Temporal Dynamics of Antioxidant Defence System in Relation to Polyamine Catabolism in Rice under Direct-Seeded and Transplanted Conditions

Six rice cultivars viz. PR120, PR116, Feng Ai Zan, PR115, PAU201 and Punjab Mehak 1 under the direct-seeded and transplanted conditions were used to investigate the involvement of antioxidative defence system in relation to polyamine catabolism in temporal regulation of developing grains. Activities of ascorbate peroxidase(APx), guaiacol peroxidase(GPx), catalase(CAT), superoxide dismutase(SOD), polyamine oxidases(PAO) and contents of ascorbate, α-tocopherol, proline and polyamines increased gradually until mid-milky stage and then declined towards maturity stage under both planting conditions. The transplanted condition led to higher activities of antioxidative enzymes(APx, GPx and CAT) and contents of ascorbate, α-tocopherol and proline whereas the direct-seeded condition had elevated levels of PAO and SOD activities and contents of polyamines, lipid peroxide and hydrogen peroxide. Cultivars Feng Ai Zan and PR120 exhibited superior tolerance over other cultivars by accumulating higher contents of ascorbate, α-tocopherol and proline with increasing level of PAO and SOD activities under the direct-seeded condition. However, under the transplanted condition PR116 and PAU201 showed higher activities of antioxidative enzymes with decreasing content of lipid peroxide. Therefore, we concluded that under the direct-seeded condition, enhancements of polyamines content and PAO activity enabled rice cultivars more tolerant to oxidative stress, while under the transplanted condition, antioxidative defence with decreasing of lipid peroxide content was closely associated with the protection of grains by maintaining membrane integrity during rice grain filling. The results indicated that temporal dynamics of H2O2 metabolic machinery was strongly up-regulated especially at the mid-milky stage.     

灌浆结实期高温对水稻籽粒蔗糖及降解酶活性的影响

选用水稻品种越光和笹锦为材料,在开花后设置自然温度和高温两种处理,研究了不同温度处理下籽粒蔗糖、果糖和葡萄糖含量的动态变化以及蔗糖合酶、液泡型转化酶和细胞壁结合型转化酶活性的差异。高温处理下,蔗糖合酶活性值大于转化酶活性值,且与淀粉积累速率呈显著正相关,表明蔗糖合酶在蔗糖的分解和淀粉的合成过程中起着重要作用;高温下两品种籽粒的蔗糖含量明显增加,但分解的葡萄糖和果糖含量并未相应增加,表明高温有利于籽粒中蔗糖的积累,而不利于籽粒中蔗糖的分解。高温下蔗糖合酶、液泡型转化酶和细胞壁结合型转化酶活性明显下降,表明蔗糖分解速率的下降与蔗糖合酶和转化酶活性下降有关。     

Effect of High Temperature on Sucrose Content and Sucrose Cleaving Enzyme Activity in Rice Grain During the Filling Stage

Global warming is due to continuous increase in greenhouse effect caused by human productive activities. Much attention has been paid to the influence brought by the increase of temperature on crop growth. Many researches in China and other countries are …     

粤西1号芒果成熟阶段的蔗糖代谢

以粤西1号为试材研究了芒果果实成熟阶段淀粉、葡萄糖、果糖和蔗糖的含量以及蔗糖代谢相关酶-酸性转化酶(AI)、中性转化酶(NI)、蔗糖合成酶(SS)和蔗糖磷酸合成酶(SPS)的活性变化.结果表明:采收前淀粉含量逐渐减少,采收后快速减少;葡萄糖含量采收后呈先增后减的趋势,果糖含量采收前缓慢增加,采收后开始迅速增加,蔗糖变化与果糖变化趋势一致,完熟时含量最多的糖为蔗糖(62.55 mg,g),果糖次之(44.45 mg/g),葡萄糖最少(2.87mg/g).AI、NI、SS、SPS在采收前活性差别不大,在后熟阶段,AI、SS、SPS活性显著增加,且SPS活性达到最高值.粤西1号糖代谢以积累蔗糖为主,2种酶SPS和SS起关键作用.     

不同气孔密度水稻剑叶光合特性及蔗糖代谢研究

以叶片不同气孔密度(高、中和低气孔密度)水稻品系为试验材料,在齐穗期研究水稻剑叶光合特性、糖分含量和蔗糖代谢酶活性。结果表明,水稻叶片中蔗糖含量、蔗糖磷酸合成酶活性与气孔密度呈极显著正相关;蔗糖合成酶活性与气孔密度呈显著正相关。高气孔密度水稻叶片Rubisco活性、净光合速率、蔗糖含量、蔗糖磷酸合成酶和蔗糖合成酶活性显著高于低气孔密度水稻;不同叶片气孔密度与中性转化酶呈显著负相关。研究认为,气孔密度增加提高了水稻叶片光合效率,促进了蔗糖积累及蔗糖代谢酶活性提高,有利于更多光合产物——蔗糖向籽粒转运。     

Carbohydrate metabolism during tuber initiation in potato: A transient surge in invertase activity marks the stolon to tuber transition

Summary The relationship between carbohydrate metabolism and tuber initiation in potato was determined by monitoring changes in the amount of starch and sugars along with the activities of sugar metabolizing enzymes upon transfer of plants to tuber-inducing conditions (TI; short days, cold nights) from non-inducing conditions (NTI: long days, warm nights). Switch to TI conditions caused an immediate slow-down in plant growth and triggered swelling of stolon tips, which went on to develop into tubers. Leaves of plants moved to TI conditions accumulated less starch and sugar while their stolon tips showed a sudden upsurge in starch content and a sharp decline in sugars even before any tip swelling was detectable. These changes were paralleled by a transient surge in the activity of cell wall invertase (74%) and soluble invertase (30%) in stolon tips of plants transferred to TI conditions in two unrelated cultivars under different experimental conditions. As the surge in invertase activity faded, it was replaced by a substantial increase in sucrose synthase activity as the tuber enlargement proceeded. The transient increase in invertase activity just prior to tuber initiation appears to mark a turning point in the transition of stolon tip to tuber.     

Selenium Alleviates Carbohydrate Metabolism and Nutrient Composition in Arsenic Stressed Rice Plants

This study reported the influence of selenium (Se) on carbohydrate composition and some related enzymes and nutrient compositions of arsenic (As) stressed rice plants. Rice plants of cultivar PR126 were grown on soil amended with As in a range of 25–100 μmol/kg with and without 0.5 or 1.0 mg/kg Se. Total soluble sugars (TSS) and reducing sugars (RS) increased in leaves of As stressed plants at the tillering and grain filling stages whereas sucrose and starch contents showed the reverse trend. Se supplementation to As stressed plants further increased TSS and RS, and enhanced sucrose phosphate synthase activity in rice leaves, thus improving sucrose content and the tolerance to As stress of the plants. Se alone or in combination with As resulted in lower As accumulation in rice husk and grains, and the highest reduction was observed in Se applied at 1.0 mg/kg compared to the corresponding As treatments alone. As may limit the accumulations of Na, Mg, K, Ca, Fe, Zn and Mn in rice grains, which are essential for humans. Binary application of different combinations of As and Se protected the plants against As and increased the mineral content in rice grains. Addition of Se in As treated soil significantly alleviated As stress by enhancing grain yields compared to the corresponding As treatment. It is concluded that Se induced amelioration of the toxic impact of As in rice either by modulating carbohydrate composition and/or nutrient uptake is one of the mechanisms to alleviate As stress in plants.     

外源二乙基二硫代氨基甲酸钠 对花期淹水大豆碳代谢的影响

      为了研究二乙基二硫代氨基甲酸钠（DDTC）对大豆碳水化合物代谢的调节作用,为大豆抗渍涝栽培提供技术支持,本研究以渍涝敏感型大豆品种徐豆18和耐渍涝品种南农1138-2为材料,采用盆栽的方式,花期淹水前喷施DDTC,调查大豆叶片光合特性、蔗糖代谢酶活性及叶片、根系和根瘤中碳水化合物含量。结果表明：淹水胁迫降低了植株地上部、根系和根瘤干物重,叶片净光合速率（Pn）、气孔导度（Gs）、胞间二氧化碳浓度（Ci）、蒸腾速率（Tr）,根系中淀粉含量、根瘤中蔗糖和淀粉含量。其中,南农1138-2植株干物重,叶片光合气体交换参数以及根系中淀粉含量比徐豆18下降幅度小,而两品种间根瘤中蔗糖和淀粉含量下降幅度基本一致。与之不同,叶片中蔗糖磷酸合成酶（SPS）、蔗糖合成酶（SS）（合成方向）活性、SS（分解方向）、酸性转化酶（AI）活性,叶片可溶性糖、蔗糖和淀粉含量,根系中可溶性糖、蔗糖含量增加。其中,南农1138-2叶片中蔗糖合成酶（SS）（分解方向）和酸性转化酶（AI）活性、可溶性糖、蔗糖和淀粉含量增幅较小,而叶片中蔗糖磷酸合成酶（SPS）、SS（合成方向）活性增幅较大。DDTC处理提高了淹水胁迫下两品种植株干物重,缓解了淹水胁迫对大豆植株生长的抑制;提高了叶片光合气体交换参数,叶片中SPS和SS（合成方向）活性,并降低了SS（分解方向）和AI活性,促进了淹水胁迫下叶片中蔗糖向根系和根瘤的转运;DDTC处理也增加了根系中可溶性糖、蔗糖和淀粉含量,根瘤中蔗糖和淀粉含量,表明DDTC处理在调控淹水胁迫下大豆植株生长和碳代谢生理过程中起着重要的作用。     

Impeded Carbohydrate Metabolism in Rice Plants under Submergence Stress

The detrimental effects of submergence on physiological performances of some rice varieties with special references to carbohydrate metabolisms and their allied enzymes during post-flowering stages have been documented and clarified in the present investigation. It was found that photosynthetic rate and concomitant translocation of sugars into the panicles were both related to the yield. The detrimental effects of the complete submergence were recorded in generation of sucrose, starch, sucrose phosphate synthase and phosphorylase activity in the developing panicles of the plants as compared to those under normal or control (i.e. non-submerged) condition. The accumulation of starch was significantly lower in plants under submergence and that was correlated with ADP-glucose pyrophosphorylase activity. Photosynthetic rate was most affected under submergence in varying days of post-flowering and was also related to the down regulation of Ribulose bisphosphate carboxylase activity. However, under normal or control condition, there recorded a steady maintenance of photosynthetic rate at the post-flowering stages and significantly higher values of Ribulose bisphosphate carboxylase activity. Still, photosynthetic rate of the plants under both control and submerged conditions had hardly any significant correlation with sugar accumulation and other enzymes of carbohydrate metabolism like invertase with grain yield. Finally, plants under submergence suffered significant loss of yield by poor grain filling which was related to impeded carbohydrate metabolism in the tissues. It is evident that loss of yield under submergence is attributed both by lower sink size or sink capacity (number of panicles, in this case) as well as subdued carbohydrate metabolism in plants and its subsequent partitioning into the grains.     

高、低糖甘蔗品种苗期糖分含量及代谢相关酶活性分析

以高糖(GT35)和低糖(B8)甘蔗品种苗期不同部位的叶和茎为材料,采用HPLC技术测定蔗糖、葡萄糖和果糖含量,分析2个甘蔗品种叶和茎中蔗糖代谢相关酶活性与糖分含量的相关性和差异性。结果表明:2个甘蔗品种苗期叶中蔗糖含量与SPS和SS-s酶活性呈显著正相关,茎中己糖含量与NI酶活性呈显著正相关,茎中蔗糖含量与SPS酶活性呈显著正相关,而与SS-c、SAI和CIN酶活性呈显著负相关。GT35茎中蔗糖含量显著高于B8,叶中蔗糖含量显著低于B8。GT35茎中SPS和叶中SS-s酶活性均显著高于B8,茎中SS-s和SS-c酶活性低于B8,其中只有幼茎中SS-c酶活性与B8差异不显著。说明苗期叶中高SS-s酶活性,茎中高SPS酶活性,低SS-s和SS-c酶活性,可能是调节高糖甘蔗品种苗期茎中蔗糖积累的重要因素。     

橡胶树‘热研7-33-97’和‘PR107’胶乳和树皮中蔗糖代谢相关的酶活与基因表达研究

对2个产胶水平差异明显的橡胶树品种（‘热研7-33-97’和‘PR107’）产胶旺季时2个与产胶直接相关的碳库（胶乳和茎干树皮）中蔗糖代谢相关的酶活、基因表达以及胶乳生理参数、树皮非结构性碳水化合物（NSC）含量进行了比较研究。结果表明：‘PR107’干胶产量的季节性变动明显大于‘热研7-33-97’,但均在9月底出现一个产胶高峰;2个品种胶乳的总固形物、无机磷和蔗糖含量差异不明显,但‘热研7-33-97’的干胶含量显著高于‘PR107’;树皮NSC的主要成分为可溶性糖（占80%以上）,但品种间的NSC及其相关组分（淀粉、可溶性糖和还原性糖）的差异均不显著;胶乳中,蔗糖合成酶（Sus）主要催化蔗糖合成,品种间的中碱性转化酶（NIN）和Sus酶活差异均不显著;树皮中,‘热研7-33-97’的NIN酶活显著大于‘PR107’,而其他蔗糖代谢相关酶活品种间的差异不显著;在胶乳中,HbSWEET10a基因的表达量高,并且‘热研7-33-97’>‘PR107’,还是唯一在品种间表达显著差异的蔗糖代谢相关基因;在树皮中,‘PR107’的HbSus3和细胞壁转化酶基因HbCIN2的表达显著高于‘热研7-33-97’,HbSUT5显著低于‘热研7-33-97’,而其他4个蔗糖代谢相关基因的表达在品种间差异不显著。本研究结果为深入探究蔗糖代谢调控与胶乳再生的互作奠定了基础。     

土壤水分对水稻光合速率与物质运转的影响

 以6个中熟水、陆稻品种为材料，研究了在不同土壤水分状况下水稻光合速率与物质运转的特点。叶片光合速率随土水势的下降而减小，抗旱性品种减小的幅度小于抗旱性较差的品种。长期低土壤水分处理后，群体光合能力的下降主要是由于叶面积的减小。低土壤水分处理导致叶片中蔗糖和总可溶性糖含[CM(45*2]量的增加，淀粉含量的减少。淀粉/可溶性糖、淀粉/蔗糖与相对产量(处理/对照)呈极显著正相关(r=0.9854、 r=0.9412)。低土壤水分处理下，抽穗期茎鞘中可用性糖(可溶性糖+淀粉)累积量减少，抽穗后运转率增加，抗旱性品种的相对累积量(处理/对照)高于抗旱性较差的品种，但运转率的大小与品种的抗旱性无明显联系。     

马铃薯不同品种淀粉积累生理基础研究——淀粉积累与磷酸化酶、蔗糖转化酶的关系

本试验以高淀粉品种晋薯２号、中淀粉品种内薯３号、低淀粉品种紫花白为材料，对不同生育时期块茎淀粉含量、淀粉积累速度与淀粉磷酸化酶、蔗糖转化酶活性的关系进行了研究，其结果表明：全生育期３个品种淀粉磷酸化酶活性均呈单峰曲线变化；不同生育时期，各品种淀粉含量与淀粉磷酸化酶活性无明显规律性；不同生育时期，各品种淀粉积累速度与淀粉磷酸化酶活性呈正相关。淀粉含量和温度是影响淀粉磷酸化酶活性的两个重要因素，而温度又起主导作用。全生育期，３个品种的蔗糖转化酶均呈苗期为最高，中期显著下降，后期略有回升的变化趋势；不同生育时期，各品种淀粉含量与蔗糖转化酶活性均呈显著负相关；不同生育时期，各品种淀粉积累速度与蔗糖转化酶活性亦呈负相关。因此，蔗糖转化酶活性可作为马铃薯高淀粉育种早期选择的重要生理指标。     

ABA对水分胁迫下水稻籽粒灌浆的调节

 利用水平分根系统研究了水稻抽穗后水分胁迫条件下ABA对水稻品种汕优63叶片光合特性、糖代谢和籽粒灌浆过程的调节。当一半根系受干旱胁迫时，叶片的水势和相对含量变化很小，但叶片和籽粒内ABA的含量显著增加。整个根系受干旱胁迫时，叶片的水势和相对含水量随着胁迫的增加而显著地降低，ABA含量则显著增加。半干和全干处理的水稻叶片的光合速率和RuBP羧化酶／氧化酶（Rubisco）的活性显著下降。叶片内蔗糖和淀粉的积累和代谢与叶片内ABA的含量和水分状况存在密切的关系，低浓度的ABA和高的叶水势与高的蔗糖积累和蔗糖磷酸合成酶活性相一致，但当叶片ABA的浓度增加时，即使叶片的水势与对照相似，叶片的蔗糖和淀粉积累以及蔗糖磷酸合成酶的活性则显著地下降，叶片的α－和β－淀粉酶活性似与ABA和水分状况关系不密切。ABA对籽粒灌浆有显著的抑制作用，ABA对籽粒灌浆的抑制作用至少部分与其对催化水稻籽粒内蔗糖代谢和淀粉积累的关键酶蔗糖合成酶的抑制有关。另外，干旱处理对叶片光合速率和蔗糖与淀粉积累的抑制程度显著高于半干旱处理，说明干旱本身也对这些过程有直接的效应。根据这些结果，作者认为，当水稻遇到干旱胁迫时，ABA对其光合作用、糖代谢以及籽粒的发育过程都有极其显著的调节作用。还就ABA对这些生理过程调节的可能性进行了讨论     

氮肥对水稻胚乳淀粉品质、相关酶活性及基因表达量的影响

【目的】旨在为阐明灌浆成熟期氮素营养对水稻淀粉品质影响机理以及建立优质高产水稻栽培技术提供理论依据。【方法】选用4个籽粒直链淀粉含量差异显著的粳稻品种,通过盆栽试验研究氮素营养对稻米淀粉组分和蛋白质含量及稻米蛋白质水解对淀粉黏滞特性的影响,并分析氮素营养对灌浆过程中籽粒蔗糖合酶(SuSy)、蔗糖磷酸合酶(SPS)、蔗糖酸性转化酶(AI)活性及OsGBSSⅠ、OsISAⅠ、OsSBEⅠ、OsSBEⅡ基因转录表达量的影响。【结果】结果表明,籽粒支链淀粉含量对氮素营养很敏感,灌浆成熟期氮素营养能改变籽粒淀粉组分含量;施氮条件下,高直链淀粉含量品种蒸煮食味品质下降更加明显,去除稻米蛋白质可明显提高稻米黏滞特性,蛋白质对淀粉黏滞特性的影响很大;增加灌浆成熟期氮素营养能显著或极显著提高籽粒SuSy和AI活性,显著抑制籽粒SPS活性;灌浆成熟期氮素营养能改变灌浆不同时期籽粒OsGBSSⅠ、OsISAⅠ和OsSBEⅡ基因转录表达量,以致灌浆过程中这些基因的转录表达量变化动态发生改变,但OsSBEⅠ基因转录表达量不因氮素营养而发生改变;受氮素营养的影响,灌浆起始期籽粒OsGBSSⅠ基因表达量明显上调,而灌浆中后期明显下调;氮素营养明显抑制灌浆成熟期籽粒OsISAⅠ基因和灌浆中后期的OsSBEⅠ基因转录表达,显著提高灌浆前期和中期的籽粒OsSBEⅠ基因转录表达量;氮素营养能抑制灌浆起始期籽粒OsSBEⅡ基因的转录表达,而提高灌浆中后期的基因转录表达。【结论】灌浆成熟期氮素营养除了通过蛋白质含量对淀粉品质产生影响外,还通过调控淀粉合成相关的酶活性和基因表达量等生理环节对淀粉含量和精细结构起作用,最终改变稻米黏滞特性和食味品质。     

脱落酸对水稻愈伤组织糖类代谢及相关酶活性的影响

以水稻愈伤组织为材料,研究了外源脱落酸（ABA）对糖类代谢及相关酶活性的影响。 ABA处理可促进淀粉和非还原糖在愈伤组织中的积累,以10~12 mg/L ABA作用效果最为显著。愈伤组织在处理培养期间,淀粉和非还原糖含量的动态变化受ABA影响明显。ABA能降低还原糖水平以及果糖与葡萄糖的比例,抑制磷酸葡萄糖同分异构酶（PGI）和转化酶的活性,增加磷酸葡萄糖变构酶（PGM）和淀粉磷酸化酶活性。试验结果表明,ABA对糖类代谢和相关酶活性的影响效果在处理的1~7 d最为明显。推测非还原糖含量的升高可能是ABA促进其积累和抑制转化酶活性所致,淀粉含量升高的原因可能是ABA增强了淀粉磷酸化酶的活性,果糖与葡萄糖比值的降低可能是由于ABA对PGI活性抑制和对PGM活性促进的双重影响。     

Effects of Weak Light on Starch Accumulation and Starch Synthesis Enzyme Activities in Rice at the Grain Filling Stage

Photosynthetic product in rice plant is transported to grains mainly in the form of sucrose. Sucrose in the grains eventually becomes starch through a series of enzymatic reactions [1-2]. The starch weight accounts for 90% of the brown rice [3-4], so the course of the rice grain filling is the biochemical process in which starch is mainly formed, and the factor of light is indispensable at the course. Much research has been conducted to study the influence of the light on the grain starch synt…     

菠萝果实发育过程中糖积累与其代谢酶的关系

以菠萝[Ananas comosus(L.)Merr.]"巴厘"和"台农11号"两个品种为试验材料,测定不同发育时期果实中蔗糖、葡萄糖和果糖含量以及蔗糖代谢相关酶--酸性转化酶(AI)、中性转化酶(NI)、蔗糖合成酶(SS)和蔗糖磷酸合成酶(SPS)的活性,并对果实中糖积累与酶活性的关系进行分析.结果表明:2个品种果实中糖积累动态十分相似,即果实发育前期,糖积累缓慢;进入成熟期,蔗糖迅速积累,而己糖"巴厘"果实中前期变化较少,后期稍有积累,"台农11号"果实中前期稍有积累,后期基本不变.2个品种果实中蔗糖含量与SS和SPS活性相关系数分别达到正的显著或极显著水平,NI活性与2个品种果实中蔗糖含量呈极显著负相关,蔗糖含量与AJ的活性相关性品种间差异明显.     

4种蔗糖分解酶在甘薯块根品质形成中的作用

本研究以2个在可溶性糖和淀粉含量上存在显著差异的鲜食型甘薯品种为材料,对其块根中的可溶性糖（蔗糖、葡萄糖和果糖）和淀粉含量以及4种蔗糖分解酶的活性进行测定与相关性分析,以明确调控甘薯块根中可溶性糖和淀粉含量的关键蔗糖分解酶种类。结果表明：（1）明确甘薯块根中活性最高的蔗糖分解酶为蔗糖合成酶（Sus）,其次为细胞质转化酶（CIN）和液泡转化酶（VIN）,细胞壁转化酶（CWIN）的活性最低。（2）本研究表明,除了已报道的VIN外,CIN活性也和己糖含量之间呈显著正相关,高己糖含量的甘薯品种块根中具有更高的VIN和CIN活性;（3）甘薯块根中Sus活性和淀粉含量之间呈极显著正相关,高淀粉甘薯品种具有更高的Sus活性;（4）CWIN活性非常低,其在块根发育中的具体作用有待进一步深入研究。总之,蔗糖分解酶对甘薯品质有重要影响,其中Sus是调控甘薯块根淀粉含量的关键酶类,而CIN和VIN是调控块根中己糖含量的关键酶类。     

Factors that determine grain weight in rice under high-yielding aerobic culture: The importance of husk size

In water-saving rice culture, yield is unstable because spikelet number per unit area and grain weight fluctuate according to water availability. In this study, we investigated the factors that determine grain weight in aerobic culture. We grew four rice varieties in non-puddled, unsaturated (aerobic) soils with a soil water potential at 20-cm depth kept above −60 kPa and in continuously flooded culture in two years. We found a significant variety × water interaction in grain weight in 2009: weights under aerobic culture were 6% and 13% larger than under flooded culture in Sasanishiki and IRAT109, respectively, versus 4% and 10% smaller in Habataki and Takanari. There was no significant variety × water interaction in grain weight in 2010. Sink activity (grain sucrose synthase activity) and source capacity (biomass production and nonstructural carbohydrate content in vegetative tissue) per plant during ripening were higher under aerobic culture than under flooded culture in both years. However, an excessive increase in spikelet number per unit area in Takanari under aerobic culture in 2009 reduced the source capacity per spikelet and single husk size, decreasing grain weight. In 2010, frequent soil drying under aerobic culture during the late reproductive period (around 20 days preceding heading) reduced single husk size, thereby decreasing grain weight. We found that sink activity and source capacity per plant could be both higher under aerobic culture during the ripening period, producing larger grain weight at a soil water potential above-40 kPa at a 20-cm depth relative to those under flooded culture. In contrast, greater drying under aerobic culture during the late reproductive period reduced single husk size, thereby reducing grain weight.