不同品质类型小麦籽粒游离氨基酸、蛋白质及其组分的动态变化

为了解小麦氮素营养代谢的动态变化规律,给优质专用小麦品种选育与应用提供科学依据,选用不同品质类型的四个小麦品种(系)济南17、PH82-2-2、PH97-4、PH97-5,在两种施氮水平和两种追肥时期下研究了籽粒游离氨基酸、蛋白质及其组分的动态变化.结果表明,籽粒发育初期游离氨基酸含量高,随籽粒发育其含量逐渐下降;蛋白质含量积累呈现"高-低-高"的变化趋势,品种之间差异显著,成熟期蛋白质含量高的品种一般有较高的游离氨基酸和蛋白质积累水平.不同品质类型小麦籽粒蛋白质组分含量变化动态基本一致,灌浆初期清蛋白含量较高,随籽粒发育逐渐下降,灌浆中后期下降趋缓;球蛋白总体含量水平较低,随籽粒发育缓慢下降,灌浆末期略有上升.醇溶蛋白在籽粒发育前期积累较少,花后14 d快速积累;谷蛋白在花后7 d已有一定的积累,随后其含量逐渐上升,强筋高蛋白品种有较高的醇溶蛋白和谷蛋白积累水平,在成熟籽粒蛋白质中谷蛋白和醇溶蛋白所占比例也高,弱筋低蛋白品种有相对较低的醇溶蛋白和谷蛋白积累,醇溶蛋白和谷蛋白所占比例也低.     

Evaluation of drought and heat stressed grain sorghum (Sorghum bicolor) for ethanol production

Sorghum is one of the most drought-tolerant grain crops and is used in biofuel production. Since sorghum is often exposed to drought and high temperature (heat) stress, this study investigated the effect of stress applied at different phenological stages of crop development on the glucose levels in grain and subsequent ethanol production. Short season sorghum hybrid DK-28E grown under controlled environment was exposed to drought and heat stress at five different stages of growth, namely: (1) pre-flowering (boot leaf emergence) to flowering, (2) flower to seed-set, (3) seed-set to early seed-fill, (4) early seed-fill to mid seed-fill, and (5) mid seed-fill to late seed-fill stage. Drought stress at any of the growth stages did not statistically affect either the glucose content or the ethanol production compared to the control (337 mg/g), although the ethanol yield increased up to 4.5% (352 mg/g) in flowering to seed-set stage. Heat stress, on the other hand, significantly reduced the glucose release and ethanol yield compared to the control (322 mg/g). Marginal ethanol yield reduction by 9% (293 mg/g) and 8.3% (295 mg/g) was seen in plants stressed during early seed-fill to mid seed-fill and mid seed-fill to late seed-fill stages, respectively. The results suggest that the phenological stage when sorghum is exposed to stress affects the ethanol yield. Overall, from the point of ethanol yield, it appears that grain sorghum cultivated in semi-arid regions where heat and drought stress are prevalent can be used for biofuel production. However, economic viability of ethanol production, especially of grain sorghum from high temperature stress needs to be investigated.     

灌浆前期高温对小黑麦籽粒淀粉积累及其合成相关酶活性的影响

为探讨灌浆前期高温对六倍体小黑麦籽粒淀粉积累及其相关合成酶活性的影响,以4个春性小黑麦品种（系）为材料,在开花后5~14 d于田间搭建温棚进行高温胁迫,对其籽粒淀粉积累动态、淀粉合成相关酶活性进行了研究。结果表明,开花后5~14 d高温胁迫导致小黑麦千粒重和容重均显著下降,其中,千粒重下降5.95%~20.47%,容重下降3.33%~9.49%;籽粒中的总淀粉、支链淀粉与直链淀粉含量均显著降低,依次下降6.84%~17.97%、7.70%~24.60%及2.27%~7.17%,直/支链淀粉比有所升高,但差异不显著。在高温胁迫期间,小黑麦籽粒中与淀粉合成相关的腺苷二磷酸葡萄糖焦磷酸化酶（AGPP）、可溶性淀粉合成酶（SSS）、淀粉分支酶（SBE）及淀粉去分支酶（DBE）活性均较对照有所上升,高温胁迫结束后,AGPP、SSS、DBE的活性均较对照有所下降。推测灌浆前期高温胁迫导致灌浆中后期小黑麦籽粒中淀粉合成相关酶活性的下降是籽粒中淀粉含量降低以及千粒重、容重下降的主要原因。     

Late defoliation and wheat yield: Little evidence of post-anthesis source limitation

The effect of source reduction on yield and protein content of bread wheat under well-watered and mild drought stress condition in a semi-arid climate was studied. Field experiments were conducted at the Tehran University research farm during 2003–2004 and 2004–2005 growing seasons. Mild drought stress was imposed when plants were at the second node stage by repeatedly withholding watering and re-irrigating when they showed symptoms of wilting or leaf rolling. Partial defoliations (all leaf lamina other than flag leaves were removed) were imposed at booting and anthesis; complete defoliation was imposed at anthesis (defoliation treatments were applied to all plants of each plot). Drought stress caused a significant increase in the remobilization of pre-anthesis reserves to the grain. Defoliation did not significantly affect remobilization. Grain yield and 1000-grain weight was reduced slightly by drought stress, but in most cases it was not significantly reduced by defoliation. Significant changes were not observed for grain protein content between defoliated and control plots. The results suggested that grain yield of the cultivar used under the condition tested is more controlled by sink than source strength.     

Physicochemical control of durum wheat grain filling and glutenin polymer assembly under different temperature regimes

Durum wheat is grown in the Mediterranean area where drought and high temperature frequently prevail and impact grain texture, composition and yield. The purpose of this work was to examine the effect of high temperature on grain development and final composition according to the timing of exposure. High temperature (up to 27.5 °C) was applied either during the linear grain filling or drying phases or during whole grain development. The dynamics of grain dry mass, water, glutenin polymers, and protein bodies during grain development were determined. Irrespective of high temperature timing, the arrest of grain filling was observed at 45.9% grain moisture content. At that point, starch granules included in endosperm cells reached their physical packing limit, limiting further deposits. HT applied before physiological maturity shortened the duration of grain filling and resulted in a significant increase in grain protein concentration and in the proportion of vitreous grain. Late formation of sodium dodecyl sulfate (SDS)-insoluble glutenin polymers below 32% grain moisture content was also favored. The ability of wheat storage protein to form a viscoelastic matrix embedding starch granules at the beginning of grain desiccation is proposed to be mandatory for gaining vitreous grains and a high proportion of SDS-insoluble glutenin polymers.     

Effect of heat stress on wheat proteins during kernel development in wheat near-isogenic lines differing at Glu-D1

Two near-isogenic lines of the wheat variety Lance having Glu-D1a (HMW-GS 2 + 12) and Glu-D1d (HMW-GS 5 + 10) were subjected to several regimes of heat stress. In 2001, the temperature regimes were (i) 20/16 (day/night, °C) from planting to maturity, (ii) 20/16 except for a 3-day heat treatment of 35/20, 25 days after anthesis and (iii) 20/16 until 25 DAA, after which plants were subjected to 40/25 until maturity. In 2002, treatments (i) and (iii) were the same while treatment (ii) used a temperature of 40/25 °C for 3 days at 25 DAA. Seed was collected at 3-day intervals starting from 16 days after anthesis and analyzed for protein composition by SE-HPLC. The line with the Glu-D1d allele showed an earlier polymerization of glutenin than its allelic counterpart and a higher molecular weight of glutenin at maturity, this being deduced from measurements of the percentage of unextractable polymeric protein. It is postulated that the timing and rate of glutenin polymerization, and the timing of high temperature application may be the key factors contributing to an explanation of the effect of heat stress on functionality.     

水氮互作对小麦籽粒蛋白质组分和品质的影响

为了给强筋小麦高产优质高效栽培提供理论依据,以高产强筋小麦品种济麦20为材料,研究了施氮量和灌溉量对小麦籽粒蛋白质组分和品质的影响.结果表明,施氮量由120 kg*ha-1(N1)增加至240 kg*ha-1(N2),籽粒蛋白质含量、清蛋白和球蛋白含量增加,(醇溶蛋白+麦谷蛋白含量)/(清蛋白+球蛋白含量)比值(谷醇/清球比值)降低,面团稳定时间缩短.同一施氮量条件下,由不灌水(W0)到灌2水(W1),籽粒蛋白质含量增加,谷醇/清球比值降低,面团稳定时间缩短;由灌2水(W1)到灌3水(W2)和5水(W3),籽粒蛋白质含量降低,蛋白质组分中清蛋白和球蛋白含量升高,醇溶蛋白和麦谷蛋白含量降低,谷醇/清球比值降低,面团稳定时间亦缩短.以上结果表明,在本试验条件下,增加施氮量和灌水量导致面团稳定时间缩短,原因是清蛋白和球蛋白占总蛋白含量的比例,即谷醇/清球比值降低.     

吐丝后不同阶段干旱胁迫对糯玉米子粒产量和淀粉品质的影响

以苏玉糯5号和渝糯7号为材料,试验对照和干旱处理下土壤相对含水量分别为75％和55％,干旱胁迫时间分别为子粒建成期和灌浆充实期,研究结实期不同阶段(子粒建成期和灌浆充实期)干旱胁迫对糯玉米子粒产量和淀粉品质的影响.结果表明,结实期干旱胁迫显著降低子粒产量,且降幅子粒建成期大于灌浆充实期.干旱胁迫下淀粉含量显著降低,总蛋...     

Thermal Resistance of Common Rice Maintainer and Restorer Lines to High Temperature During Flowering and Early Grain Filling Stages

Fifteen common rice maintainer lines and 26 high-yielding restorer lines were used to evaluate their thermal resistance and fertility during flowering and early grain filling stages.The rice plants were subjected to high temperature stress(39-43 ℃) for 1-15 d from main stem flowering.Based on the heat stress index,they were divided into thermal resistant lines,semi-thermal resistant lines,semi-thermal sensitive lines and thermal sensitive lines.Therefore,the maintainer lines K22B,Bobai B and V20B belonged to thermal resistant lines,whereas II-32B,Zhongzhe B and Zhong 9B belonged to thermal sensitive lines.For rice restorer lines,Minghui 63 had the highest thermal resistance,followed by R207,P32,P929,and the lowest thermal resistant lines P62-2-2,R8006 and P51.The correlation analysis indicated that the heat stress index was significantly correlated with seed-setting rate and abortive grain rate under heat stress,but not under natural conditions.This indicated that heat stress occurred during flowering and early grain filling stages mainly decreased the seedsetting rate and significantly increased the abortive grain rate in both rice maintainer and restorer lines.     

Yield response of rice (Oryza sativa L.) genotypes to drought under rainfed lowland: 3. Plant factors contributing to drought resistance

A series of experiments were conducted in drought-prone northeast Thailand to examine the magnitude of yield responses of diverse genotypes to drought stress environments and to identify traits that may confer drought resistance to rainfed lowland rice. One hundred and twenty eight genotypes were grown under non-stress and four different types of drought stress conditions.Under severe drought conditions, the maintenance of PWP of genotypes played a significant role in determining final grain yield. Because of their smaller plant size (lower total dry matter at anthesis) genotypes that extracted less soil water during the early stages of the drought period, tended to maintain higher PWP and had a higher fertile panicle percentage, filled grain percentage and final grain yield than other genotypes. PWP was correlated with delay in flowering (r=−0.387) indicating that the latter could be used as a measure of water potential under stress. Genotypes with well-developed root systems extracted water too rapidly and experienced severe water stress at flowering. RPR which showed smaller coefficient of variation was more useful than root mass density in identifying genotypes with large root system.Under less severe and prolonged drought conditions, genotypes that could achieve higher plant dry matter at anthesis were desirable. They had less delay in flowering, higher grain yield and higher drought response index, indicating the importance of ability to grow during the prolonged stress period.Other shoot characters (osmotic potential, leaf temperature, leaf rolling, leaf death) had little effect on grain yield under different drought conditions. This was associated with a lack of genetic variation and difficulty in estimating trait values precisely.Under mild stress conditions (yield loss less than 50%), there was no significant relationship between the measured drought characters and grain yield. Under these mild drought conditions, yield is determined more by yield potential and phenotype than by drought resistant mechanisms per se.     

干旱条件下土壤盐分对大豆生长及光合作用的影响

以盆栽大豆为材料,设置不同程度的土壤盐分(NaCl)、干旱及旱盐组合处理,然后测定各处理大豆植株的株高、生物量、光合作用指标以及植株的水分状况和Na~+、K~+含量,探索干旱条件下土壤盐分对大豆生长的影响及可能机制。结果表明:干旱和盐胁迫均可导致大豆叶片的净光合速率降低和生长量的减少,干旱还导致光合机构的严重损伤。但是,干旱和适量土壤盐分(100~150 mmol·L~(-1)NaCl)组合处理的大豆植株,其生长量、净光合速率和PSⅡ最大光化学效率都显著高于单一干旱处理。同时,旱盐组合处理的大豆叶片RWC、Na~+含量也高于单一干旱处理,水势和渗透势低于后者,且叶片Na~+含量与其渗透势降低显著相关。综合分析表明,在干旱条件下,土壤适量NaCl的存在使大豆能够吸收和积累更多Na~+等盐离子作为渗透调节物质,来降低渗透势、提高吸水能力,以改善植株的水分状况和光合性能,保持植株较高的生长速率,即土壤中适量盐分(NaCl)的存在可减轻干旱对大豆的负效应。     

Adapting wheat in Europe for climate change

Increasing cereal yield is needed to meet the projected increased demand for world food supply of about 70% by 2050. Sirius, a process-based model for wheat, was used to estimate yield potential for wheat ideotypes optimized for future climatic projections for ten wheat growing areas of Europe. It was predicted that the detrimental effect of drought stress on yield would be decreased due to enhanced tailoring of phenology to future weather patterns, and due to genetic improvements in the response of photosynthesis and green leaf duration to water shortage. Yield advances could be made through extending maturation and thereby improve resource capture and partitioning. However the model predicted an increase in frequency of heat stress at meiosis and anthesis. Controlled environment experiments quantify the effects of heat and drought at booting and flowering on grain numbers and potential grain size. A current adaptation of wheat to areas of Europe with hotter and drier summers is a quicker maturation which helps to escape from excessive stress, but results in lower yields. To increase yield potential and to respond to climate change, increased tolerance to heat and drought stress should remain priorities for the genetic improvement of wheat.     

灌水模式对春小麦光合性能和干物质生产的影响

为探寻宁夏引黄灌区春小麦节水栽培的适宜灌水模式,比较分析了6个不同灌水处理(W1:二棱水+开花水;W2:二棱水;W3:拔节水;W4:二棱水+孕穗水;W5:二棱水+拔节水+孕穗水+开花水+灌浆水;W6:二棱水+拔节水+孕穗水+开花水)下春小麦光合性能、物质积累与转运、产量性状及水分利用的差异。结果表明,灌二棱水、孕穗水或开花水的W1、W4、W5、W6处理明显增加春小麦花后旗叶光合速率、蒸腾速率和气孔导度,灌二棱水和生育后期少灌水的W1、W2、W4处理明显增加花后旗叶胞间CO2浓度。灌水次数减少会显著增加旗叶SPAD值,并降低叶面积系数,其中以缺少二棱水的W3处理最为明显。灌有二棱水和孕穗水的W4处理明显降低抽穗至开花旗叶SPAD值,增加开花至灌浆旗叶SPAD值和抽穗至开花叶面积系数,灌开花水明显增加开花至灌浆旗叶面积。灌二棱水、拔节水、孕穗水、开花水明显促进了干物质积累,灌水次数增加明显增加了叶干物质积累量,并降低茎鞘干物质比例;缺少二棱水的W3处理最不利于叶、穗干物质积累,但提高了开花至成熟叶干物质比例;灌浆水对各器官干物质积累影响不明显。二棱期至开花期缺少灌水的W1、W3、W2处理促进了开花前贮藏同化物在花后向籽粒的转运。籽粒产量随灌水次数的增加而增加;缺少二棱水明显降低穗粒数、收获指数,但增加千粒重;增加灌水次数降低了灌水利用效率,且以灌二棱水影响最为明显。综上所述,缺少二棱水对春小麦产量影响最明显,孕穗水、开花水影响次之,灌浆水影响不明显。     

Environmental factors associated with glutenin polymer assembly during grain maturation

This study aims to understand the environmental factors, focusing on rain and fungal infection, affecting the assembly of glutenin polymers during grain maturation. Spring wheat was grown in the field and grains were sampled from 50% grain moisture until maturity. Grain moisture content, protein content, size of glutenin polymers, the presence of proteases, and the amount of DNA from common wheat pathogenic fungi were analysed.Rain influenced the rate of grain desiccation that occurred parallel to the rate of glutenin polymer assembly. Rapid desiccation contributed to faster glutenin polymer assembly than gradual desiccation. Severe reduction in the glutenin polymer size coincided with increased grain moisture due to rain. Furthermore, increased fungal DNA followed by presence of gluten-degrading proteases was observed in the grain after humid conditions. The presence of gluten-degrading proteases was presumably involved in reducing the size of glutenin polymers in grain.Our study gave new insight into how environmental conditions could be associated with the assembly of glutenin polymers during grain maturation. The results suggest that rain and/or fungal proteases play an important role in reducing the molecular size of glutenin polymers.     

Yield and yield-attributing traits of rice (Oryza sativa L.) under lowland drought and suitability of early vigor as a selection criterion

Fifteen experiments testing seven sets of genotypes under irrigated non-stress and drought stress situations were conducted from 2004 to 2006 with the objective to study the effect of different severity levels of drought on tolerant and susceptible advanced breeding lines, current varieties, and traditional and improved donors, and to study the effect of selection for high early vigor on yield. With the onset of drought, a yield reduction due to a decrease in biomass was observed in both susceptible and tolerant lines. However, after an initial reduction in yield, tolerant lines were able to sustain a yield loss more effectively than susceptible lines by better maintenance of biomass and higher harvest index. Under intermittent drought stress, genotypes with the ability to maintain higher biomass, reflected in terms of high early vigor recorded 3 wk after transplanting, recorded higher grain yield than genotypes with low early vigor. Lines with high early vigor yielded higher under both irrigated non-stress and intermittent drought stress situations. Under intermittent stress, plants with high vigor, low vigor, or random plants with high or low vigor coming from tolerant × susceptible crosses yielded higher than did the plants with the same vigor coming from susceptible × susceptible crosses. Under both irrigated non-stress and intermittent drought stress situations, early vigor showed significant genetic correlation with grain yield and could be included as an efficient selection criterion to improve yield in the lowlands.     

灌浆期不同阶段高温胁迫对春小麦籽粒生长的影响

灌浆期短暂高温胁迫严重影响小麦产量。为了明确灌浆期不同阶段短暂高温胁迫对春小麦籽粒生长的影响,以强筋春小麦品种龙麦26和龙麦30为试材,在人工气候室(25℃/15℃)精确控温和人工温室形成绝对高温胁迫的条件下,分析了灌浆期不同阶段5d短暂高温胁迫对粒重的影响。结果表明,在灌浆期不同阶段短暂高温胁迫处理中,两个小麦品种均表现为:前期高温胁迫对粒重影响最大,达极显著水平(P0.01);随着高温胁迫处理时期的后移,粒重降低幅度逐渐减小;后期(花后25d)高温胁迫对粒重影响不显著。灌浆期缩短是导致粒重降低的主要原因。     

不同生育时期干旱胁迫对水稻产量与碾米品质的影响

以粳稻品种农大3号为材料,利用盆栽的方式人工控制土壤水分(水势),研究了不同生育时期干旱胁迫对水稻产量及碾米品质的影响。各时期干旱均导致产量下降,其敏感程度依次为：孕穗中期（抽穗前11~20 d）>孕穗后期（抽穗前1~10 d）>分蘖中期（移栽后11~20 d）>孕穗前期（抽穗前21~30 d）>分蘖初期（移栽后1~10 d）>分蘖后期（移栽后21~30 d）>乳熟期（抽穗后31~40 d）>抽穗开花期（抽穗后1~10 d）>灌浆期（抽穗后21~30 d）>蜡熟期（抽穗后41~50 d）。抽穗前干旱胁迫的敏感时期主要是孕穗中期、孕穗后期和分蘖中期。抽穗后,抽穗到乳熟期均较蜡熟期对干旱敏感,但对土壤水分变化较抽穗前迟钝。抽穗前干旱胁迫对糙米率和精米率影响最大的时期为孕穗中期,其次为孕穗后期。灌浆期到蜡熟期干旱胁迫,糙米率和精米率明显下降,抽穗后11~40 d干旱胁迫对整精米率的影响最大,其次为抽穗前1~20 d,其他时期干旱虽对整精米率有一定影响,但影响较小。     

Heat and drought stress on durum wheat: Responses of genotypes,yield, and quality parameters

Heat and/or drought stress during cultivation are likely to affect the processing quality of durum wheat (Triticum turgidum L. ssp. durum). This work examined the effects of drought and heat stress conditions on grain yield and quality parameters of nine durum wheat varieties, grown during two years (2008–09 and 2009–10). Generally, G and E showed main effects on all the parameters whereas the effects of G × E were relatively small. More precipitation in Y09–10 may account for the large differences in parameters observed between crop cycles (Y08–09 and Y09–10). Combined results of the two crop cycles showed that flour protein content (FP) and SDS sedimentation volume (SDSS) increased under both stress conditions, but not significantly. In contrast the gluten strength-related parameters lactic acid retention capacity (LARC) and mixograph peak time (MPT) increased and decreased significantly under drought and heat stress, respectively. Drought and heat stress drastically reduced grain yield (Y) but significantly enhanced flour yellowness (FY). LARC and the swelling index of glutenin (SIG) could be alternative tests to screen for gluten strength. Genotypes and qualtiy parameters performed differently to drought and heat stress, which justifies screening durum wheat for both yield and quality traits under these two abiotic stress conditions.     

外源水杨酸对高温胁迫下小麦籽粒灌浆特性的影响

为探究外源水杨酸(SA)对高温胁迫下小麦产量形成特性的影响,以扬麦18为试验材料,于2018-2020年进行盆栽试验,分别在小麦抽穗期(SAHG)、开花期(SAAG)、花后5 d(SA5G)以及花后10 d(SA10G)对其喷施水杨酸(SA),在灌浆期(花后15～19 d)采用被动式模拟增温方法对其进行高温胁迫,分析不同处理对小麦产量及其相关指标的影响。结果表明,与常温对照(NN)相比,灌浆期高温胁迫(NG)显著降低了小麦的产量和千粒重,不同时期喷施SA显著降低了高温胁迫下产量和千粒重的下降幅度,不同处理缓解效果表现为SAAG>SA5G>SA10G>SAHG。外源SA处理缓解了高温胁迫对小麦不同穗位强、弱势粒粒重的损害,对上部小穗及弱势粒粒重的影响较为显著,从而缓解了高温胁迫下小麦产量的降低。喷施SA显著降低了灌浆期高温胁迫下籽粒蔗糖含量、可溶性糖含量、蔗糖合成酶(SS)活性的降低幅度,提高了籽粒中吲哚乙酸(IAA)、脱落酸(ABA)、细胞分裂素(CTK)的含量。推测喷施SA通过调控小麦籽粒内源激素水平提高籽粒糖类物质含量,进而缓解高温胁迫对产量影响。     

Enclosed stigma contributes to higher spikelet fertility for rice(Oryza sativa L.) subjected to heat stress

With global warming, rice plants may be subjected to heat stress more regularly during the heatsensitive flowering stage, causing spikelet sterility and grain yield loss.Stigma exsertion is considered to increase pollen reception and promote female reproductive success.The aim of this study was to investigate the role of stigma exsertion on spikelet fertility at high temperatures.Five rice cultivars(Liangyoupeijiu, Shanyou 63, Huanghuazhan, Nagina 22, and IR64) with differing degrees of stigma exsertion were cultivated and exposed to high temperature at anthesis.Heat-tolerant cultivars did not always show a high percentage of spikelets with exserted stigmas, and vice versa.Irrespective of the presence of more pollen grains on exserted stigmas, spikelets with exserted stigmas did not show greater spikelet fertility than spikelets with fewer exserted stigmas or hidden stigmas under heat stress.GA3 application augmented the percentage of spikelets with exserted stigmas;however, it did not increase spikelet fertility under heat stress.Spikelet fertility of whole panicles was negatively correlated with the percentage of spikelets with exserted stigmas, but positively with that with hidden stigmas.Viability of the hidden stigmas was less reduced than that of exserted stigmas under heat stress, suggesting that hidden stigmas have an advantage in maintaining viability.Heat stress delayed anther dehiscence and reduced the viabilities of both exserted stigmas and pollens, thereby causing low spikelet fertility.Together, these results suggest that high spikelet fertility does not depend on stigma exsertion and that enclosed stigma generally contributes to higher spikelet fertility and heat tolerance under high-temperature conditions during flowering in rice.