of the embryo, the upper endosperm, and the bottom endosperm. Using non-targeted metabolomics and whole-genome RNA sequencing (RNA-seq), a nearly complete catalog of expressed metabolites and genes was generated. Results showed that the embryo impaired the storage of sucrose, amino acid, starch, and storage proteins in the bottom endosperm of NB by enhancing the expression of sugar, amino acids, and peptide transporters, and declining the expression of starch, prolamin, and glutelin synthesis-related genes. Importantly, the competitive advantage of the developing embryo in extracting the nutrients from the endosperm, transformed the bottom endosperm into an "exhaustive source" by diverting the carbon (C) and nitrogen (N) metabolism from synthetic storage to secondary pathways, resulting in impaired filling of the bottom endosperm and subsequently the formation of chalky tissue. In summary, this study reveals that embryo-induced metabolic shift in the endosperm is associated with the occurrence of grain chalkiness, which is of relevance to the development of high-quality rice by balancing the embryo–endosperm interaction. [ABSTRACT FROM AUTHOR]

With the intensification of global warming, rice production is facing new challenges. Field evidence indicates that elevated temperature during rice grain-filling leads to the further deterioration of grain quality. In order to clarify the potential regulatory mechanism of elevated temperature on the formation of rice quality, the DIA mass spectrometry method under the background of field warming was conducted to investigate the regulatory effects of high temperature on grain development and material accumulation pathways. The results showed that a total of 840 differentially expressed proteins were identified during the grain-filling process under elevated temperature. These differentially expressed proteins participated in carbon metabolism, amino acid biosynthesis, signal transduction, protein synthesis, and alternately affected the material accumulation of rice grains. The significant up-regulation of PPROL 14E, PSB28, granule-bound starch synthase I, and the significant down-regulation of 26.7 kDa heat shock protein would lead to the component difference in grain starch and storage proteins, and that could be responsible for the degradation of rice quality under elevated temperature. Results suggested that proteins specifically expressed under elevated temperature could be the key candidates for elucidating the potential regulatory mechanism of warming on rice development and quality formation. In-depth study on the metabolism of storage compounds would be contributed in further proposing high-quality cultivation control measures suitable for climate warming. [ABSTRACT FROM AUTHOR]

Our previous study has shown that nitrogen plays an important role in dealing with significantly increased chalkiness caused by elevated temperature. However, the role of nitrogen metabolites has not been given sufficient attention, and its regulatory mechanism is not clear. This study investigated the effects of high temperature and nitrogen fertilizer on the synthesis of grain storage protein and further explored the quality mechanism under the actual scenario of field warming. Results showed that increased temperature and nitrogen fertilizer could affect the activities of nitrogen metabolism enzymes, namely, glutamate synthetase, glutamine synthetase, glutamic pyruvic transaminase, and glutamic oxaloacetic transaminase, and the expressions of storage protein synthesis factor genes, namely, GluA and GluB , and subfamily genes, namely, pro14, BiP1 , and PDIL1 , which co-induced the changes of storage protein synthesis in rice grains. Furthermore, the increased temperature changed the balance of grain storage substances which may lead to the significantly increased chalky rate (197.67%) and chalkiness (532.92%). Moreover, there was a significant negative correlation between prolamin content and chalkiness, indicating that nitrogen fertilizer might regulate the formation of chalkiness by affecting the synthesis of prolamin. Results suggested that nitrogen application could regulate the related core factors involved in nitrogen metabolism pathways, which, in turn, affects the changes in the storage protein components in the grain and further affects quality. Therefore, as a conventional cultivation measure, nitrogen application would have a certain value in future rice production in response to climate warming. [ABSTRACT FROM AUTHOR]

Background: Large panicle rice has a large sink capacity, but inferior spikelet filling is poor in this variety of rice due to asynchronous grain filling. The understanding of the factors that cause asynchronous grain filling will help to propose a model for how to regulate the rice inferior spikelets grain filling. Results: In this study, two large panicle rice varieties, W1844 and CJ03, with the same sink capacity but with differences in asynchronous grain filling were used. The difference in the grain filling rate between superior and inferior spikelets in W1844 was much smaller than that in CJ03. We found that superior spikelet filling was initiated earlier in W1844 than in CJ03. The source-to-sink translocation rate of sucrose during the grain filling stage was more efficient in W1844 than in CJ03, and the gene expression levels of sucrose transporters (OsSUTs) were higher in W1844 functional leaves than in those of CJ03. In addition, carbon output, the transport ratio, and the contribution rate from the stem and sheath to the panicle were much higher at the early filling stage than at later filling stages in W1844. Conclusion: Efficient sugar translocation can satisfy high sink strength, and also the strong sink activity can facilitate the sugar unloading in spikelets. All the above results indicate that an efficient sugar translocation rate at the early grain filling stage can improve sink strength and inferior grain filling initiation. Strategies to limit asynchronous grain filling in rice were also discussed based on our findings. [ABSTRACT FROM AUTHOR]

Climate change threatens rice production by increasing the frequency of extreme temperatures during grain-filling stage (T GF). Nitrogen application can significantly affect rice yield and quality. However, the effects of interaction between T GF and nitrogen application rate on yield and quality of rice remains unclear. The objectives of this study were to comprehensively evaluate the quantitative regulatory mechanism between T GF and nitrogen application rate on rice yield and quality formation. A two-year field experiment with three varieties, five sowing dates, and three nitrogen application rates was conducted at four ecological sites in the middle and lower reaches of the Yangtze River, China in 2019 and 2020. Grain yield, milling, appearance, and eating quality were quantified for each treatment. This study demonstrated that T GF varied from 17.8 ℃ to 31.4 ℃ in different sites and sowing dates and decreased by an average of 2.1 ℃ for every 15 days the sowing date was delayed. The grain yield, head rice rate, chalky grain rate, chalkiness and taste value were all related to T GF through a quadratic function. The optimum T GF for both yield and quality of indica hybrid rice was 24.2–26.5 ℃. In terms of grain yield, nitrogen agronomic efficiency increased linearly with T GF , and nitrogen application could only confer a small increase in yield or even reduce yield under extreme low T GF. Nitrogen application alleviated the adverse effects of high T GF on rice quality, but exacerbated the deterioration of quality under low T GF. For milling and appearance quality and eating quality, the critical T GF leading to such opposite regulatory effect were identified as 23.4 ± 0.4 ℃ and 29.5 ± 0.5 ℃, respectively. The variations of the ratio of amylopectin to amylose and protein content altered texture and starch gelatinization properties, mainly explaining the differences in eating quality caused by T GF and nitrogen application rate. The present study suggests that T GF should be considered when formulating nitrogen management strategies for high-quality rice, which is crucial for alleviating the deterioration of rice quality caused by extreme T GF under global climate change. [Display omitted] • The optimum temperature during grain filling stage (T GF) for indica hybrid rice was 24.2–26.5 ℃. • Nitrogen agronomic efficiency was significantly positively correlated with T GF. • Nitrogen application could alleviate the adverse effects of high T GF on rice quality. • Nitrogen application exacerbated the deterioration of quality under low T GF. [ABSTRACT FROM AUTHOR]

Rice glutelin is sensitive to temperature and nitrogen, however, the regulatory mechanism of glutelin response to temperature and nitrogen is unclear. In this study, we conducted the open field warming experiment by the Free-air temperature enhancement facility and application of nitrogen during grain filling. In three-year field warming experiments, glutelin relative content was significantly increased under elevated temperature and application of nitrogen. Temperature and nitrogen and their interaction increased the glutelin accumulation rate in the early and middle grain filling stages (10-25d after flowering), but decreased the glutelin accumulation rate in the middle and late grain filling stages (25-45d after flowering). Elevated temperature promoted pro-glutelin levels whereas application of nitrogen under warming increased the amount of α-glutelin. At the transcriptional level, the expression levels of the glutelin-encoding genes and protein disulphide isomerase-like enzyme (PDIL1-1), glutelin precursor accumulation 4 (GPA4), glutelin precursor mutant 6 (GPA2), glutelin precursor accumulation 3 (GPA3) and vacuolar processing enzyme (OsVPE1) of glutelin folding, transport and accumulation-related genes were up-regulated by nitrogen under natural temperature as early as 5d after flowering. However, elevated temperature up-regulated glutelin-encoding genes before 20d after flowering, and the expression of endoplasmic reticulum chaperone (OsBip1), OsPDIL1-1 , small GTPase gene (GPA1), GPA2-GPA4 and OsVPE1 were significantly increased post 20d after flowering under warming. In addition, the increase in glutelin content worsened grain quality, particularly chalkiness and eating quality. Overall, the results were helpful to understand glutelin accumulation and provide a theoretical basis for further study the relationship between rice quality and glutelin under global warming. • Elevated temperature and additional nitrogen and their interaction significantly increased the glutelin relative content. • Temperature and nitrogen and their interaction increased the glutelin accumulation rate in the early and mid-filling periods. • Elevated temperature promoted the level of pro-glutelin whereas further additional nitrogen improved the level of α-glutelin. • Glutelin synthesis and maturity were regulated by elevated temperature at 5-20d and 20-30d after flowering, respectively. [ABSTRACT FROM AUTHOR]

Abstract Due to the sensitivity of chalk formation to high temperatures, the global warming will further threaten the improvement of high chalky grain rate in the southern japonica region of China. There have been many studies reported on the role of starch in the formation of chalkiness, but the relationship between nitrogen metabolites and chalkiness has not been given sufficient attention. In this study, we conducted the open field warming experiment and investigated the effects of temperature increase on grain protein metabolism. Results showed under the warming range of predicted global temperature increase (1.9 °C–4.0 °C), field warming apparently accelerated the development of grain endosperm, and the number of protein bodies were increased in both rice varieties, Furthermore, we clarified that the increased temperature regulated the key regulatory factors in protein synthesis and metabolism pathway, and the synthesis, transport, folding, assembly process of grain protein were further affected, and that further lead the component changes of grain storage protein. In conclusion, chalky traits are closely related to the grain storage materials and in this study, increased temperature induced the changes in the accumulation balance of storage proteins, amino acids and starches, which coordinately controls the formation of grain chalkiness. Highlights • We simulated climate warming with the field-warming system in actual rice field. • Endosperm and protein bodies were apparently increased under elevated temperatures. • Elevated temperature regulated regulatory factors in protein anabolism pathway. • Elevated temperatures affect balance of storage material and induce the chalkiness. • Proteins may play a certain role in chalkiness formation under elevated temperature. [ABSTRACT FROM AUTHOR]

This paper reports the effect of nitrogen on amino acid composition of leaf and grain for japonica rice during the grain filling stage, using pot experiments with three N rate treatments. Dynamic changes of free amino acids (FAA) and protein amino acids (PAA) in leaves and grains were monitored. Results showed that FAA in leaves decreased with the progress of grain filling. FAA in grains increased at first 14 days after anthesis and then decreased to a nearly constant level. PAA in leaves showed a diminishing trend as leaf senesced after anthesis. Conversely, PAA in grain exhibited an increasing trend with the progress of grain filling. N had significantly positive effect on contents of FAA and PAA in leaves and grains. There was no obvious trend of the variation of FAA in leaves and grains with N rates. PAA in leaves was more sensitive to N fertilization. By contrast, PAA in grains was more stable across N rates, especially at the late stage of grain filling. In addition, composition of leaf PAA was different from that of grain, indicating the reconstruction of amino acids during the process of translocation of N compounds from leaves to grains. [ABSTRACT FROM AUTHOR]

• We investigated the dynamics of internode dry matter (DM) from 21 days before anthesis to maturity for japonica rice. • We found a sharp reduction of internode DM between 20 and 30 days after anthesis (DAA). • Non-structural carbohydrates in internodes were significantly remobilized between 20–30 DAA. • A marked increase in internode DM at maturity was detected. • Internode DM can be viewed as an integrative index for source-sink balance. Storage of non-structural carbohydrates (NSC) in internode parenchyma before anthesis functions as a major carbon source for rice grain filling. In this study, two-year field and pot experiments with three japonica cultivars and two nitrogen (N) fertilization modes were performed, and dynamics of dry matter (DM) in internodes were quantified from 21 day before anthesis (DBA) to 60 day after anthesis (DAA; maturity). Results showed that in comparison to leaves and sheaths, DM in internodes varied dramatically, with a clear pattern of increasing at 21 DBA, peaking at 10 DAA, then dramatically dropping until 20 or 30 DAA, and finally refilling at maturity. The sharp reduction of dry matter in the internodes corresponded with the fastest rate of grain filling, indicating the importance of internode reserves for yield formation. On the other hand, the high rate of refilling in internodes at late stage implies the excess production of leaf photosynthetic assimilates. Similarly, starch content in internodes showed a pattern of decreasing at 20–30 DAA. To clarify the indicative meaning of internode reserves, we calculated their remobilization rate at 20 DAA and refilling rate at 60 DAA. Genotypic variations existed for the three cultivars, with a range of 15.17 %–36.37 % for remobilization rate and 54.36 %–104.33 % for refilling rates in 2018. Overall, N topdressing significantly increased the translocation of internode reserves for all three cultivars. Furthermore, our calculation showed that only about 10 % of the grain yield was formed during the second month of rice grain filling, indicating this period is less important for grain yield. Our findings help to clarify the significance of internode DM dynamics for rice management and should be beneficial to enhance our knowledge of source and sink relations, the fundamental component of crop yield formation. [ABSTRACT FROM AUTHOR]