The causes and impacts for heat stress in spring maize during grain filling in the North China Plain--A review

High-temperature stress （HTS） at the grain-filling stage in spring maize （Zea mays L.） is the main obstacle to increasing productivity in the North China Plain （NCP）. To solve this problem, the physiological mechanisms of HTS, and its causes and impacts, must be understood. The HTS threshold of the duration and rate in grain filling, photosynthetic characteristics （e.g., the thermal stability of thylakoid membrane, chlorophyll and electron transfer, photosynthetic carbon assimilation）, water status （e.g., leaf water potential, turgor and leaf relative water content） and signal transduction in maize are reviewed. The HTS threshold for spring maize is highly desirable to be appraised to prevent damages by unfavorable temperatures during grain filling in this region. HTS has negative impacts on maize photosynthesis by damaging the stability of the thylakoid membrane structure and degrading chlorophyll, which reduces light energy absorption, transfer and photosynthetic carbon assimilation. In addition, photosynthesis can be deleteriously affected due to inhibited root growth under HTS in which plants decrease their water-absorbing capacity, leaf water potential, turgor, leaf relative water content, and stomatal conductance. Inhibited photosynthesis decrease the supply of photosynthates to the grain, leading to falling of kernel weight and even grain yield. However, maize does not respond passively to HTS. The plant transduces the abscisic acid （ABA） signal to express heat shock proteins （HSPs）, which are molecular chaperones that participate in protein refolding and degradation caused by HTS. HSPs stabilize target protein configurations and indirectly improve thylakoid membrane structure stability, light energy absorption and passing, electron transport, and fixed carbon assimilation, leading to improved photosynthesis. ABA also induces stomatal closure to maintain a good water status for photosynthesis. Based on understanding of such mechanisms, strategies for alleviating HTS at the grain-filling stage in spring maize are summarized. Eight strategies have the potential to improve the ability of spring maize to avoid or tolerate HTS in this study, e.g., adjusting sowing date to avoid HTS, breeding heat-tolerance varieties, and tillage methods, optimizing irrigation, heat acclimation, regulating chemicals, nutritional management, and planting geometric design to tolerate HTS. Based on the single technology breakthrough, a com- prehensive integrated technical system is needed to improve heat tolerance and increase the spring maize yield in the NCP.

Changes in Activities of the Key Enzymes Related to Starch Synthesis in Rice Grains During Grain Filling and Their Relationships with the Filling Rate and Cooking Quality

With 10 rice cultivars (lines) as materials, the changes in activities of adenosine diphosphoglucose pyrophosphorylase(ADPGPase), starch synthase (SSase) and starch branching enzyme (Q-enzyme) in the grains during grain filling, and theirrelationships with the filling rate, gel consistency (GC), alkali spreading value (ASV) and amylose content (AC) werestudied. The results showed that changes in activities of ADPGPase, SSase and Q-enzyme exhibited a single peak duringgrain filling, and the time of the activity peaks for the former two enzymes was earlier than that of the maximum grain-fillingrate (Tmax), and the time reaching the peak for Q-enzyme was synchronous with Tmax. The activities at early grain fillingstage, and the mean and maximum activities of each enzyme during grain filling period were positively and significantly orvery significantly correlated with the mean and maximum grain filling rate and starch content (mg grain-1) in the grains.Activities of ADPGPase at all grain filling stages and those of Q-enzyme at the early and mid filling stages were notsignificantly correlated the cooking quality (GC, ASV and AC). SSase activities at the early filling stage were significantlyand negatively correlated with GC and ASV, and positively correlated with AC. Activities of SSase at mid and late grainfilling stages and Q-enzyme at the late filling stage were significantly and positively correlated with GC and ASV, andnegatively correlated with AC. Spraying zeatin or abscisic acid at early grain filling stage could obviously regulate theactivities of ADPGPase, SSase and Q-enzyme in the grains.

Changes in Enzyme Activities Involved in Starch Synthesis and Hormone Concentrations in Superior and Inferior Spikelets and Their Association with Grain Filling of Super Rice

The changes in activities of key enzymes involved in sucrose-to-starch conversion and concentrations of hormones in superior and inferior spikelets of super rice were investigated and their association with grain filling was analyzed.Four super rice cultivars,Liangyoupeijiu,IIyou 084,Huaidao 9 and Wujing 15,and two high-yielding and elite check cultivars,Shanyou 63 and Yangfujing 8,were used.The activities of sucrose synthase （SuSase）,adenosine diphosphoglucose pyrophosphorylase （AGPase）,starch synthase （StSase） and starch branching enzyme （SBE）,and the concentrations of zeatin ＋ zeatin riboside （Z ＋ ZR）,indole-3-acetic acid （IAA） and abscisic acid （ABA） in superior and inferior spikelets were determined during the grain filling period and their relationships with grain filling rate were analyzed.Maximum grain filling rate,the time reaching the maximum grain-filling rate,mean grain filling rate and brown rice weight for superior spikelets showed a slight difference between the super and check rice cultivars,but were significantly lower in the super rice than in the check rice for inferior spikelets.Changes of enzyme activities and hormone concentrations in grains exhibited single peak curves during the grain filling period.The peak values and the mean activities of SuSase,AGPase,StSase and SBE were lower in inferior spikelets than in superior ones,as well as the peak values and the mean concentrations of Z ＋ ZR and IAA.However,the peak value and the mean concentration of ABA were significantly higher in inferior spikelets than in superior ones and greater in the super rice than in the check rice.The grain filling rate was positively and significantly correlated with the activities of SuSase,AGPase and StSase and the concentrations of Z ＋ ZR and IAA.The results suggested that the low activities of SuSase,AGPase and StSase and the low concentrations of Z ＋ ZR and IAA might be important physiological reasons for the slow grain filling rate and light grain weight of inferior spikelets in super rice.

Combined effect of shading time and nitrogen level on grain filling and grain quality in japonica super rice

There is limited information about the combined effect of shading time and nitrogen （N） on grain filling and quality of rice. Therefore, two japonica super rice cultivars, Nanjing 44 and Ningjing 3, were used to study the effect of shading time and N level on the characteristics of rice panicle and grain filling as well as the corresponding yield and quality. At a low N level （150 kg N ha^-1, 150N）, grain yield decreased （by 21.07-26.07%） under the treatment of 20 days of shading before heading （BH） compared with the no shading （NS） treatment. These decreases occurred because of shortened panicle length, decreased number of primary and secondary branches, as well as the grain number and weight per panicle. At 150N, in the treatment of 20 days of shading after heading （AH）, grain yield also decreased （by 9.46-10.60%） due to the lower grain weight per panicle. The interaction of shading and N level had a significant effect on the number of primary and secondary branches. A high level of N （300 kg N ha^-1, 300N） could offset the negative effect of shading on the number of secondary branches and grain weight per panicle, and consequently increased the grain yield in both shading treatments. In superior grains, compared with 150N NS, the time to reach 99% of the grain weight （T99） was shortened by 1.6 to 1.7 days, and the grain weight was decreased by 4.18-5.91% in 150N BH. In 150N AH, the grain weight was 13.39-13.92% lower than that in 150N NS due to the slow mean and the maximum grain-filling rate （GRmean and GRmax）. In inferior grains, grain weight and GRmean had a tendency of 150N NS〉150N BH〉150N AH. Under shaded conditions, 300N decreased the grain weight due to lower GReen both in superior and inferior grains. Compared with 150N NS, the milling and appearance qualities as well as eating and cooking quality were all decreased in 150N BH and 150N AH. Shading with the high level of 300N improved the milling quality and decreased the number of chalky rice kernels, but the eating and cooking quality was reduced with increased chalky area and overall chalkiness. Therefore, in the case of short term shading, appropriate N fertilizer could be used to improve the yield and milling quality of rice, but limited application of N fertilizer is recommended to achieve good eating and cooking quality of rice.

Effects of nitrogen fertilizer and chemical regulation on spring maize lodging characteristics, grain filling and yield formation under high planting density in Heilongjiang Province, China

Now,lodging is a major constraint factor contributing to yield loss of maize (Zea mays L.) under high planting density.Chemical regulation and nitrogen fertilizer could effectively coordinate the relationship between stem lodging and maize yield,which significantly reduce lodging and improve the grain yield.The purpose of this study was to explore the effects of chemical regulation and different nitrogen application rates on lodging characteristics,grain filling and yield of maize under high density.For this,we established a field study during 2017 and 2018 growing seasons,with three nitrogen levels of N100 (100 kg ha^(–1)),N200 (200 kg ha^(–1)) and N300 (300 kg ha^(–1)) at high planting density (90 000 plants ha^(–1)),and applied plant growth regulator (Yuhuangjin,the mixture of 3% DTA-6 and 27% ethephon) at the 7th leaf.The results showed that chemical control increased the activities of phenylalanine ammonia-lyase (PAL),tyrosine ammonia-lyase (TAL),4-coumarate:Co A ligase (4CL),and cinnamyl alcohol dehydrogenase (CAD),and increased the lignin,cellulose and hemicellulose contents at the bottom of the 3rd internode,which significantly reduced the lodging percentage.The lignin-related enzyme activities,lignin,cellulose and hemicellulose contents decreased with the increase of nitrogen fertilizer,which significantly increased the lodging percentage.The 200 kg ha^(–1) nitrogen application and chemical control increased the number,diameter,angle,volume,and dry weight of brace roots.The 200 kg ha^(–1) nitrogen application and chemical control significantly increased the activities of ADP-glucose pyrophosphorylase (AGPase),soluble starch synthase (SSS) and starch branching enzyme(SBE),which promoted the starch accumulation in grains.Additional,improved the maximum grain filling rate (V_(max)) and mean grain filling rate (V_(m)),which promoted the grain filling process,significantly increased grain weight and grain number per ear,thus increased the final yield.

Top-grain filling characteristics at an early stage of maize(Zea mays L.) with different nitrogen use efficiencies

Maize genotypes vary significantly in their nitrogen use efficiencies（NUEs）.Better understanding of early grain filling characteristics of maize is important,especially for maize with different NUEs.The objectives of this research were（i）to investigate the difference in apical kernel development of maize with different NUEs,（ii）to determine the reaction of apical kernel development to N application levels,and（iii）to evaluate the relationship between apical kernel development and grain yield（GY）for different genotypes of maize.Three maize hybrid varieties with different NUEs were cultivated in a field with different levels of N fertilizer arranged during two growing seasons.Kernel fresh weight（KFW）,volume（KV）and dry weight（KDW）of apical kernel were evaluated at an early grain filling stage.Ear characteristics,GY and its components were determined at maturity stage.Apical kernel of the high N and high efficiency（HN-HE）type（under low N,the yield is lower,and under higher N,the yield is higher）developed better under high N（N210 and N240,pure N of 210 and 240 kg ha^–1）than at low N（N120 and N140,pure N of 120 and 140 kg ha^–1）.The low N and high efficiency（LN-HE）type（under low N,the yield is higher,while under higher N,the yield is not significantly higher）developed better under low N than at high N.The double high efficiency（D-HE）type（for both low and high N,the yield is higher）performed well under both high and low N.Apical kernel reacted differently to the N supply.Apical kernel developed well at an early grain filling stage and resulted in a higher kernel number（KN）,kernel weight（KW）and GY with better ear characteristics at maturity.

Mapping QTL for Heat-Tolerance at Grain Filling Stage in Rice

A mapping population of 98 lines (backcross inbred lines, BILs) derived from a backcross of Nipponbare/Kasalath// Nipponbare was planted at two experimental sites, Nanjing and Nanchang, and treated with high and optimal temperature during grain filling, respectively. The grain weight heat susceptibility index [GWHSI= (grain weight at optimum temperature-grain weight at high temperature) / grain weight at optimum temperature × 100] was employed to evaluate the tolerance of rice to heat stress. A genetic linkage map with 245 RFLP markers and a mixed linear-model approach was used to detect quantitative trait loci (QTLs) and their main effects, epistatic interactions and QTL× environment interactions (Q×E). The threshold of LOD score=2.0 was used to detect the significance of association between marker and trait. A total of 3 QTLs controlling heat tolerance during grain filling were detected, on chromosomes 1, 4 and 7, with LOD scores of 8.16, 11.08 and 12.86, respectively, and they explained the phenotypic variance of 8.94, 17.25 and 13.50 %, correspondingly. The QTL located in the C1100-R1783 region of chromosome 4 showed no QTL× environment interaction and epistatic effect, suggesting that it could be stably expressed in different environments and genetic backgrounds, and thus it would be valuable in rice breeding for heat tolerance improvement. This QTL allele, derived from Kasalath reduced 3.31% of the grain weight loss under heat stress. One located between R1613-C970 on chromosome 1 and the other between C1226-R1440 on chromosome 7, with additive effect 2.38 and 2.92%, respectively. The tolerance alleles of both these QTLs were derived from Nipponbare. Both of these QTLs had significant QTL× environment interactions, and the latter was involved in epistatic interaction also. Eight pairs of epistatic effect QTLs were detected, one pair each on chromosomes 1,2,3, 5, 7, 8, 10 and 12. The results could be useful for elucidating the genetic mechanism of heat-tolerance and the development of new rice varieties with heat tolerance during grain filling phase.

Relationship Between Lodging Resistance and Chemical Contents in Culms and Sheaths of Japonica Rice During Grain Filling

To understand the relationship between lodging resistance and chemical component contents in culms and leaf sheaths of rice, the physical strength and maximum bearing capacity of culm, and the contents and amounts of potassium （K） silicon （Si） and soluble sugars in culms and leaf sheaths were investigated using four japonica rice varieties with different lodging resistance characteristics during grain filling. There were significant differences in the total amounts of K, Si and soluble sugars in culms and leaf sheaths among the tested rice varieties. The difference in the total amount of Si was greater than that of K or soluble sugars. The physical strength and maximum bearing capacity of culm continuously decreased from heading to ripening, with a rapid decrease at the dough stage. However, the contents and total amounts of K and Si in culms and the Si content in leaf sheaths gradually increased and an accumulation of K and Si in culms was exhibited, whereas the content and total amount of K and the total amount of Si in leaf sheaths gradually decreased and an exportation of K and Si in leaf sheaths was presented. The physical strength was positively and significantly correlated with the total amounts of K and Si in culms during grain filling except that at the heading stage, the total amount of soluble sugars in culms at the heading and milky stages, the total amounts of Si and soluble sugars in leaf sheaths at the heading stage, the total amount of K in leaf sheaths at the heading and milky stages, and the maximum bearing capacity during grain filling. It is suggested that the lodging resistance of japonica rice would be improved by increasing the amount of soluble sugars in plants at the early filling stage, and enhancing the amount of Si in plants during grain filling through topdressing Si fertilizer at the early filling stage.

Effects of High Temperature on Antioxidant Enzymes and Lipid Peroxidation in Flag Leaves of Wheat During Grain Filling Period

On the basis of the phytotron, the effects of high temperature （daily average temperature 25, 30, 35 and 40℃, respectively） on antioxidant enzymes and lipid peroxidation in flag leaves of wheat at 50% relative air moisture during grain fastest filling stage [19-21 days after anthesis （DAA）] were studied. The wheat cultivars tested were Yangmai 9 with weak-gluten and Yangmai 12 with medium-gluten. Compared with 25℃, the higher the temperature was, the higher was the MDA content in flag leaves, while lower were the SOD, POD, and CAT activities. SOD and CAT activities in Yangmai 12 appeared to be more sensitive to high temperature than that in Yangmai 9. But POD activity in Yangmai 12 was less sensitive to high temperature. MDA content in Yangmai 12 was higher than that in Yangmai 9. The 1000-grain weight declined with increase in temperature.

Effects of Panicle Nitrogen Fertilization on Non-Structural Carbohydrate and Grain Filling in Indica Rice

Grain filling, a crucial stage of grain yield formation in rice, is usually affected by the panicle nitrogen （N） fertilization. Field and pot culture experiments were conducted to explore the underlying mechanisms of N effect. Two rice cultivars with high lodging resistance were grown in the field and pot. Four panicle N fertilization treatments were conducted in 2006 and repeated in 2007. The result showed that medium level of panicle N fertilization treatment （NM） enhanced the accumulation and translocation of non-structural carbohydrate （NSC） in the stem and sheath. Compared with non-nitrogen treatment （NO）, NM promoted the translocation of labeled ^13C from stem and sheath to grain. But, low level of panicle N fertilization treatment （NL） and high level of panicle N fertilization treatment （NH） showed the negative effect. The endosperm cell, grain length, and grain width of NM increased more quickly than that of NO from 4 to 10 d after anthesis. During the early period of grain filling, sucrose-phosphate synthase （EC 2.4.1.14, SPS） activity were significantly higher for the NM treatment than those of the NL and NH treatments. Sucrose synthase （EC 2.4.1.13, SuSase） activity in the grains was substantially enhanced by NM, with the duration of higher activity being longer than those of the other treatments. At maturing stage, NM significantly increased the filled grain number, the seed-setting rate, and the grain weight compared with NL and NH. The results suggest that NM have a positive effect on the activities of enzymes of physiological importance, thereby increasing the grain size and promoting grain filling.

Effect of Temperature at Grain Filling Stage on Activities of Key Enzymes Related to Starch Synthesis and Grain Quality of Rice

Three japonica rice varieties with different cooking and eating quality were grown at high temperature in the greenhouse and natural field. Effects of temperature at the grain filling stage on these varieties were investigated in terms of the activities of key enzymes related to starch synthesis and cooking and eating quality of rice grain. The high temperature at the grain filling stage increased protein content, and decreased amylose content and taste meter value of rice; inferior grain quality varieties showed a greater magnitude of the increase or decrease than the superior ones. Reaction of rapid visco analyser profiles to the temperature varied with rice varieties. The activities of adenosine diphosphoglucose pyrophosphoryiase （AGPP）, soluble starch synthase （SSS） and starch branching enzyme （SBE） gradually increased to a peak value, and thereafter declined as grain filling progressed. Enzyme activities in different varieties differed in a same filling stage, and also in the time when the enzyme activity reached a maximum. AGPP and SSS were insensitive to the environmental temperature, but SBE was comparatively sensitive to the temperature, and its activity declined when temperature was too high or too low.

Effects of Early- and Late-Sowing on Starch Accumulation and Associated Enzyme Activities During Grain Filling Stage in Rice

The environmental temperature occurring during the grain filling stage is an important factoraffecting starch synthesis and accumulation in rice. We investigated starch accumulation, amylaseactivity and starch granule size distribution in two low-amylose japonica rice varieties, Nanjing 9108 andFujing 1606, grown in the field at different filling temperatures by manipulating sowing date. The two ricevarieties exhibited similar performances between two sowing dates. Total starch, amylose andamylopectin contents were lower at the early-filling stage of T1 treatment (Early-sowing) compared withthose at the same stage in T2 treatment (Late-sowing). In contrast, at the late-filling stage, when fieldtemperatures were generally decreasing, total starch and amylopectin contents in T1 were highercompared to those in T2. The ideal temperature for strong activity of ADP-glucose pyrophosphorylaseand soluble starch synthase was about 22℃. A higher temperature from the heading to maturity stagesin T1 increased the activities of starch branching enzyme and suppressed the activities of granule boundstarch synthetase and starch debranching enzyme. We found that rice produced larger-sized starchgranules under the T1 treatment. These results suggested that due to the early-sowing date, the hightemperature (30℃) occurring at the early-filling stage hindered starch synthesis and accumulation,however, the lower temperatures (22 ℃) at the late-filling stage allowed starch synthesis and accumulationto return to normal levels.

Changes in Activities of Key Enzymes for Starch Synthesis and Glutamine Synthetase in Grains of Progenies from a Rice Cross During Grain Filling

The progenies differed in amylose and protein contents in grains, which derived from a rice cross, Dongnong 423×Toukei 180, were used to study changes in the activities of ADP-glucose pyrophosphorylase （AGPP）, soluble starch synthetase （SSS）, starch branching enzyme （SBE） and glutamine synthetase （GS） in rice grains during grain filling. The activities of AGPP, SSS and SBE gradually increased and then declined as a single-peak curve with the process of grain filling in the progenies with high and low amylose contents in grains. The progenies with high amylose content peaked earlier in the AGPP, SSS and SBE activities and had higher AGPP, SSS and SBE activities at the early grain filling stage than those with low amylose content. The GS activity peaked earlier and was higher at the late stage of grain filling in the progenies with high protein content than in those with low protein content. It is suggested that the activities of key enzymes for starch synthesis and glutamine synthetase could be changed in oriented breeding for amylose and protein contents in grains.

Effect of Different Irrigation Patterns on Transportation and Allocation of Carbohydrate During Grain Filling of Rice

Using intersubspecific hybrid rice Xieyou 9308 and Liangyou Peijiu as the tested materials, the effects of differentirrigation patterns on transportation and allocation of carbohydrate during grain filling stage was observed by the designwith three level of soil water content in irrigated field. The results showed that in the conventional flooding and thealternate dry-watering cultivations, the exported rate of stored carbohydrate from stem and photophate from the leaves were60 and 90 % respectively. The exported rate of carbohydrate was decreased significantly (P<0.01) in the non-floodingcultivation. There was no significant difference between the conventional flooding and dry-wet alternation treatments interm of the exported rate of carbohydrate. The filling grains were the major sinks for carbohydrate storage during grainfilling stage. Grains received nearly 50% of stored carbohydrate from leaf sheath and 80% of photophate from leaves. Atthe nonflooding condition the absorbing rates of grains were significantly decreased by 10 % from leaf sheath and 20 %from leaves photophate. Water stress leaded much decrease in absorbing ability for inferior grains, which might be one ofthe main causes for low seed-settling rate in non-flooding cultivation.

Relationship Between Variation in Activities of Key Enzymes Related to Starch Synthesis During Grain Filling Period and Quality of Eating and Cooking in Rice

Four japonica rice varieties with significant differences in quality of eating and cooking were used in the experiment. The varieties showed differences in amylose and amylopectin contents at different grain filling stages, which were attributed to the accumulative speed of starch at different grain filling stages. During grain filling period, the varieties had no difference in the time when the activities of ADPglocose pyrophosphorylase （AGPP） and soluble starch synthesis （SSS） reached a maximum, but had difference in the time when the activity of starch branching enzyme （SBE） reached a maximum, in which the inferior quality varieties were earlier than the high quality ones, and high quality varieties still kept high enzyme activities at the late stage of grain filling. The correlation and correlative degree between AGPP, SSS, SBE and amylose content, amylopectin content, taste meter value, and RVA properties varied with the different stages of grain filling. The correlation between SSS activity and taste meter value was not significant during the whole period of grain filling, but the activities of AGPP and SBE had significant or highly significant correlation with taste meter value. It was helpful for improving quality of eating and cooking of japonica rice to use the materials with low enzyme activity at the early stage of grain filling or high enzyme activity at the late stage as parents.

The Effect of Plastic-Covered Ridge and Furrow Planting on the Grain Filling and Hormonal Changes of Winter Wheat

Although plastic-covered ridge and furrow planting（RF） has been reported to produce substantial increases in the grain weight of winter wheat,the underlying mechanism is not yet understood.The present study used two cultivars,Xinong 538 and Zhoumai 18,and RF and traditional flatten planting（TF,control） with the objective of investigating the effect of RF on wheat grain filling and the possible relationship of hormonal changes in the wheat grains under RF to grain filling.The results indicated that RF significantly increased the grain weight,although the effects on grain filling were different： RF significantly increased the grain-filling rate and grain weight of inferior grains,whereas RF had no significant effect on grainfilling rate and grain weight of superior grains.The final grain weight of inferior grains under RF was 39.1 and 50.7 mg for Xinong 538 and Zhoumai 18,respectively,3.6 and 3.4 mg higher than the values under TF.However,the final grain weight of superior grains under RF was only 0.6 and 0.8 mg higher than under TF for Xinong 538 and Zhoumai 18,respectively.RF significantly decreased the ethylene and gibberellic acid content in the inferior grains and increased the indole-3-acetic acid,abscisic acid and zeatin ＋ zeatin riboside content in the inferior grains;however,no significant difference between RF and TF was observed for the hormonal content in the superior grains.Based on these results,we concluded that RF significantly modulated hormonal changes in the inferior grains and,thus,affected the grain filling and grain weight of the inferior grains;in contrast,RF had no significant effect on grain filling,grain weight and hormonal changes in the superior wheat grains.

Variations in Carbohydrate and Protein Accumulation among Spikelets at Different Positions Within a Panicle During Rice Grain Filling

The accumulation dynamics of kernel components for spikelets at different positions within a rice panicle were investigated during grain filling to understand the physiological reasons for the variation of grain quality.Two rice cultivars,Yangdao 6 （indica） and Yangjing 9538 （japonica）,were field-grown,and the grain filling characters and contents of starch,soluble sugar,and protein of the spikelets at different positions were studied.There were significant differences in matter accumulation among spikelets at different positions during grain filling.The early-flowering spikelets presented dominance over the late-flowering spikelets in initial time and initial rate of accumulation.At the initial and mid filling stages,the contents and the rates of starch and amylose accumulation in spikelets decreased with the flowering sequence,but soluble sugar content （SSC） exhibited the opposite trend.The difference in SSC among the spikelets of Yangjing 9538 was greater than that of Yangdao 6,but amylose content in mature spikelets showed no obvious relationship to their flowering sequence.The crude protein content （CPC） of early-flowering spikelets decreased more rapidly than that of late-flowering ones at the initial filling stage,and CPC in the spikelets on the secondary branch was higher than that on the primary branch,but CPC in early-flowering ones was lower than that in late-flowering across the whole grain filling period.Grain water content （GWC） of early-flowering spikelets decreased more rapidly than that of late-flowering spikelets on the same branch at the initial and mid filling stages,especially for the top grain on each primary branch.The results suggested that poor grain filling of late-flowering spikelets may be attributed to their low biological activity rather than carbohydrate supply limitation.

Nitrogen application affects maize grain filling by regulating grain water relations

Grain water relations play an important role in grain filling in maize. The study aimed to gain a clear understanding of the changes in grain dry weight and water relations in maize grains by using hybrids with contrasting nitrogen efficiencies under differing nitrogen levels. The objectives were: 1) to understand the changes in dry matter and percent moisture content(MC) during grain development in response to different nitrogen application rates and 2) to determine whether nitrogen application affects grain filling by regulating grain water relations. Two maize hybrids, high N-efficient Zhenghong 311(ZH311) and low N-efficient Xianyu 508(XY508), were grown in the field under four levels of N fertilizer: 0, 150, 300, and 450 kg N ha;during three growing seasons. Dry weight, percent MC and water content(WC) of basal-middle and apical grains were investigated. The difference in the maximum WC and filling duration of basal-middle and apical grains in maize ears resulted in a significant difference in final grain weight. Grain position markedly influenced grain drying down;specifically, the drying down rate of apical grains was faster than that of basal-middle grains. Genotype and grain position both influenced the impact of nitrogen application rate on grain filling and drying down. Nitrogen rate determined the maximum grain WC and percent MC loss rate in the middle and the late grain-filling stages, thus affecting final grain weight. The use of high N-efficient hybrids, combined with the reduction of nitrogen application rate, can coordinate basal-middle and apical grain drying down to ensure yield. This management strategy could lead to a win-win situation in which the maximum maize yield, efficient mechanical harvest and environmental safety are all achieved.

Identification of microRNAs regulating grain filling of rice inferior spikelets in response to moderate soil drying post-anthesis

The grain filling of inferior spikelets is much less complete than that of superior spikelets in rice cultivars with large panicles and numerous spikelets and is promoted by moderate soil drying(MD)post-anthesis.A growing body of evidence has shown that microRNAs function in regulating grain development.However,little is known about the mechanism of microRNA control of grain filling of inferior spikelets in response to MD.In this study,grain filling of inferior spikelets was promoted by MD treatment in Nipponbare.Small-RNA profiling at the most active grain-filling stage was conducted in inferior spikelets under control(CK)and MD treatment.Of 521 known and 128 novel miRNAs,38 known and 9 novel miRNAs were differentially expressed between the CK and MD treatments.Target genes of differentially expressed miRNAs were involved in multiple developmental and signaling pathways associated with catalytic activity,carbohydrate metabolism,and other functions.Both miR1861 and miR397 were upregulated by MD,leading to a decrease in OsSBDCP1 and OsLAC,two negative regulators of SSIIIa activity and BR signaling,respectively.In contrast,miR1432 abundance was reduced by MD,resulting in upregulation of OsACOT and thus an elevated content of both ABA and IAA.These results suggest that both starch synthesis and phytohormone biosynthesis are regulated by differentially expressed miRNAs in inferior spikelets in response to MD treatment.Our results suggest the molecular mechanisms by which miRNAs regulate grain filling in inferior spikelets of rice under moderate soil drying,providing potential application in agriculture to increase rice yields by genetic approaches.