This study tested the hypothesis that the interaction between polyamines and ethylene may mediate the effects of soil drying on grain filling of rice (Oryza sativa L.). Two rice cultivars were pot grown. Three treatments, well-watered, moderate soil drying (MD), and severe soil drying (SD), were imposed from 8 d post-anthesis until maturity. The endosperm cell division rate, grain-filling rate, and grain weight of earlier flowering superior spikelets showed no significant differences among the three treatments. However, those of the later flowering inferior spikelets were significantly increased under MD and significantly reduced under SD when compared with those which were well watered. The two cultivars showed the same tendencies. MD increased the contents of free spermidine (Spd) and free spermine (Spm), the activities of S-adenosyl-L-methionine decarboxylase and Spd synthase, and expression levels of polyamine synthesis genes, and decreased the ethylene evolution rate, the contents of 1-aminocylopropane-1-carboxylic acid (ACC) and hydrogen peroxide, the activities of ACC synthase, ACC oxidase, and polyamine oxidase, and the expression levels of ethylene synthesis genes in inferior spikelets. SD exhibited the opposite effects. Application of Spd, Spm, or an inhibitor of ethylene synthesis to rice panicles significantly reduced ethylene and ACC levels, but significantly increased Spd and Spm contents, grain-filling rate, and grain weight of inferior spikelets. The results were reversed when ACC or an inhibitor of Spd and Spm synthesis was applied. The results suggest that a potential metabolic interaction between polyamines and ethylene biosynthesis responds to soil drying and mediates the grain filling of inferior spikelets in rice.
Abscisic acid mediates the effect of post-anthesis soil drying on grain filling through regulating the activities of key enzymes and expressions of genes involved in sucrose-to-starch conversion in rice spikelets. This study investigated if abscisic acid (ABA) would mediate the effect of post-anthesis soil drying on grain filling through regulating the key enzymes in sucrose-to-starch conversion in rice (Oryza sativa L.) spikelets. Two rice cultivars were field-grown. Three treatments, well-watered (WW), moderate soil drying (MD), and severe soil drying (SD), were imposed from 6 days after full heading until maturity. When compared with those under the WW, grain filling rate, grain weight, and sink activity, in terms of the activities and gene expression levels of sucrose synthase, ADP glucose pyrophosphorylase, starch synthase, and starch branching enzyme, in inferior spikelets were substantially increased under the MD, whereas they were markedly decreased in both superior and inferior spikelets under the SD. The two cultivars showed the same tendencies. Both MD and SD increased ABA content and expression levels of its biosynthesis genes in spikelets, with more increase under the SD than the MD. ABA content was significantly correlated with grain filling rate and sink activities under both WW and MD, while the correlations were not significant under the SD. Application of a low concentration ABA to WW plants imitated the results under the MD, and applying with a high concentration ABA showed the effect of the SD. The results suggest that ABA plays a vital role in grain filling through regulating sink activity and functions in a dose-dependent manner. An elevated ABA level under the MD enhances, whereas a too high level of ABA under the SD decreases, sink activity.
In wheat stems, the levels of fructan-dominated water-soluble carbohydrates (WSC) do not always correlate well with grain yield. Field drought experiments were carried out to further explain this lack of correlation. Wheat (Triticum aestivum) varieties, Westonia, Kauz and c. 20 genetically diverse double haploid (DH) lines derived from them were investigated. Substantial genotypic differences in fructan remobilization were found and the 1-FEH w3 gene was shown to be the major contributor in the stem fructan remobilization process based on enzyme activity and gene expression results. A single nucleotide polymorphism (SNP) was detected in an auxin response element in the 1-FEH w3 promoter region, therefore we speculated that the mutated Westonia allele might affect gene expression and enzyme activity levels. A cleaved amplified polymorphic (CAP) marker was generated from the SNP. The harvested results showed that the mutated Westonia 1-FEH w3 allele was associated with a higher thousand grain weight (TGW) under drought conditions in 2011 and 2012. These results indicated that higher gene expression of 1-FEH w3 and 1-FEH w3 mediated enzyme activities that favoured stem WSC remobilization to the grains. The CAP marker residing in the 1-FEH w3 promoter region may facilitate wheat breeding by selecting lines with high stem fructan remobilization capacity under terminal drought.
This study was to test the hypothesis that polyamines (PAs) and ethylene and their interactions may be involved in mediating the post-anthesis development of spikelets in rice (Oryza sativa L.). Six rice cultivars differing in grain filling rate were field-grown, and the changing patterns of PAs and ethylene levels in rice spikelets during the filling and their relations with grain filling rates were investigated. The results showed that inferior spikelets had much greater ethylene evolution rate and 1-aminocylopropane-1-carboxylic acid (ACC) concentration than superior spikelets. Opposite to ethylene production, superior spikelets showed much higher freespermidine (Spd) and free-spermine (Spm) concentrations than inferior spikelets. Grain filling rate was very significantly and negatively correlated with ethylene evolution rate and ACC concentration, whereas positively correlated with free-Spd and free-Spm concentrations and with the ratio of free-Spd or free-Spm to ACC. Application of Spd, Spm, or aminoethoxyvinylglycine (an inhibitor of ethylene synthesis by inhibiting ACC synthesis) to panicles at the early grain filling stage significantly reduced ethylene evolution rate and ACC concentration, while significantly increased Spd and Spm concentrations, grain filling rate and grain weight of inferior spikelets. Application of ACC, ethephon (an ethylene-releasing agent), or methylglyoxal-bis (guanylhydrazone) (an inhibitor of Spd and Spm synthesis) showed the opposite effects. The results suggest that antagonistic interactions between PAs (Spd and Spm) and ethylene may be involved in mediating grain filling. A higher ratio of free-Spd or free-Spm to ethylene in rice spikelets could enhance grain filling.
Alternate wetting and moderate soil drying irrigation (WMD) has been widely adopted in rice (Oryza sativa L.) production for saving irrigation water and increasing grain yield. However, limited information is available about the effect of WMD on rice grain quality. The main purpose of this study was to evaluate the characteristics of grain quality in response to WMD. Two rice varieties, Yangdao 6 (YD6, indica) and Hanyou 8 (HY8, japonica), were grown in the field, with two water management treatments, well‐watered (WW) and WMD from 10 d after transplanting to maturity. In comparison with WW, WMD prominently increased grain yield by 7.57 to 9.72%, and improved some grain quality parameters such as milling, appearance, and eating and cooking qualities, including increases in head rice and decreases in chalkiness. The two varieties showed the same tendency. However, WMD had a negative impact on some nutritional quality traits in rice grains, such as reduction in the contents of amino acids and the micronutrient elements Cu, Fe, Mn, Mo, Se, and Zn. The WMD treatment also markedly decreased As content and exhibited no significant effect on Cd content in grains. The results demonstrated that WMD has an overall impact on rice grain quality with both positive and negative effects.
This study investigated the possibility that abscisic acid (ABA) may mediate the effect of post-anthesis soil drying on carbon remobilization from the stem and grain filling through regulating fructan hydrolysis and starch biosynthesis in wheat (Triticum aestivum L.). Two wheat cultivars showing difference in plant height were pot-grown. Three treatments, well watered (WW), moderate soil drying (MD) and severe soil drying (SD, were imposed from 9 days post-anthesis until maturity. The results showed that ABA contents in stems and grains, the remobilization of carbon reserves from the stem and its contribution to grain yield, and activities and gene expression levels of the enzymes involved in fructan hydrolysis in stems were increased with the increase in soil drying. When compared with the WW, the MD substantially increased, but the SD markedly decreased, grain filling rate, grain weight, and starch biosynthetic ability, in terms of activities and gene expression levels of ADP glucose pyrophosphorylase, soluble starch synthase, and starch branching enzyme in grains. The two cultivars showed the same tendency. ABA content was positively correlated with the remobilization of carbon reserves, and positively correlated with grain filling rate and starch biosynthetic ability under both WW and MD treatments. Exogenous application of a low concentration ABA to WW plants imitated the results under the MD, and applying with a high concentration ABA showed the effect of the SD. In summary, ABA enhances fructan hydrolysis in wheat stems and regulates starch biosynthesis in the grain.
Background Brassinosteroids (BRs) are a new group of plant hormones and play important roles in plant growth and development. However, little information is available if BRs could regulate spikelet development in rice (Oryza sativa L.) especially under soil-drying conditions. This study investigated whether and how BRs mediate the effect of soil-drying on spikelet differentiation and degeneration in rice. A rice cultivar was field-grown and exposed to three soil moisture treatments during panicle development, that is, well-watered (WW), moderate soil-drying (MD) and severe soil-drying (SD). Results Compared with the WW treatment, the MD treatment enhanced BRs biosynthesis in young panicles, increased spikelet differentiation and reduced spikelet degeneration. The SD treatment had the opposite effects. Changes in expression levels of key rice inflorescence development genes (OsAPO2 and OsTAW1), ascorbic acid (AsA) content, and activities of enzymes involved AsA synthesis and recycle, and amount of nonstructural carbohydrates (NSC) in young panicles were consistent with those in BRs levels, whereas hydrogen peroxide (H2O2) content showed opposite trend. Knockdown of the BRs synthesis gene OsD11 or application of a BRs biosynthesis inhibitor to young panicles markedly decreased OsAPO2 and OsTAW1 expression levels, BRs and AsA contents, activities of enzymes involved AsA synthesis and recycle, NSC amount in rice panicles and spikelet differentiation but increased the H2O2 content and spikelet degeneration compared to the control (the wide type or application of water). The opposite effects were observed when exogenous BRs were applied. Conclusions The results suggest that BRs mediate the effect of soil-drying on spikelet differentiation and degeneration, and elevated BRs levels in rice panicles promote spikelet development under MD by enhancing inflorescence meristem activity, AsA recycle and NSC partitioning to the growing panicles.
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