水稻胚乳中淀粉合成相关酶活性的变化对支链淀粉精细结构的影响

分析了水稻（Oryza sativa L．）籽粒发育过程淀粉生物合成途径中的关键酶——ADP-葡萄糖焦磷酸化酶、可溶性淀粉合酶、淀粉分支酶以及淀粉脱支酶活性变化,同时研究了淀粉结构形成动态．与野生型晚粳9522相比,转基因晚粳9522中直链淀粉的合成被显著抑制,而总淀粉含量和籽粒终重量没有改变．淀粉生物合成途径中关键酶活性表达时间不一致,存在明显的时段特征,这与淀粉积累动态密切相关．可溶性淀粉合酶活性表达最早,其在灌浆前期驱动淀粉合成起始;而淀粉粒结合态淀粉合酶在胚乳发育的中期活性最大．两水稻实验材料间,除淀粉脱支酶活性变化有所不同外,ADP-葡萄糖焦磷酸化酶和淀粉分支酶活性的变化没有明显差异．并且,支链淀粉的分支模式在水稻籽粒发育过程中变化较大,且与品种有关．以上结果揭示,支链淀粉的合成要先于直链淀粉,并且在控制支链淀粉各分支的形成过程中有不同的酶在起特异的作用．     

冬小麦籽粒淀粉合成相关酶活性的日变化

蔗糖向淀粉的转化是决定小麦籽粒产量的重要因素.田间条件下研究了两个小麦(Triticum aestivum L.)品种"鲁麦22"和"鲁麦14"籽粒淀粉合成相关酶:蔗糖合酶(sucrose synthase,SS)、腺苷二磷酸葡萄糖焦磷酸化酶(ADP-glucose pyrophosphorylase,ADPGPPase)、可溶性淀粉合酶(soluble starch synthase,SSS)、束缚态淀粉合酶(starch granule-bound synthase,GBSS)的活性以及籽粒ATP含量的日变化.结果表明,上述酶活性呈现明显的昼夜变化特征,酶活性一般在白天较低,而在夜间呈现较高活性,而籽粒ATP含量趋势相反.相关分析表明,白天较低的酶活性可能与气温超过其适宜温度有关.对籽粒淀粉合成相关酶活性日变化的可能因子进行了讨论.     

冬小麦籽粒淀粉合成相关酶活性的日变化

蔗糖向淀粉的转化是决定小麦籽粒产量的重要因素。田间条件下研究了两个小麦(TriticumaestivumL.)品种“鲁麦22”和“鲁麦14”籽粒淀粉合成相关酶:蔗糖合酶(sucrosesynthase,SS)、腺苷二磷酸葡萄糖焦磷酸化酶(ADP-glucosepyrophosphorylase,ADPGPPase)、可溶性淀粉合酶(solublestarchsynthase,SSS)、束缚态淀粉合酶(starchgranule-boundsynthase,GBSS)的活性以及籽粒ATP含量的日变化。结果表明,上述酶活性呈现明显的昼夜变化特征,酶活性一般在白天较低,而在夜间呈现较高活性,而籽粒ATP含量趋势相反。相关分析表明,白天较低的酶活性可能与气温超过其适宜温度有关。对籽粒淀粉合成相关酶活性日变化的可能因子进行了讨论。     

花后高温对不同耐热性小麦品种籽粒淀粉形成的影响

以耐热性不同的2个小麦品种济麦20和鲁麦21为材料,分别于花后5～9d（T1）和15～19d（T2）进行高温处理,研究了小麦花后不同阶段高温对籽粒淀粉积累、淀粉粒分布及相关酶活性的影响。结果表明,花后高温显著降低籽粒淀粉积累量;显著降低籽粒淀粉及支链淀粉含量,但提高直链淀粉含量、直／支链淀粉比例。处理间比较,他处理对籽粒淀粉积累的影响程度较T1处理大。品种间比较,高温对济麦20的影响程度较鲁麦21大。高温使A型淀粉粒的体积、数量和表面积比例显著增加,B型淀粉粒的体积、数量和表面积比例显著降低。T1处理后,两品种籽粒蔗糖含量、蔗糖合酶（SS）和腺苷二磷酸葡萄糖焦磷酸化酶（AGPP）、可溶性淀粉合酶（SSS）、束缚态淀粉合酶（GBSS）和淀粉分支酶（SBE）活性均略高于对照;但济麦20、鲁麦21上述指标分别于花后15、20d开始低于对照。他处理后,两品种籽粒蔗糖含量、SS、AGPP、SSS、GBSS和SBE活性显著低于对照,济麦20上述指标的降幅较鲁麦21大。与其它淀粉合成相关酶相比,高温对籽粒GBSS活性的影响程度较小。两品种处理间籽粒蔗糖含量、SS、AGPP、SSS、GBSS及SBE活性的变化趋势,与淀粉积累量的变化趋势基本一致。说明灌浆期高温使籽粒淀粉积累量降低,一方面是由于籽粒蔗糖供应较低引起糖源不足;另一方面则是由于灌浆中后期淀粉合成相关酶活性下降使淀粉合成受抑所致。     

结实期土壤水分亏缺影响水稻籽粒灌浆的生理原因

通过分析结实期土壤水分亏缺对水稻(Oryza sativa)籽粒中蔗糖向淀粉合成的生理代谢中关键酶活性及籽粒灌浆的调节作用, 探讨土壤水分亏缺影响水稻籽粒灌浆的生理机制。结果表明, 适度土壤水分亏缺诱导了灌浆高峰期(花后15-20天)水稻籽粒中蔗糖合成酶、腺苷二磷酶葡萄糖焦磷酸化酶、可溶性淀粉合成酶及淀粉分支酶活性的增加, 提高了籽粒灌浆中前期(花后10-20天)籽粒中淀粉积累速率和籽粒灌浆速率。但在灌浆后期(花后20-30天)籽粒中, 上述关键酶活性下降较快, 籽粒活跃灌浆期明显缩短, 灌浆前中期灌浆速率的增加不能完全补偿灌浆期缩短带来的同化物积累损失, 导致水分亏缺处理水稻籽粒充实不良, 结实率、籽粒重和产量显著降低。研究认为, 灌浆期土壤水分亏缺引起的灌浆后期籽粒中蔗糖向淀粉合成代谢中一些关键酶活性快速下降和籽粒内容物的供应不足是籽粒淀粉积累总量减少、粒重降低的主要生理原因。     

谷物籽粒淀粉研究进展

谷物籽粒是人类最基本的粮食,其胚乳淀粉与品质有密切的关系.本文就谷物胚乳淀粉的合成积累过程、谷物淀粉合成酶的作用与特性、淀粉特性与品质的关系以及基因工程在改良作物淀粉品质方面的研究进展进行了综述,并对谷物籽粒淀粉进一步的研究提出了展望,为谷物淀粉品质育种提供参考.     

灌浆期高温和水分逆境对冬小麦籽粒蛋白质和淀粉含量的影响

灌浆期高温和水分逆境是影响小麦籽粒产量和品质的关键气候因子。以扬麦9号、徐州26和豫麦34三个小麦品种为材料,利用人工气候室模拟灌浆期高温和水分胁迫环境,研究了花后高温及温度和水分互作对小麦籽粒蛋白质和淀粉形成的影响。结果表明,高温显著提高了小麦籽粒蛋白质含量及清蛋白、球蛋白和醇溶蛋白含量,但降低了谷蛋白含量,导致麦谷蛋白／醇溶蛋白比值降低。高温显著降低了籽粒总淀粉和支链淀粉含量及支／直比。籽粒蛋白质和淀粉及其组分形成所需的适宜昼夜温差随小麦品质类型而异,但温度水平对籽粒蛋白质和淀粉的影响较温差大。在高温和水分逆境下,温度对籽粒蛋白质和淀粉含量的影响较水分逆境大,且存在显著的互作效应。小麦籽粒蛋白质含量均表现为干旱〉对照〉渍水,以高温干旱最高,适温渍水最低;淀粉含量为对照〉干旱〉渍水,以适温对照最高,而高温渍水最低。高温和水分逆境显著提高了籽粒醇溶蛋白含量而降低了谷蛋白含量及支链淀粉含量,使蛋白质谷／醇比和淀粉支／直比降低,以高温渍水对籽粒蛋白质和淀粉组分的影响最为显著。不同品种之间,高蛋白小麦籽粒蛋白质和组分的形成受高温和水分逆境的影响更大,而低蛋白品种籽粒淀粉形成显著受温度和水分逆境的调节。分析表明,在高温和水分逆境下籽粒蛋白质含量与清蛋白和醇溶蛋白显著正相关,籽粒淀粉含量与谷蛋白、支链淀粉含量及支／直比显著正相关。     

糯玉米SW70和SW22是一类新的淀粉合成酶基因功能缺失多突变体

糯玉米是一类相对于粮食玉米的重要天然突变体,其淀粉合成的遗传机制复杂而又多样。以糯玉米自交系5形22、普通玉米自交系5003、SW22（♀）和5003（♂）的杂种定向选育的糯玉米自交系SW70（自交4代）为材料,研究了自交系授粉后1、2、4周时,SW22、SW70和5003籽粒中ADP-葡萄糖焦磷酸化酶（AGPase）、淀粉合酶（SS）、淀粉分支酶（SBE）和淀粉去分支酶（DBE）共4个家族21个成员编码基因表达水平的变化。结果表明：1）SW22中多个淀粉合成酶编码基因的表达动态与SW70及5003中有显著差异。差异主要存在于淀粉合酶SS和淀粉去分支酶DBE基因家族：2）SW22及SW70籽粒中有明显的阢转录产物累积,但累积水平均显著低于5003籽粒中：3）SW22及SW70中颗粒结合淀粉合成酶编码基因Gbssm和异淀粉酶型去分支酶编码基因Iso2完全不表达．意味着SW22胚乳中GbssIIb基因和Iso2基因的功能完全缺失。目前,有关玉米籽粒中GbSSIIb和异淀粉酶型DBEIso2同时突变或单个基因突变均未见报道。推测淀粉合成酶基因GbssIIb和异淀粉酶型DBEIso2功能缺失以及耽功能部分缺失．共同导致了SW22和SW70籽粒中直链淀粉合成部分缺陷。     

糯玉米胚乳淀粉粒粒度分布形成的酶学机理

以7个糯玉米品种为材料,测定其籽粒发育过程中淀粉粒粒度分布及淀粉合成相关酶活性的变化,分析两者之间的关系。结果表明,随着籽粒发育,糯玉米淀粉粒平均粒径逐渐增大,可溶性淀粉合成酶(SSS)和淀粉分支酶(SBE)活性呈单峰曲线变化。籽粒发育前期,小淀粉粒(≤7.4μm)所占体积较大;随着籽粒发育,小淀粉粒所占体积减少,大淀粉粒(>7.4μm)所占体积增多;籽粒发育后期,大淀粉粒所占体积较大。相关分析表明, SSS和SBE活性与大淀粉粒体积增大速率和平均粒径增大速率均呈显著或极显著正相关。因此, SSS和SBE是影响糯玉米胚乳淀粉粒粒度分布形成的主要酶, SSS和SBE活性越高,淀粉粒平均粒径越大,大淀粉粒所占体积越多。     

水稻籽粒成熟过程中淀粉合成及水解酶活力的变化

作者为了要探求水稻自开花至成熟期间籽粒中所含有关澱粉形成和水解酶系活力的变迁以及这些变迁与淀粉累积的关系,以晚粳稻“长萁铁桿青”作材料,在开花後分期取样,剥出穗上籽粒,研磨後制成酶提取液;在pH 6.8及30℃条件下测定各个时期籽粒的澱粉形成酶及水解酶系的活力。测定澱粉形成酶系活力的酶作用液中加有葡萄糖-1-磷酸钾盐,少量可溶性澱粉及失水蘋果酸钠缓冲液。在测定澱粉水解酶系活力的酶作用液中只是不加葡萄糖-1-磷酸。最后均用蒽酮法测定酶作用液中的澱粉量。试验结果发现水稻籽粒中,在开花後即有合成澱粉的活力,以後逐日增加,至乳熟後期糊     

Effects of the activities of key enzymes involved in starch biosynthesis on the fine structure of amylopectin in developing rice (Oryza sativa L.) endosperms

The dynamic changes of the activities of enzymes involving in starch biosynthesis, including ADP-glucose pyrophosphorylase (AGPase), soluble starch synthases (SSS), starch branching enzyme (SBE) and starch debranching enzymes (DBE) were studied, and changes of fine structure of amy- lopectin were characterized by isoamylase treatment during rice grain development, using trans anti-waxy gene rice plants. The relationships between the activities of those key enzymes were also analyzed. The amylose synthesis was significantly inhibited in transgenic Wanjing 9522, but the total starch content and final grain weight were less affected as compared with those of non-transgenic Wanjing 9522 rice cultivar. Analyses on the changes of activities of enzymes involving in starch bio- synthesis showed that different enzyme activities were expressed differently during rice endosperm development. Soluble starch synthase is relatively highly expressed in earlier stage of endosperm de- velopment, whilst maximal expression of granule-bound starch synthase (GBSS) occurred in mid-stage of endosperm development. No obvious differences in changes of the activities of AGPase and SBE between two rice cultivars investigated, except the DBEs. Distribution patterns of branches of amy- lopectin changed continually during the development of rice grains and varied between two rice culti- vars. It was suggested that amylopectin synthesis be prior to the synthesis of amylose and different enzymes have different roles in controlling syntheses of branches of amylopectin.     

稻米淀粉品质形成的关键酶及其分子生物学研究进展

稻米淀粉的形成是影响水稻产量和品质的决定性因素之一。因此,开展稻米淀粉形成过程中所涉及关键酶的研究是非常必要的。随着分子生物学技术的快速发展,有关稻米淀粉品质的研究也越来越深入,并取得了较大进展。该文对水稻淀粉品质形成过程中的关键酶及其分子生物学研究进展进行了较为详尽的综述,主要包括ADP葡萄糖焦磷酸化酶、淀粉合成酶、淀粉分支酶和淀粉去分支酶等,并对该领域的发展趋势进行了展望。     

稻米淀粉品质形成的关键酶及其分子生物学研究进展

稻米淀粉的形成是影响水稻产量和品质的决定性因素之一。因此, 开展稻米淀粉形成过程中所涉及关键酶的研究是非常必要的。随着分子生物学技术的快速发展, 有关稻米淀粉品质的研究也越来越深入, 并取得了较大进展。该文对水稻淀粉品质形成过程中的关键酶及其分子生物学研究进展进行了较为详尽的综述, 主要包括ADP葡萄糖焦磷酸化酶、淀粉合成酶、淀粉分支酶和淀粉去分支酶等, 并对该领域的发展趋势进行了展望。     

Starch metabolism in leaves

Starch is the most abundant storage carbohydrate produced in plants. The initiation of transitory starch synthesis and degradation in plastids depends mainly on diurnal cycle, post-translational regulation of enzyme activity and starch phosphorylation. For the proper structure of starch granule the activities of all starch synthase isoenzymes, branching enzymes and debranching enzymes are needed. The intensity of starch biosynthesis depends mainly on the activity of AGPase (adenosine 5'-diphosphate glucose pyrophosphorylase). The key enzymes in starch degradation are beta-amylase, isoamylase 3 and disproportionating enzyme. However, it should be underlined that there are some crucial differences in starch metabolism between heterotrophic and autotrophic tissues, e.g. is the ability to build multiprotein complexes responsible for biosynthesis and degradation of starch granules in chloroplasts. The observed huge progress in understanding of starch metabolism was possible mainly due to analyses of the complete Arabidopsis and rice genomes and of numerous mutants with altered starch metabolism in leaves. The aim of this paper is to review current knowledge on transient starch metabolism in higher plants.     

稻米品质形成的生理基础研究进展

综述了水稻(Oryza sativa L.)米质形成的生理基础的研究进展,主要包括稻米品质形成期的源库特征、淀粉合成的关键酶以及淀粉粒的形态、结构及其糊化特性等几个内容,从植物生理学角度来揭示稻米品质形成的规律有着重要的理论意义。     

Genetic and Biochemical Evidence for the Involvement of α-1,4 Glucanotransferases in Amylopectin Synthesis

We describe a novel mutation in the Chlamydomonas reinhardtii STA11 gene, which results in significantly reduced granular starch deposition and major modifications in amylopectin structure and granule shape. This defect simultaneously leads to the accumulation of linear malto-oligosaccharides. The sta11-1 mutation causes the absence of an α-1,4 glucanotransferase known as disproportionating enzyme (D-enzyme). D-enzyme activity was found to be correlated with the amount of wild-type allele doses in gene dosage experiments. All other enzymes involved in starch biosynthesis, including ADP-glucose pyrophosphorylase, debranching enzymes, soluble and granule-bound starch synthases, branching enzymes, phosphorylases, α-glucosidases (maltases), and amylases, were unaffected by the mutation. These data indicate that the D-enzyme is required for normal starch granule biogenesis in the monocellular alga C. reinhardtii.     

Different isoforms of starch-synthesizing enzymes controlling amylose and amylopectin content in rice (Oryza sativa L.)

Starch, composed of amylose and amylopectin, greatly influences rice cooking and textural quality, which in turn is controlled by various isoforms of several enzymes. Activity of one or more isoforms of starch‐synthesizing enzymes results in various forms of starch structure based on the amylopectin chain length and average external, internal and core chain length distribution and hence results in varying physicochemical and cooking quality. Since the synthesis of starch is highly complex, it is crucial but essential to understand its biosynthetic pathway, starch structure and effects on the physicochemical properties that control eating and cooking quality, and alongside conduct research on gene/QTL mapping for use in marker-assisted selection (MAS) with a view to improve and select cultivars with most desirable range and class of rice starch properties. This article presents the updates on current understanding of the coordination among various enzymes/isoforms towards rice starch synthesis in endosperm and their effect on rice grain physicochemical, cooking and eating qualities. The efforts in identifying regions responsible for these enzymes by mapping the gene/QTLs have provided a glimpse on their association with physicochemical and cooking properties of rice and, hence, improvement is possible by modifying the allelic pattern, resulting in down or nil regulation of a particular enzyme. The clear understanding of the tissue specific coordination between enzyme isoforms and their subsequent effect in controlling eating and cooking properties will enhance the chances to manipulate them for getting desired range of amylose content (AC) and gelatinization temperature (GT) in improved cultivars through combining desired alleles through MAS.     

LOCALIZATION OF STARCH BIOSYNTHETIC AND DEGRADATIVE ENZYMES IN MAIZE LEAVES

The cellular distribution of the starch biosynthetic and degradative enzymes in protoplasts prepared from maize leaf mesophyll and bundle sheath cells was investigated. In conformity with the cellular distribution of starch, starch biosynthetic enzymes (soluble starch synthase, ADPglucose pyrophosphorylase, branching enzyme and starch Phosphorylase) were exclusively localized in the bundle sheath cells. In contrast, starch degradative enzymes (α-amylase, β-amylase and debranching enzyme) were present in both types of leaf cells. Isolated chloroplasts from bundle sheath cells were shown to contain 100% of the starch biosynthetic enzymes. However, approximately 60% of the activity of degradative enzymes and 67% of the activity of starch Phosphorylase was localized in bundle sheath chloroplasts.     

转基因技术改良稻米淀粉品质的研究进展

淀粉是稻米胚乳的主要组成成分, 而淀粉含量和性质的差异又直接决定稻米的品质。早期关于稻米淀粉的研究局限于其组成成分、品种间含量的差异以及稻米淀粉合成的经典遗传学方面。随着现代分子生物学和转基因技术的发展, 稻米品质、淀粉的生物合成及其分子调控成为研究热点。随着淀粉合成相关基因的克隆、分子特性及其表达调控的研究取得了较大进展, 人们也开始尝试利用现代基因工程技术, 高效地改良稻米品质, 如提高或降低淀粉含量及改变支链淀粉的结构。本文从综述稻米淀粉的组成、结构和淀粉合成相关酶的研究进展入手, 探讨了转基因技术改良稻米品质的研究进展和发展前景。