Enhanced non-structural carbohydrate metabolism and transport contribute to yield improvement in salt-tolerant rice under brine irrigation.

More than 4 billion inhabitants in Asia depend on rice for 35–60% of the calories consumed in their diets, but new rice cultivars frequently do not reach expected yields because of poor rice grain filling. Here, we quantified the activities of enzymes involved in starch metabolization in rice to investigate the mechanisms regulating the accumulation and translocation of stem non-structural carbohydrates (NSC) under different levels of nitrogen fertilizer application. A pot experiment was conducted using two rice cultivars, Liangyoupeijiu (LYPJ) and Shanyou63 (SY63), under high and low nitrogen applications. Compared with high nitrogen application (HN), low nitrogen application (LN) increased stem NSC concentration before the heading stage and NSC translocation during the grain filling stage; concomitantly, LN significantly shortened the active grain filling period and increased the grain filling rate in superior spikelets. Compared with the LYPJ cultivar, SY63 exhibited a higher grain weight, higher grain filling percentage, and higher stem NSC concentration before heading and greater NSC translocation after heading. During the period between panicle initiation and heading, the activities of adenosine diphosphate-glucose pyrophosphorylase (AGP), starch synthase (StS), and starch branching enzyme (SBE), all enzymes involved in starch synthesis, increased under the LN treatment and positively correlated with increases in stem NSC. During grain filling, the activities of enzymes involved in starch-to-sucrose conversion [α-amylase, β-amylase, and sucrose phosphate synthase (SPS)] increased under the LN treatment and positively correlated with stem NSC remobilization. Overall, the investigated enzymes exhibited higher activities in SY63 than in LYPJ. Our results suggest that low nitrogen increases the activities of AGP, StS, SBE, α-amylase, β-amylase, and SPS, leading to increased accumulation and remobilization of stem starch and NSC in SY63. We conclude that calculated reductions in nitrogen application and the choice of an appropriate cultivar may improve rice grain yields via enhanced stem NSC accumulation and translocation, thereby reducing the costs and increasing the sustainability of rice production.

Quantifying the effects of short-term heat stress at booting stage on nonstructural carbohydrates remobilization in rice

Remobilisation of non-structural carbohydrates (NSC) from leaves and stems and unloading into developing grains are essential for yield formation of rice. In present study, three recombinant inbred lines of rice, R91, R156 and R201 have been tested for source-flow-sink related attributes determining the nature of NSC accumulation and translocation at two nitrogen levels in the field. Compared to R91 and R156, R201 had lower grain filling percentage, harvest index, and grain yield. Meanwhile, R201 had significantly lower stem NSC translocation during grain filling stage. Grain filling percentage, harvest index, and grain yield showed the consistent trend with stem NSC translocation among the three lines. In comparison with R91 and R156, R201 had similarity in leaf area index, specific leaf weight, stem NSC concentration at heading, biomass, panicles m-2, spikelets per panicle, remobilization capability of assimilation in stems, sink capacity, sink activity, number and cross sectional area of small vascular bundles, greater number and cross sectional area of large vascular bundles, and higher SPAD, suggesting that source, flow, and sink were not the limiting factors for low stem NSC translocation and grain filling percentage of R201. However, R201 had significant higher stem and rachis NSC concentrations at maturity, which implied that unloading in the developing grains might result in low NSC translocation in R201. The results indicate that stem NSC translocation could be beneficial for enhancement of grain yield potential, and poor unloading into caryopsis may be the possible cause of low stem NSC translocation, poor grain filling and yield formation in R201.

Stem non-structural carbohydrates (NSCs) and its relationship with yield formation was investigated under low nitrogen (LN) and normal nitrogen (NN) treatments, using 46 recombinant inbred lines from Zhenshan 97 × Minghui 63 (Oryza sativa). Apparent contribution of transferred NSC to grain yield (AC(NSC) ) ranged from approximately 1 to 28% under LN and from 1 to 15% under NN. Concentration and total mass of NSC in stem (TM(NSC) ) at heading, apparent transferred mass of NSC (ATM(NSC) ) and AC(NSC) were larger under LN compared with NN. However, there was no significant difference in the apparent ratio of transferred NSC from stems to grain (AR(NSC) ). ATM(NSC) was positively correlated with grain yield, 1000-grain weight and AC(NSC) under both nitrogen levels, whereas AR(NSC) was highly correlated with harvest index and AC(NSC) . Leaf area contributed more strongly to grain yield compared with ATM(NSC) under both LN and NN. ATM(NSC) showed larger direct effects on grain yield under LN compared with NN. TM(NSC) at heading, small vascular bundles (SVBs) and spikelets per m(2) under LN had positive direct effects on ATM(NSC) . SVB and spikelets per m(2) under LN had larger and positive direct effects, and large vascular bundles had negative direct effects on AR(NSC) . TM(NSC) at heading and SVB under LN had positive direct effects on AC(NSC) . In brief, LN supply increased stem NSC accumulation and translocation to developing grain. Components of the source-sink-flow system showed different effects on NSC translocation and contribution to yield formation, depending on genotype and nitrogen level.

Non-structural carbohydrates (NSC) accumulation and photosynthesis traits were studied in two rice (Oryza sativa L.) genotypes maintained under control (22/30 °C - night/day) and at high night temperatures (HNT) (28/30 °C) conditions from heading to milk stage. Rice cultivars were Nagina22 - N22 and BRS Querência - Quer, which are tolerant and sensitive to high temperatures, respectively. The source-sink flow related attributes were tested to understand the nature of NSC accumulation and translocation. Compared to N22, Quer maintained higher stem starch in glucose on seventh day after heading and at milk stage independently of imposed temperatures. However, the levels of starch in glucose were lower for N22 meanwhile their total sugar concentration (TSC) were higher at control and at HNT at milk stage as compared to Quer. N22 maintained unaltered the spikelet sterility and 1000-grain weight across environments showing a consistent trend with its stem NSC translocation. Both genotypes showed similarity in some gas exchange and chlorophyll fluorescence performance suggesting unaffected photosystem II photochemistry, linear electron flux, and CO2 assimilation. Beyond indicating that source functioning was not the limiting factor for low TSC and starch in glucose levels found in N22 on seventh day after heading stage. Moreover, our data suggest that the higher translocation capacity shown by N22 can be involved in their lower spikelet sterility and 1000-grain weight stability across the environments. These results indicate that selecting genotypes with higher capacity to stem NSC translocation at HNT could lead to more grain yield stability in future climate scenarios.

Rice production is threatened by climate change, particularly heat stress (HS). Nonstructural carbohydrates (NSCs) remobilization is a key physiological mechanism that allows rice plants to cope with HS. To investigate the impact of short-term HS on the remobilization of nonstructural carbohydrates (NSCs) in rice, two cultivars (Huaidao-5 and Wuyunjing-24) were subjected to varying temperature regimes: 32/22/27 °C as the control treatment, alongside 40/30/35 °C and 44/34/39 °C, for durations of 2 and 4 days during the booting, flowering, and combined stages (booting + flowering) within phytotrons across the years 2016 and 2017. The findings revealed that the stem's NSC concentration increased, while the panicle's NSCs concentration, the efficiency of NSCs translocation from the stem, and the stem NSC contribution to grain yield exhibited a consistent decline. Additionally, sugar and starch concentrations increased in leaves and stems during late grain filling and maturity stages, while in panicles, the starch concentration decreased and sugar concentration increased. The heat-tolerant cultivar, Wuyunjing-24, exhibited higher panicle NSC accumulation under HS than the heat-sensitive cultivar, Huaidao-5, which had more stem NSC accumulation. The flowering stage was the most vulnerable to HS, followed by the combined and booting stages. Heat degree days (HDDs) were utilized to quantify the effects of HS on NSC accumulation and translocation, revealing that the flowering stage was the most affected. These findings suggest that severe HS makes the stem the primary carbohydrate storage sink, and alleviation under combined HS aids in evaluating NSC accumulation, benefiting breeders in developing heat-tolerant rice varieties.

The translocation of nonstructural carbohydrates (NSC) from leaf sheaths to filling grains after anthesis contributed greatly to the grain yield of cereal crops. In this study, the effect of nitrogen (N) supply levels on the accumulation and translocation of NSC in leaf sheath tissues and its relationship with the initiation and progression of leaf senescence during grain filling was investigated using two rice (Oryza sativa L.) genotypes, namely, premature flag leaf senescence mutant (psf) and its wild‐type. Three N treatment levels were used to examine N‐supply induced alteration in the activities of several key enzymes involved in NSC translocation and N assimilation in different leaf sheaths. The results show that the NSC translocation rate in leaf sheaths under low nitrogen (LN) treatment was significantly higher than those under normal nitrogen (NN) and high nitrogen (HN) treatments. However, the positive effect of LN on the NSC translocation in leaf sheath was closely associated with its negative effect on grain yield, due to accelerated leaf senescence and shortened leaf longevity. Comparatively, the upper‐positional sheath had a lower NSC amount and higher NSC translocation rate than the lower‐leaf sheaths after heading. High N suppressed sucrose‐phosphate synthase (SPS) activity in leaf sheaths, but enhanced the activity of key enzymes involving in N assimilation in leaf sheaths. The upper sheath had higher activity of sucrose‐metabolizing enzymes and lower activity of N‐assimilating enzymes. Hence, the upper‐leaf sheath had a relatively weak N assimilation and stronger NSC translocation than the lower‐leaf sheaths.

Excessive nitrogen application in rice production leads to low translocation efficiency of photosynthates in stems and lodging, resulting in inadequate grain filling and yield loss. The application of nanotechnology and growth regulators provides a feasible way to solve these problems. In this study, field experiments were conducted to explore the effects of foliar spraying of different concentrations of carbon dots and melatonin combinations on rice growth and development, photosynthesis, accumulation and translocation of non-structural carbohydrates (NSC) in stems and leaves, stem strength and yield under reduced nitrogen input. The results showed that the combination of carbon dots and melatonin improved rice yield and stem lodging resistance under the condition of 20% nitrogen reduction. Specifically, application of 0.3 mg mL-1 carbon dots and 100 μmol mL-1 melatonin increased the grain yield by 7.5%-26.8%. Foliar spraying of carbon dots and melatonin increased the photosynthetic rate and leaf area index of rice leaves, thereby increasing the content of stem NSC and plant biomass by 37.2%-153.8% and 4.5%-23.5%, respectively, and promoted NSC translocation from stems (51.9%-127.1% apparent transferred mass) and leaves (66.7%-161.6% apparent transferred mass) to grains by increasing the number and cross-sectional area of large and small vascular bundles of peduncle, the activities of α-amylase, β-amylase and SPS in stems, and upregulating the expression of sugar translocation related genes in stems, leaves and grains, and consequently improved the grain filling percentage and yield. In addition, foliar spraying of carbon dots and melatonin increased cellulose, hemicellulose and lignin contents in the basal stem by 10.8%-34.3%, 10.8%-42.1%, 17.2%-71.9%, and increased the basal internode stem diameter (+5.1%-10.8%), stem wall thickness (+6.3%-21.4%) and breaking resistance (+8.7%-66.7%), thus improving the lodging resistance of rice plants.

Nitrogen (N) fertilizer is a critical element that affects rice yield. However, its effects on dry matter accumulation (DMA), N accumulation, and their physiological mechanisms with grain yield and N utilization efficiency still lack in-depth study. Three large-scale japonica rice varieties—Jinxiangyu 1, Nanjing 46, and Huaidao 5—were used in two field experiments with varying N fertilizer application rates to examine grain yield and N utilization efficiency. The results showed that: (1) In the range of 0~360 kg ha−1 total N application rate (TNAR), the rice yields of the three cultivars were maximum under the TNAR at 270 kg ha−1. The optimal TNAR for the highest yield of Jinxiangyu 1, Nanjing 46, and Huaidao 5 were calculated based on quadratic regressions with values of 305.5 kg ha−1, 307.6 kg ha−1, and 298.0 kg ha−1, and the corresponding yields were 10.3 t ha−1, 10.6 t ha−1 and 10.2 t ha−1, respectively. The N utilization efficiency decreased gradually with the increase in TNAR, and the recovery efficiency decreased from 35.7~38.19% to 29.61~31.59%. (2) The yield was significantly positively correlated with DMA and N accumulation from the heading stage (HD) to the maturity stage (MA). The DMA and N accumulation of HD-MA were significantly positively correlated with leaf photosynthetic rate, non-structural carbohydrate (NSC) accumulation in stems, root oxidation activity, zeatin (Z) + zeatin riboside (ZR) contents in roots, and nitrate reductase (NR) and glutamate synthase (GOGAT) activity in HD. (3) In the range of 0~216 kg ha−1 panicle N application rate (PNAR), the rice yield was maximum under the PNAR at 108 kg ha−1. The optimal PNAR for the highest yield of Jinxiangyu 1 was calculated based on the quadratic regression with values of 139.5 kg ha−1, and the highest yield was 9.72 t ha−1. The leaf photosynthetic rate, NSC accumulation in stems, root oxidation activity, Z + ZR contents in roots, and NR activity in leaves in rice were higher under 108 kg ha−1 PNAR. Excessive application of panicle fertilizer reduced the above physiological indicators and rice yield. The above results showed that optimizing N fertilizer could increase the leaf photosynthetic rate, NSC accumulation in stems, root oxidation activity, Z + ZR contents in roots, and NR activity from HD to MA, which was beneficial to improving DMA and N uptake during HD-MA, thus improving grain yield and N utilization efficiency in rice.

Phloem unloading and loading are associated with stem non-structural carbohydrates (NSCs) accumulation and remobilization in rice (Oryza sativa L.). Four rice recombinant inbred lines (R032, R191, R046, and R146) derived from a cross between Zhenshan 97 and Minghui 63 were used to investigate the contributions of stem large and small vascular bundles (SVBs) to NSCs accumulation and translocation. Before heading, the parenchyma cells in stem cortex tissues (PCs) surrounding SVBs had higher starch density than those surrounding large vascular bundles (LVBs). Moreover, the protein levels of sucrose transporters (SUTs), cell wall invertase, sucrose synthase, and adenosine diphosphate glucose pyrophosphorylase, as well as the phloem plasmodesma densities were higher in SVBs than those in LVBs. After heading, starch density decreased more in PCs surrounding SVBs than in LVBs. Also, the protein levels of SUTs, α-amylase, sucrose phosphate synthase and sucrose synthase, the phloem plasmodesma densities in SVBs were higher than those in LVBs. The correlations of the number and total cross-sectional area of SVBs with mass and contribution to yield of transferred NSCs were higher than those of LVBs. Our results suggest that SVBs may have higher contributions to pre-anthesis stem NSCs accumulation and post-anthesis translocation than LVBs, which is potentially attributed to the high level of protein and enzyme involved in stem unloading and loading via apoplastic and symplastic pathways.

Non-structural carbohydrates (NSCs) are an important energy source for plant growth and metabolism. Analysis of NSC changes can provide important clues to reveal the adaptation mechanisms of plants to a specific environment. Although considerable differences have been reported in NSCs in response to nitrogen (N) application among crop species and cultivars, previous studies have mostly focused on the differences in leaves and stems. However, the effects of N on the characteristics of accumulation and translocation of NSC in maize with different levels of N tolerance remain unclear. To determine differences in the N levels, two cultivars (N-efficient ZH311 and N-inefficient XY508) were grown in field pots (Experiment I) and as hydroponic cultures (Experiment II) and were supplemented with different concentrations of N fertilizer. In both experiments, low-N stress significantly increased the accumulation of NSCs in maize vegetative organs and increased the translocation rate of NSCs in the stems and their apparent contribution to yield, thereby reducing the yield loss caused by low-N stress. N application had a greater effect on starch content in the vegetative organs of ZH311, but had less effect on soluble sugar (SS) and NSC content in the whole plant and starch content in the ears. ZH311 could convert more starch into SS under low N conditions to adapt to low N environments than XY508, while ensuring that grain yield and starch quantity were not affected. This is evidently an important physiological mechanism involved in this cultivar’s tolerance to low N conditions.

Grain-filling is a crucial process that determines final grain yield in rice (Oryza sativa L.). To understand the genetic basis of dynamics of grain-filling, quantitative trait locus (QTL) analysis was conducted using time-related phenotypic data on grain-filling collected from a population of 155 recombinant inbred lines (F12), derived from a cross between Milyang 23 and Akihikari. Two QTLs detected on chromosomes 8 and 12 were strongly associated with increased filling percentage per panicle. These QTLs were not linked with those controlling spikelet numbers per panicle. This result confers the possibility of improving grain-filling together with an enlargement of sink size. The QTL for filling percentage per panicle on chromosome 8 exactly overlapped that for non-structural carbohydrate (NSC) content in the culm and leaf sheaths during grain-filling, and the Milyang 23 allele associated with increased grain-filling percentage per panicle was associated with decreased NSC content. Therefore, this QTL may be directly involved in NSC translocation from the culm and leaf sheaths to panicle. In addition, the Milyang 23 alleles of QTLs associated with greater spikelet number per panicle on chromosomes 1 and 6 were also related with a reduction in NSC content in the culm and leaf sheaths during grain-filling. These results indicate that NSC dynamics during grain-filling is partly dependent on sink size. NSC accumulation in the culm and leaf sheaths at the heading stage was mainly controlled by different genetic regulations from NSC dynamics during grain-filling. Nitrogen dynamics during grain-filling may also be involved in carbohydrate dynamics.

Delaying panicle nitrogen application to emergence of 3rd leaf from flag leaf increases the grain-filling ability and yield of large-panicle rice by increasing stem nonstructural carbohydrates at heading

Non-structural carbohydrates (NSC) play an important role in yield formation. In this paper, the relationships of NSC accumulation and translocation with yield formation were investigated under elevated CO2 concentrations ([CO2]) and nitrogen (N) application rates. A japonica rice (Oryza sativa L.) cultivar, “Nanjing 9108,” was grown at three [CO2]—Ambient (T0), ambient + 160 μmol·mol−1 (T1), and ambient + 200 μmol·mol−1 (T2)—in open-top chambers (OTC), with three levels of N fertilizer application rates—10 gN·m−2 (N1), 20 gN·m−2 (N2), and 30 gN·m−2 (N3)—Which were set in OTCs using pot experiments. The results showed that the concentration of NSC (CNSC) and the total mass of NSC stored in stems (TMNSC) under T1 and T2 were higher than those in the T0 treatment, and the CNSC and TMNSC of N3 were lower than those of N1 and N2 at the heading stage. The CNSC and the TMNSC were significantly positively correlated with the stem biomass during the growth period and were notably negatively correlated with the N content in leaves (Nleaf) at the heading and filling stages. The seed setting rate was significantly positively related to the apparent transferred mass of NSC from stems to grains (ATMNSC) at the filling stage, while the relationship between yield and the ATMNSC was not statistically significant. Although there was no difference in the apparent contribution of transferred NSC to grain yield (ACNSC) between treatments, NSC stored in stems further accumulated obviously during the late filling stage, implying that the grain yield of “Nanjing 9108” was predominantly sink-limited. We concluded that elevated [CO2] improved the concentration of NSC at the rice heading stage. The interaction between elevated [CO2] and N fertilizer rates significantly influenced the concentration of NSC at the filling stage. Rice stems NSC reaccumulated at the late grain filling stage, which implies a restriction on NSC transference to grain.

Use of marginal coastal saline lands for rice cultivation is one of the options to sustain the net rice cropped area and rice production. On-farm farmer’s participatory field trial was conducted in Bangladesh during 2012 and 2013 in non-gher and gher of medium saline ecosystems. The objective of the study was to identify suitable Boro variety for medium saline soils. Salt tolerant rice varieties, BRRI dhan47 and Binadhan-8 were evaluated along with widely adopted BRRI dhan28 in non-gher area. BRRI dha47, Binadhan-8 and Binadhan-10 were tested in saline ghers. Crops were irrigated with water of 1.89 to 3.5 dS m-1 at different growth stages respectively. In saline non-gher, where soil salinity went up to 6.35 dS m-1, the highest grain yield was produced by Binadhan-8 (4.57 t ha-1) followed by BRRI dhan28 (4.23 t ha-1). The highest grain yield was recorded in Binadhan-10 (5.14 t ha-1) in ghers which was 6 and 12% higher than BRRI dhan47 and Binadhan-8 respectively at soil salinity of 7.43 dS m-1. Farmers preferred Binadhan-10 as a salt tolerant variety for its higher grain yield and medium grain quality.Bangladesh Rice j. 2014, 18(1&2): 18-23