水稻淀粉合成关键酶的研究进展

淀粉是稻米的主要成份,其组成和性质决定着稻米品质。在水稻淀粉合成过程中,淀粉合成关键酶发挥了重要的作用。综述了ADP-葡萄糖焦磷酸化酶（AGPase）、淀粉合成酶（SS）、淀粉分支酶（SBE）及淀粉脱支酶（DBE）的功能及调控机理,并对目前研究存在的问题进行了探讨。     

Identification of allelic variations of puroindoline genes controlling grain hardness in wheat using a modified denaturing PAGE

The amount of long chains (LC) of amylopectin in high-amylose rice is thought to be one of the important determinants of its quality when cooked. A wide range of differences in LC content have been reported in rice varieties, which can be clearly divided into four classes based on LC and apparent amylose content: namely, amylose and LC-free, low or medium-amylose and low-LC, high-amylose and medium-LC, high-amylose and high-LC. However, genetic factors controlling LC content have not been fully understood. Here, we performed quantitative trait loci (QTL) analysis of LC content using 157 recombinant inbred lines (RILs) derived from a cross of a low-LC cultivar, Hyogokitanishiki, and a high-LC line, Hokuriku 142. By analyzing randomly selected 15 RILs, it was shown that high LC content (≥11%) was associated with high setback viscosity (≥200 RVU), and that low LC (≤ 3%) was associated with low setback viscosity (≤ 130 RVU), as measured by a Rapid Visco Analyzer. With setback viscosity as an indicator for LC content, QTL analysis was conducted using 60 DNA markers including a CAPS marker that distinguished Wx a and Wx b alleles coding for granule-bound starch synthase I (GBSSI or Wx protein), the enzyme working for amylose biosynthesis. Only one QTL with a peak log of likelihood score at the wx locus was detected, and no line showing setback viscosity corresponding to the medium-LC class appeared. The fact that wx mutants of Hokuriku 142 lacked LC in their rice starch supports the view that the functional Wx allele is indispensable for LC synthesis in addition to amylose synthesis in rice endosperm. We suggest three possible reasons why no line with medium-LC content was observed. First, the locus (loci) responsible for generation of medium-LC may be located very close to the wx locus and not able to be dissected by the population and DNA markers we used. Second, there may be special QTLs for medium-LC cultivars that do not exist in low- or high-LC cultivars. Third, medium-LC cultivars may have an as-yet unidentified Wx allele with lower capability in LC synthesis compared to the Wx allele in high-LC cultivars.     

水稻籽粒淀粉结构及其生物合成与米质关系研究进展

淀粉是水稻籽粒的主要成分,也是人们食物的主要能量来源。许多研究者开始关注淀粉的生物合成途径,以期通过控制这一途径,来改善植物淀粉合成的数量和质量。然而,这一目标实现需要广泛了解淀粉的生物合成以及淀粉的理化性质与淀粉颗粒结构之间的关系。对于淀粉颗粒的结构、组成及支链淀粉的链长分布,前人已做较深入的研究,并且对淀粉合成过程中的关键酶与米质主要指标的关系也进行了剖析。此文就淀粉合成和淀粉颗粒结构与稻米品质的关系展开综述,为水稻栽培和稻米品质改良提供理论依据。     

Starch characterisation and variability in GBSS loci of synthetic hexaploid wheats and their durum and <Emphasis Type="Italic">Aegilops tauschii</Emphasis> parents

Greater variability in starch properties is found in lower ploidy wheats than in commercial hexaploid wheats. This paper reports on the starch properties and variability in granule bound starch synthase (GBSS) loci of 17 diploid (Aegilops tauschii) and 12 tetraploid (durums) potential progenitors of wheat, compared with 29 synthetic hexaploid wheats produced from such progenitors. Starch properties examined were granule size distribution, swelling power, amylose content, gelatinisation and amylose-lipid dissociation properties. A PCR screening method was able to detect the presence or absence of each of the three GBSS genes. It also detected polymorphisms in eight diploids and nine hexaploids, all displaying the same 25 bases deletion in the D genome allele of GBSS. Two tetraploids and five hexaploids were null 4A for GBSS. There was little difference in the amylose contents and amylose-lipid dissociation peak temperatures of the synthetic hexaploids and the lower ploidy wheats. The synthetic hexaploids showed intermediate swelling power values with the durums giving the highest swelling powers. The durums also had higher B granule contents than the A. tauschii accessions, but not as high as the synthetics. However, the A. tauschii samples gave the highest gelatinisation peak temperatures. The presence of the null 4A mutation was positively correlated with swelling power, amylose content and DSC measurements. The new smaller D genome allele of GBSS was associated with slightly higher swelling power. These results confirm the value of wheat progenitor lines as sources of new starch properties for hexaploid wheat. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

QTL for rice grain quality based on a DH population derived from parents with similar apparent amylose content

A doubled haploid (DH)population consisting of 135 lines, derived from an indica (IR64) and a japonica (Azucena) rice with a similar apparent amylose content (AAC), was used to investigate the genetic factors affecting cooking and eating quality of rice. AAC,gelatinization temperature (GT), gel consistency (GC) and six starch pasting viscosity parameters were measured for quantitative trait loci (QTL) analysis using 193 molecular markers mapped on the DH population. A total of 17 QTLs were detected for the 9 traits, with at least one QTL and as many as 3 QTLs for each individual trait. No QTL for the measured parameters was found at the wx locus,possibly because of the similar AAC between the parents. Several QTLs with important effects on the variations in the measured parameters were detected in the present study which have not been found in earlier reports based on populations derived from parents with different AAC and wxgene alleles. Two interesting loci could be deduced from the present study according to the marker order compared with other genetic linkage maps. A QTL flanked by Amy2A and RG433 on the end of the long arm of chromosome 6, identified for GT, set back and consistency viscosity, might cover the gene encoding starch branching enzyme I. Similarly, a QTL flanked by RG139 and RZ58on chromosome 2, detected for hot paste viscosity and breakdown viscosity, might cover the gene encoding starch branching enzyme III. Generally, traits significantly correlated with each other shared identical QTL, but it was not true in some cases. The fine molecular mechanisms underlying these traits await further elucidation for the improvement of eating and cooking quality of rice. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effect of dosage at the soft starch (h) locus on starch granule volume in maize

Larger particle volume is beneficial for many aspects of maize starch processing, and may improve the performance of some starch attributes. Of the numerous endosperm carbohydrate mutants, the soft starch (h) mutant appears to have the greatest influence on starch particle volume. The objective of this study was to determine the gene dosage effect upon starch particle volume at the h locus. Two inbreds, B37 and Oh43, were studied in 1992 and 1993, and their hybrid B37 × Oh43 was studied in 1993. Appropriate crosses were made to establish four endosperm genotypes ++/+, ++/h, hh/+ and hh/h. Starch was extracted from mature kernels collected for each endosperm genotype and analysed for particle volume. The source of variation attributable to fitting two straight regression lines on number of h alleles was highly significant (P < 0.01) in all experiments, resulting in a horizontal‐line relationship among the three normal genotypes (++/+,++/h and hh/+) at 14.452 μm³3 and a regression line of slope 1.622 μm³ passing from hh+ to hh/h. There is no dosage effect of starch particle volume at the h locus; complete recessiveness of the h allele for starch particle volume is the same as the visual phenotypic observation of soft starch.     

水稻籽粒淀粉品质的影响因素及其机制研究进展

淀粉作为水稻籽粒的主要贮藏物质,其生物学特性直接决定着稻米的品质。淀粉形成的生理生化过程极为复杂,容易受到内源和外源因素的影响。为了充分了解水稻籽粒淀粉的发育机制,从发育、生理、遗传和分子生物学角度,归纳了水稻淀粉的生物学特性、淀粉形成的生理机制,并总结了关键酶、基因和激素之间的关系,分析了环境因素（如温度、光照、水分、氮素等）对水稻淀粉品质的影响,指出影响水稻淀粉品质的生物学机制研究中存在的不足。本研究为今后深入研究稻米淀粉的合成过程理清思路,并提出了提高稻米淀粉品质可行方法,为水稻避免不良环境因素提供相关知识,同时为作物遗传改良提供参考。     

高抗性淀粉粳稻新品系稻米淀粉特性

【目的】比较分析高低抗性淀粉含量水稻品种（系）间淀粉主要特性差异。【方法】利用上海农业科学院最新培育的抗性淀粉含量有极显著差异的粳稻品种（系）“降糖稻1号”和 “金丰”,比较分析了支链淀粉链长分布和稻米RVA谱特征值及淀粉分子结构差异。【结果】结果表明,降糖稻1号短链部分所占相对比率明显低于金丰,而长链部分所占相对比例,明显高于金丰;降糖稻1号的最高黏度值,热浆黏度值,崩解值显著或极显著低于金丰,而消减值极显著高于金丰;淀粉颗粒热稳定性也较金丰高;降糖稻1号淀粉晶体主要为B型晶体和B、V混合晶型,而金丰淀粉主要为A型晶体和A、V混合晶型。【结论】高低抗性淀粉含量水稻品种（系）间支链淀粉的链长分布、直链淀粉含量、稻米RVA谱特性值和淀粉晶体结构有很大的差异,在育种实践中可作为育种选择指标之一。     

Developing gene-tagged molecular markers for functional analysis of starch-synthesizing genes in rice (Oryza sativa L.)

The granule-bound starch synthase(GBSS), starch branching enzymes 1 (SBE1)and 3 (SBE3) are major enzymes involved in starch biosynthesis in rice endosperm. Available sequences of Sbe1 and Sbe3 genes encoding corresponding SBE1 and SBE3 have been used to identify homologous regions from genome databases of both the indica rice 93-11 and the japonica rice Nipponbare. Sequence diversities were exploited to develop gene-tagged markers to distinguish an indica allele from a japonica allele for both Sbe1 and Sbe3 loci. With these newly developed gene-tagged markers and available Wx gene markers, the roles of these starch-synthesizing genes (Sbe1, Sbe3, and Wx) in determining amylose content (AC) in the rice endosperm were evaluated using a double-haploid (DH)population derived from a cross between the indica rice cv. Nanjing11 and the japonica rice cv. Balilla. Only the Wx and Sbe3 loci had significant effects on the AC variation. On average, indica Wx ^a genotypes showed ∼9.1% higher AC than japonica Wx ^b genotypes, while indica Sbe3 ^a genotypes showed ∼1.0% lower AC than japonica Sbe3 ^b genotypes. A significant interaction was also observed between Wx and Sbe3 loci whereby the amylose content was 0.3% higher in Sbe3 ^a than Sbe3 ^b genotypes in the presence of the Wx ^a allele, but it was lower by 2.3% in the presence of the Wx ^b allele. Overall, polymorphisms at the Wx and Sbe3 loci together could account for 79.1% of the observed AC variation in the DH population. The use of gene-tagged markers in marker-assisted selection and gene functional analysis was also discussed. This revised version was published online in July 2006 with corrections to the Cover Date.     

不同介质对糯米淀粉糊化特性的影响

采用Brabender黏度仪,测定糯米淀粉糊在不同浓度的碱面、明矾、食盐和蔗糖条件下的黏度曲线,研究了这些因素对糯米淀粉糊化特性的影响.结果表明,添加碱面、明矾、食盐和蔗糖提高了糯米淀粉的糊化温度、热稳定性,减弱了老化程度.碱面、明矾、食盐对糯米淀粉糊化特性影响较大,而蔗糖的影响较小.     

绿豆羧甲基淀粉糊性质的研究

对绿豆羧甲基淀粉糊性质进行了详细的研究,包括糊的冻融稳定性、抗生物降解能力、抗老化能力、糊透明度、pH值、介质(蔗糖和NaCl)对糊黏度的影响。结果表明,绿豆羧甲基淀粉冻融稳定性好,抗霉菌能力及抗老化能力强,透明度高。糊的热黏度稳定性降低,冷黏度稳定增加。NaCl强电解质对绿豆羧甲基淀粉糊黏度影响较大。     

生长季节对糯玉米淀粉粒分布和热力学特性的影响

以8个糯玉米品种为材料,利用激光衍射粒度分析仪和差示扫描量热仪分析了春播和秋播糯玉米淀粉粒分布和热力学特性。结果表明,糯玉米淀粉主要由直径<13 μm的颗粒组成,占总数目的71.7%~84.3%。颗粒体积和表面积呈双峰分布,>17 μm颗粒占比例最高,9~13 μm颗粒占比例较高,<9 μm颗粒占比例最低。和春播相比,秋播处理使糯玉米粉中<17 μm颗粒体积和表面积比例升高,>17 μm颗粒体积和表面积比例降低,<9 μm的颗粒数目增多,>9 μm颗粒数目减少。淀粉热力学特征参数中,秋播处理下转变温度(起始温度、峰值温度和终值温度)、峰值指数和回生值降低,糊化范围扩大,但热焓值总体上无显著变化。结合前期研究结果,9~13 μm和13~17 μm颗粒体积比例与淀粉的结晶度、峰值黏度和崩解值显著正相关,与峰值温度和回生值显著负相关。>17 μm颗粒体积比例与结晶度、峰值黏度和崩解值极显著负相关,与峰值温度和回生值显著正相关。糯玉米淀粉粒分布在不同季节下不同,导致淀粉糊化特性和热力学特性发生变化。     

再生稻次适宜区迟播栽对不同杂交籼稻淀粉RVA谱的影响

为探讨再生稻次适宜区播栽期对杂交籼稻淀粉RVA谱的影响,以2个杂交籼稻品种,川优6203和宜香优2115为材料,在四川的隆昌和犍为2个生态点,研究不同生态条件下推迟播栽期对籼稻淀粉RVA谱特征值的影响。结果表明,播栽期的主效应对籼稻淀粉RVA谱特征值存在明显的影响。在再生稻次适宜区,推迟播栽期会导致水稻灌浆期温度降低,日照时长缩短,降雨量减少,从而降低了籼稻淀粉RVA谱的峰值黏度、热浆黏度、冷胶黏度和崩解值,提高了消减值以及峰值时间;与再生稻相比,迟播栽期处理具有较大的峰值黏度、崩解值以及较小的冷胶黏度、消减值、回复值、峰值时间和糊化温度。播栽期对籼稻淀粉RVA谱特征值的影响在不同生态点具有差异,与隆昌生态点相比,犍为生态点处理间的峰值黏度、消减值和崩解值差异更为显著。播栽期对不同品种淀粉RVA谱的影响程度不同,宜香优2115的峰值黏度和崩解值比川优6203更容易受到播栽期的影响。从淀粉RVA谱的变化趋势来看,迟播栽期处理的籼稻淀粉RVA谱表现与再生稻更为接近,蒸煮食味品质更高。     

水稻籽粒灌浆过程直链淀粉的积累及其相关酶的品种类型间差异

对籼、粳、糯3种类型水稻籽粒灌浆过程中直链淀粉含量的变化及其相关酶活性差异的比较分析表明,不论是早籼品种浙733,还是早粳品种浙农104,其直链淀粉含量在籽粒灌浆过程中的变化趋势基本相似,均表现为灌浆初期的直链淀粉含量较低,随着灌浆时间的推移,籽粒中直链淀粉含量明显升高,灌浆后期则略有降低,而对早糯品种早香糯而言,     

稻米淀粉黏滞性QTL定位及其G×E互作分析

用珍汕97B/密阳46构建RIL群体及其遗传图谱,经海南和杭州两地遗传试验,以精米粉RVA谱5个参数特征值PKV、HPV、CPV、BDV和SBV作为研究稻米淀粉黏滞性的指标,运用检测QTL主效应、上位性效应和G×E互作效应的遗传分析方法,进行QTL联合分析。结果表明, (1)在检测到涉及5个性状的9个主效应QTL中,除PKV位于第5染色体qPKV5外,其余8个QTL均位于第6染色体上;(2)5个性状均检测到位于第6染色体RM197-RZ516区间的主效应QTL,很可能它们为同一基因,该基因还与Wx基因处于相同区域;(3)检测到与PKV、HPV、CPV、BDV等4个性状有关的QTL主效基因均表现有G×E互作,且方向一致,在海南试验中有增效作用;(4)还检测到涉及5个性状的10对上位性互作效应,但均未发现有显著的上位性×环境互作效应。     

Quality variations in transgenic rice with a synthetic cry1Ab gene from Bacillus thuringiensis

In order to estimate the potential of transgenic rice, characteristics related to grain quality and starch viscosity were investigated in six japonica lines based on three primary transgenic lines containing a synthetic cry1Ab gene from Bacillus thuringiensis. No significant differences were found between the transgenic lines and the wild type, including negative lines and an untransformed line. All six transgenic lines were comparable in size, milling quality, appearance quality and physicochemical properties to the wild type that were derived from. One exception was that the lines derived from the primary transgenic line TR0‐101 had smaller grains. Crude protein contents were equivalent in all the material tested, but Cry1Ab protein was only detected in grains of transgenic rice and was undetectable in the cooked rice. The viscosity of the starch differed between the transgenic lines, the wild type and other controls, and two transgenic lines had breakdown values (BDV) and setback values (SBV) similar to the wild type. A positional effect of T‐DNA insertion on starch viscosity was found in three primary transgenic lines.     

几种红小豆理化特性及淀粉性质研究

旨在通过比较不同品种红小豆的理化特性及淀粉性质,筛选优质红小豆品种。本试验以7个不同产地不同品种红小豆为代表性的试验材料,进行理化测定,包括豆粒水分、百粒重、直链淀粉、出沙率。通过湿法提取淀粉,测定其基本性质（水分、脂肪、灰分、粒径、颗粒形貌及黏度）。结果表明：7个品种红小豆理化性质具有显著性差异,豆粒水分含量10.27%~16.35%,百粒重7.02~14.45 g,直链淀粉28.30%~35.43%以及出沙率60.40%~67.85%,理化性质测定中5号样品均为最高值。几种红小豆淀粉特性具有显著性差异,灰分0.05%~0.43%,水分8.85%~9.64%,脂肪0.04~0.13 g/100 g,粒径值549.32~1460.40 nm。峰值黏度为2459.50~4121.00 cp,最终黏度为4054.50~4765.50 cp,跌落值178.50~1257.50 cp,5 号样品回升值最高为2201.50±13.435 cp。豆粒水分、百粒重、直链淀粉含量、出沙率是影响红小豆品质的主要因子。淀粉颗粒粒度、形貌及黏度可作为红小豆加工品质评价的依据,本试验筛选出5 号‘天津红’加工品质优于其他6种红小豆。     

花后高温对稻米品质及胚乳淀粉粒结构的影响

为阐明花后高温对稻米品质的影响机理,以耐热性不同的两个水稻品系为材料,在人工气候室高温和适温条件下,研究了花后高温胁迫对稻米外观品质、加工品质、蒸煮食味品质及胚乳淀粉粒结构的影响。结果表明,与对照相比,花后高温胁迫条件下,稻米长宽比增大,垩白粒率和垩白度显著增加, 精米率和整精米率显著降低,直链淀粉含量和胶稠度降低,蛋白质含量增加;高温下淀粉RVA谱特征值最终黏度、消减值和糊化温度升高,崩解值降低,峰值时间延长,Mg/ K比值降低。相同处理条件下,热敏感品系4628稻米品质受高温影响程度大些。花后高温条件下,稻米胚乳淀粉粒排列不紧密,淀粉粒间空隙大,多为单粒淀粉体,导致光线的多角度散射,胚乳透明度降低,垩白形成。花后高温条件下,稻米蛋白质含量的增加、Mg/K降低、稻米RVA谱特征值及胚乳淀粉粒结构的变化是导致稻米蒸煮食味品质和外观品质降低的主要原因。     

刺苦草根状茎淀粉特性的研究

系统研究了刺苦草根状茎淀粉的理化性质。结果表明,刺苦草根状茎淀粉形状较一致,呈椭圆形、锥形,淀粉颗粒表面光滑,大小在8～36μm之间。X-射线呈B型结晶图样,结晶度为47.2%。刺苦草根状茎淀粉的糊化温度为66℃。与对照相比,刺苦草根状茎淀粉具有较高的透明度,较好的冻融稳定性,较易凝沉。刺苦草根状茎淀粉糊具有非牛顿流体的特性,属于剪切稀化体系。     

钾肥对高淀粉马铃薯块茎淀粉合成相关酶活性的影响

以东北地区选育的2个高淀粉马铃薯品种东农310和克新22号为试验材料,研究不同施钾量对高淀粉马铃薯品种块茎形成过程中淀粉合成相关酶活性的影响。结果表明,随着生育期的推进,不同施钾量处理马铃薯品种东农310和克新22号块茎中的腺苷二磷酸葡萄糖焦磷酸化酶(AGPase)、颗粒凝结性淀粉合成酶(GBSS)、可溶性淀粉合成酶(SSS)、淀粉分支酶(SBE)都呈现先增加、达到峰值后又开始下降的单峰曲线变化。增施钾肥对东农310和克新22号块茎淀粉合成中AGPase和GBSS活性影响比较大,而对SSS和SBE的活性影响较小。