木薯块根淀粉形态发生与积累的酶活性动态初步研究
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摘要
蔗糖的积累、运输及转化从“源”上影响库器官的发育及淀粉的积累,而库器官的发育及淀粉合成直接关系到木薯淀粉产量。本研究选用形态特征及来源差异较大的三个木薯品种(SC124、SC8、Arg7)为材料,对植后100至200天块根膨大时期淀粉及其组分的积累动态进行检测,分析不同木薯品种淀粉积累的规律及特点；研究功能叶的还原糖、蔗糖含量及蔗糖代谢关键酶的动态变化,分析源叶的生长状况、糖分积累及关键酶活性动态变化与库器官淀粉含量的关系；研究库器官中还原糖含量、蔗糖含量、蔗糖代谢及淀粉合成关键酶的动态变化,分析库器官糖分积累、蔗糖代谢及淀粉合成关键酶与块根淀粉含量的关系。结果如下：
1、对三个木薯品种植后100天至200天块根淀粉及其组分积累动态特征进行比较研究。结果表明,随着块根膨大发育,三个品种单株淀粉总量持续升高,而淀粉含量波动增加。植后120至200天总淀粉含量为Arg7>SC124>SC8, Arg7与其它两个品种差异显著。三个木薯品种块根淀粉积累量和积累速率变化随时间基本呈单峰曲线变化,SC124和SC8早于Arg720天达到积累量和积累率的峰值,且三个木薯品种峰值处的积累速率为SC124>SC8>Arg7。由此可见,SC124和SC8淀粉积累速率较快,而Arg7淀粉积累的持续期较长。三个品种块根支链淀粉含量、支链淀粉积累量和积累速率趋势与总淀粉相似,SC8各时期的支链淀粉含量均值最低,Arg7迟于SC124和SC820天达到积累量和积累率的峰值,三个品种在峰值处的积累速率为SC124>SC8>Arg7。在P<0.05水平,品种间直链淀粉含量差异不显著,而SC124和SC8块根支链淀粉和总淀粉含量与Arg7差异显著。
2、对植后100天至200天的木薯从“源”到“库”的一系列蔗糖代谢和淀粉合成关键酶活性动态及与块根淀粉积累的相关性进行研究。结果表明,三个木薯品种叶片酸性转化酶(AI)总体呈下降趋势,磷酸蔗糖合成酶(SPS)和中性转化酶(NI)都呈单峰型曲线变化；品种间比较,各时期Arg7的AI活性都较低,而SC124 SPS和NI活性都较高。相关性分析表明,SPS与NI极显著正相关,表明SPS主要与NI协同调节蔗糖的含量；叶片AI活性与还原糖含量呈极显著正相关,与块根中淀粉含量极显著负相关,推断叶片中酸性转化酶活性低,使叶片中分解产生的还原糖少,意味着有较多的蔗糖向块根中转运,有利于块根淀粉的膨大。
三个品种块根的AI、NI活性变化趋势一致,在块根膨大早期较高,而SS活性峰值出现较晚,且SC124早于SC8和Arg7达到峰值；块根NI与AI活性的极显著正相关,与淀粉含量呈显著负相关,推测NI和AI可能协同作用,分解蔗糖产生的还原糖除供作淀粉合成底物,还主要用于调节与库膨大发育有关的生长,淀粉含量却由于库器官的快速膨大而略有下降。三个品种块根AGPase活性有两个较高时期,第一个高峰时期在早、中迟熟品种中存在差异,第二个高峰时期在早、中迟熟品种中较为一致；三个品种块根SSS.和GBSS都呈单峰曲线型变化,而SBE总体呈下降趋势,Arg7的AGPase和SBE酶的活性在整个生育期显著高于其他两个品种：相关性分析表明,AGPase酶活性与支链淀粉和总淀粉含量均达到极显著相关,检测该酶活性有可能用来评价库器官积累淀粉能力,而其它淀粉合成关键酶与淀粉含量的相关性不显著。
3、以3个亲缘关系较远的高产华南木薯品种(SC124、SC8、Arg7)为材料,介绍木薯AGPase和SBE同工酶电泳及酶活性染色技术,并对木薯块根发育过程中两大家族的SBE的活性变化进行了跟踪观察,结合AGPase、SBE酶活性和淀粉含量检测,初步探讨了木薯淀粉AGPase和SBE同工酶位点与淀粉含量及其组分的关系。同工酶分析表明,木薯AGPase至少有6个等位基因位点(AGPa、AGPb、AGPc、AGPd、AGPe、AGPf).品种间具有多态性。AGPe位点只在Arg7中出现,初步判定同工酶位点AGPe可能与其较高的酶活性和淀粉含量有关。木薯分支酶有SBE-Ⅰ和SBE-Ⅱ两个基因位点,其中SBE-Ⅰ至少有3个等位基因位点(SBE-Ⅰa、SBE-Ⅰb、SBE-Ⅰc),品种间具有多态性；而SBE-Ⅱ为单一条带,SBE-Ⅱ可能不具有遗传多样性,但活性大小在品种间和发育不同阶段有差异。等位基因位点SBE-Ⅰc和SBE-Ⅱ可能对SBE酶活性和支链淀粉含量贡献较大。
4、对SC124组培苗贮藏根诱导中淀粉粒形态发生过程进行了观察。木薯贮藏根诱导初期,淀粉体内有数目不等的淀粉粒,随着淀粉粒的发育,淀粉体被膜破裂释放淀粉粒,观察到贮藏根细胞内大部分淀粉是单粒的。
对3个木薯品种(SC124、SC8、Arg7)植后120天和160天的块根进行了扫描电镜和透射观察。透射电镜观察证明：块根膨大初期(大约植后120天),多数淀粉体还未解体,因此在同时期扫描电镜下观察到的结果主要是淀粉体还未解体时的形态,多极球体等形态为多个淀粉粒在一个淀粉体发育：植后160天,大量成熟淀粉体被膜解体,淀粉体和淀粉粒同时存在；木薯块根淀粉粒主要为单粒淀粉,品种间复粒淀粉数量差异可能与淀粉品质有关。扫描电镜结果表明：植后120天,品种间淀粉体大小及空间排列差异显著,SC124淀粉体平均直径小,变异系数大,淀粉体间空隙较多,呈层状排布；SC8淀粉体大小较为一致,变异系数小,淀粉体排列紧密,呈束状排布；Arg7淀粉体大小及变异系数都较小,淀粉体排列较为疏松,成团状排布；植后160天,三个品种块根的淀粉充实程度显著增加。
透射电镜下观察到木薯淀粉体的两种增殖方式：淀粉质体被膜出泡、外凸，以出芽方式增殖；被膜向内出泡,在淀粉体内形成新的淀粉质体。淀粉体的大小、形态、空间排列及增殖方式及效率可能是影响木薯块根淀粉充实程度和品质特性直接原因。
5、对两个木薯主栽品种(SC124、SC8)自盛暑期开始遮荫,对其淀粉迅速积累的膨大期进行淀粉含量及相关酶分析,结合扫描电镜观察遮荫处理与正常光照下木薯块根贮藏淀粉结构上的变化,探讨弱光对木薯贮藏淀粉积累的影响。结果表明,该试验设计的遮荫处理下,改变了光合产物在“源”与“库”之间的分配,弱光对木薯块根淀粉含量及其组成的影响较为显著,遮荫处理提高或显著提高了SC124和SC8块根直链淀粉含量,降低了总淀粉含量和支链淀粉含量,以及支链淀粉与直支链淀粉的比率。遮荫处理AGPase活性在两个品种中均有降低；GBSS活性增加(SC8)或无显著变化(SC124)；SSS和SBE活性在两品种表现完全不同,SSS在SC124活性增加,但在SC8品种中降低,SBE活性则相反,即在SC8活性增加,但在SC124品种中降低。我们推测总淀粉含量及支链淀粉含量的下降可能与AGPase活性有关；GBSS活性增加与否与直链淀粉含量相对应；SSS和SBE活性任一酶的下降都有可能降低支链含量。从组织结构看,遮荫处理后的两品种淀粉充实度低,淀粉结构疏松,SC8单粒淀粉增多,结合淀粉含量和淀粉粒大小分析表明,遮荫处理使贮藏淀粉中有更多直径较小的淀粉粒,遮荫影响淀粉粒充实程度从而造成淀粉含量降低。
The accumulation, transportation, and conversion of sucrose affect the development of cassava "storage root" and starch accumulation from" source", and starch synthesis, which directly relates to the yield of starch. The period from 100 d to 200 d after planting, three high yield and far related cassava cultivars (SC124, SC8, and Arg7) were used for researches,' which contributed in the following aspects:the changes of root starch and its component accumulation characteristics; the changes of leaf sucrose content, reducing sugar content and enzyme activity involved in sucrose metabolism; the relation between root starch content and leaf growth, sucrose accumulation as well as key enzymes activity involved in sucrose metabolism; the changes of root sucrose content, reducing sugar content and root enzyme activity including in sucrose metabolism and starch synthesis; the relation between root starch content and sugar content along with enzymes activity including in sucrose metabolism and starch synthesis.
1. The characteristics of starch accumulation in cassava storage roots have been studied in the three cassava cultivars (SC124、SC8、Arg7) during root development from 100 d to 200 d after planting. The quantity of total starch per plant was continuously increased, but the starch content (%) was increased at intervals. The starch content (%) was significant higher in Arg7, among which it was Arg7>SC124>SC8. The content and rate of starch accumulation among the three cultivars was changed in the pattern of a single-peak curve during root mature processing, in which SC124 and SC8 was much earlier to reach the peak in content, and in the peak of rate was SC124>SC8>Arg7. Thus, the starch accumulation in SC124 and SC8 was faster than Arg7, but it lasted longer period in Arg7.The total quantity, content, accumulation rate of amylopectin was similar as those of total starch accumulation in the three cultivars. Between the cultivars, the average value of amylopectin content (%)in SC8 was much lower than other two cultivars, but the peak of accumulation rate and quantity of amylopectin was much slowly appeared in Arg7 than SC124 and SC8. At the level of P<0.05, amylose content(%) was not significantly different, but it was significantly different in amylopectin and total starch content (%) among SC124、SC8 and Arg7.
2. The enzymes involved in sucrose metabolism and starch synthesis have been investigated from "source" to" storage roots", and the relationship between enzymes and starch content of storage roots have been discussed. The results are as follows:AI activity was declined during root mature processing, and the activity of SPS and NI was changed in the pattern of a single-peak curve; The AI activity in Arg7 was much lower than that in SC8 or Arg7, and SPS and NI activity in SCI 24 was much higher than that in SC8 or Arg7; The activity of SPS was significantly correlated with the activity of NI, showing the coordinate reduation in controlling sucrose content of leaf; The activity of AI was significantly correlated with the activity of the reducing sugar content in leaves, but was negatively significantly correlated with the root starch content (%), it can be deduced that the lower activity of AI could produce less reducing sugar, which was propitious to sucrose transportation and root development.
The changes of AI and NI activity in storage roots are almost similar among three cassava cultivars, and both of them showed high activity in early developing stage. The peak of SS was much slowly appeared, but kept high level in the later developing stage. The activity of NI was significantly correlated with the AI activity in the storage roots, but was negatively significantly correlated with the root starch content, it can be deduced that the reducing sugar deprived from sucrose not only was used for substrate in starch synthase, but also play an important role in the root growth regulation, so that the starch content may declined slightly as a result of rapid swelling of root. The higher activity of AGPase was identified at two periods, and the first peak of AGPase activity arrived much earlier in SC8 than that in SC124 and Arg7, but the second peak arrived at the same period. The activity of soluble starch synthase (SSS), granule-bound starch synthase (GBSS) was changed in the pattern of a single-peak curve during root mature processing. The activity of AGPase and SBE in Arg7 cultivar with higher starch content was higher than that in SC124 and SC8. The content of total starch and amylopectin was significantly correlated with the activity of AGPase, but was not correlated with the activity of other enzymes (GBSS, SSS and SBE). The activity of AGPase may be used as standard to estimate the capability of starch accumulation in cassava storage roots.
3. The three cassava cultivars (SC124、SC8、Arg7) with far relationship and high-yielding were used for analysis of the isozyme electrophoresis. The correlation between isozyme locus, enzyme activities of SBE and AGPase, and starch content was investigated. The results showed that there were at least six isozyme loci in AGPase (AGPa、AGPb、AGPc、AGPd、AGPe、AGPf), which manifested genetic diversity among three cultivars. The locus of AGPe only appeared in Arg7, and it may closely relate with higher AGPase activity and starch content. SBE I manifested genetic diversity among three cultivars, while SBEⅡdid not exhibit genetic polymorphism, but showed different in activity among different varieties and root developing stages. The loci of SBEIc and SBEⅡmay provide much great contribution to SBE activity and amylopectin content.
4. By studying the starch development of the induced cassava storage roots in vitro of SC124, it indicated that in the early stage of starch grain development, a number of starch grains were developing in a closed amyloplast with membrane. With the maturation of amyloplast, the starch granules were released and many single granules were observed in induces cassava storage roots.
The storage roots of the three cassava cultivars (SC124、SC8、Arg7) in 120 and 160 d after planting have been scanned under SEM and TEM. Our observations by TEM confirm that the images sanned by SEM at 120 d were the amyloplasts with membrance envelope, and the morphology of starch such as momultidimensional were the starch granules at one amyloplast. On the 160 days after planting, the membrane envelop of mature amyloplast was disaggregated, and mature starch granules have been released from the amyloplasts. Both amyloplasts and starch granules can be observed in parenchyma. The starch granules in cassava mainly consist of many simple starch grains, few of them are compound starch granule. The frequency of the starch multigrain varies in different cultivars may be interrelated to starch quanlity. Our observations by SEM show that the size (the average diameters), pattern and arrangement are diversity among the three cassava cultivars:The average size of amyloplasts in SC124 are smallest, but their coefficient of variation and interspaces are larger; the granule size of amyloplasts in SC8 are relative uniform with smaller coefficient of variation and tight arrangement; however in Arg7, the granule size and coefficient of variation are smaller, while the interspaces are larger. SEM image showed that of the amyloplasts are stratiformly arranged in zylem parenchyma in SC124, sarciniformly arranged in SC8; and granularly arranged in Arg7. On the 160 days after planting, the starch granules were increased remarkablely.
Under TEM, two patterns of proliferation were observed:1, envelope dilates and invaginates to form new amyloplasts; 2, the amyloplast envelope buds to form the double membrane vesicle. The size, arrangement, the pattern and efficient of proliferation may maybe the immanent cause of character difference.
5. Two cassava cultivars SC124 and SC8 were used for the study of starch synthesis and accumulation in storage roots in response to shading light to reduce light intensity from the early stage (70 d after planting)to storage root filling stages (160 d after planting). The starch accumulation, starch structures and activities of the enzymes involved in starch synthesis have been examined at the developing stages of cassava storage roots. The results showed that the shading treatment (ST) has changed the distribution of the product from photosynthesis between the "source" and "storage" organs. The content of total starch and amylpectin was remarkably reduced, but the amylose content was significantly increased in comparison to the control (CK, no shading light). The ratio of amylose to amylopectin in ST conditions was significantly higher than that of CK. The activity of adenosine diphosphate glucose pyrophosphorylase (AGPase) in shading conditions was significantly lower than that . in control plants. The activity of GBSS was increased in SC8, but no obviously changed in SC124. The activity of SSS was increased in SC124, but was decreased in SC8, which was contrast to SBE; It was increased in SC8, but was decreased in SC124. It suggests that the low activity of AGPase resulted from shading light may relate to the lower total starch and amylopectin content. The GBSS activity was in response to the change of amylose content; and the any reduced activity of SSS, or SBE may result in lower amylose content. When the cassava was under shading conditions, the organization of starch structure became looser, the size of starch granule was significantly smaller, but the number of the single starch granule was obviously increased.
1、对三个木薯品种植后100天至200天块根淀粉及其组分积累动态特征进行比较研究。结果表明,随着块根膨大发育,三个品种单株淀粉总量持续升高,而淀粉含量波动增加。植后120至200天总淀粉含量为Arg7>SC124>SC8, Arg7与其它两个品种差异显著。三个木薯品种块根淀粉积累量和积累速率变化随时间基本呈单峰曲线变化,SC124和SC8早于Arg720天达到积累量和积累率的峰值,且三个木薯品种峰值处的积累速率为SC124>SC8>Arg7。由此可见,SC124和SC8淀粉积累速率较快,而Arg7淀粉积累的持续期较长。三个品种块根支链淀粉含量、支链淀粉积累量和积累速率趋势与总淀粉相似,SC8各时期的支链淀粉含量均值最低,Arg7迟于SC124和SC820天达到积累量和积累率的峰值,三个品种在峰值处的积累速率为SC124>SC8>Arg7。在P<0.05水平,品种间直链淀粉含量差异不显著,而SC124和SC8块根支链淀粉和总淀粉含量与Arg7差异显著。
2、对植后100天至200天的木薯从“源”到“库”的一系列蔗糖代谢和淀粉合成关键酶活性动态及与块根淀粉积累的相关性进行研究。结果表明,三个木薯品种叶片酸性转化酶(AI)总体呈下降趋势,磷酸蔗糖合成酶(SPS)和中性转化酶(NI)都呈单峰型曲线变化；品种间比较,各时期Arg7的AI活性都较低,而SC124 SPS和NI活性都较高。相关性分析表明,SPS与NI极显著正相关,表明SPS主要与NI协同调节蔗糖的含量；叶片AI活性与还原糖含量呈极显著正相关,与块根中淀粉含量极显著负相关,推断叶片中酸性转化酶活性低,使叶片中分解产生的还原糖少,意味着有较多的蔗糖向块根中转运,有利于块根淀粉的膨大。
三个品种块根的AI、NI活性变化趋势一致,在块根膨大早期较高,而SS活性峰值出现较晚,且SC124早于SC8和Arg7达到峰值；块根NI与AI活性的极显著正相关,与淀粉含量呈显著负相关,推测NI和AI可能协同作用,分解蔗糖产生的还原糖除供作淀粉合成底物,还主要用于调节与库膨大发育有关的生长,淀粉含量却由于库器官的快速膨大而略有下降。三个品种块根AGPase活性有两个较高时期,第一个高峰时期在早、中迟熟品种中存在差异,第二个高峰时期在早、中迟熟品种中较为一致；三个品种块根SSS.和GBSS都呈单峰曲线型变化,而SBE总体呈下降趋势,Arg7的AGPase和SBE酶的活性在整个生育期显著高于其他两个品种：相关性分析表明,AGPase酶活性与支链淀粉和总淀粉含量均达到极显著相关,检测该酶活性有可能用来评价库器官积累淀粉能力,而其它淀粉合成关键酶与淀粉含量的相关性不显著。
3、以3个亲缘关系较远的高产华南木薯品种(SC124、SC8、Arg7)为材料,介绍木薯AGPase和SBE同工酶电泳及酶活性染色技术,并对木薯块根发育过程中两大家族的SBE的活性变化进行了跟踪观察,结合AGPase、SBE酶活性和淀粉含量检测,初步探讨了木薯淀粉AGPase和SBE同工酶位点与淀粉含量及其组分的关系。同工酶分析表明,木薯AGPase至少有6个等位基因位点(AGPa、AGPb、AGPc、AGPd、AGPe、AGPf).品种间具有多态性。AGPe位点只在Arg7中出现,初步判定同工酶位点AGPe可能与其较高的酶活性和淀粉含量有关。木薯分支酶有SBE-Ⅰ和SBE-Ⅱ两个基因位点,其中SBE-Ⅰ至少有3个等位基因位点(SBE-Ⅰa、SBE-Ⅰb、SBE-Ⅰc),品种间具有多态性；而SBE-Ⅱ为单一条带,SBE-Ⅱ可能不具有遗传多样性,但活性大小在品种间和发育不同阶段有差异。等位基因位点SBE-Ⅰc和SBE-Ⅱ可能对SBE酶活性和支链淀粉含量贡献较大。
4、对SC124组培苗贮藏根诱导中淀粉粒形态发生过程进行了观察。木薯贮藏根诱导初期,淀粉体内有数目不等的淀粉粒,随着淀粉粒的发育,淀粉体被膜破裂释放淀粉粒,观察到贮藏根细胞内大部分淀粉是单粒的。
对3个木薯品种(SC124、SC8、Arg7)植后120天和160天的块根进行了扫描电镜和透射观察。透射电镜观察证明：块根膨大初期(大约植后120天),多数淀粉体还未解体,因此在同时期扫描电镜下观察到的结果主要是淀粉体还未解体时的形态,多极球体等形态为多个淀粉粒在一个淀粉体发育：植后160天,大量成熟淀粉体被膜解体,淀粉体和淀粉粒同时存在；木薯块根淀粉粒主要为单粒淀粉,品种间复粒淀粉数量差异可能与淀粉品质有关。扫描电镜结果表明：植后120天,品种间淀粉体大小及空间排列差异显著,SC124淀粉体平均直径小,变异系数大,淀粉体间空隙较多,呈层状排布；SC8淀粉体大小较为一致,变异系数小,淀粉体排列紧密,呈束状排布；Arg7淀粉体大小及变异系数都较小,淀粉体排列较为疏松,成团状排布；植后160天,三个品种块根的淀粉充实程度显著增加。
透射电镜下观察到木薯淀粉体的两种增殖方式：淀粉质体被膜出泡、外凸，以出芽方式增殖；被膜向内出泡,在淀粉体内形成新的淀粉质体。淀粉体的大小、形态、空间排列及增殖方式及效率可能是影响木薯块根淀粉充实程度和品质特性直接原因。
5、对两个木薯主栽品种(SC124、SC8)自盛暑期开始遮荫,对其淀粉迅速积累的膨大期进行淀粉含量及相关酶分析,结合扫描电镜观察遮荫处理与正常光照下木薯块根贮藏淀粉结构上的变化,探讨弱光对木薯贮藏淀粉积累的影响。结果表明,该试验设计的遮荫处理下,改变了光合产物在“源”与“库”之间的分配,弱光对木薯块根淀粉含量及其组成的影响较为显著,遮荫处理提高或显著提高了SC124和SC8块根直链淀粉含量,降低了总淀粉含量和支链淀粉含量,以及支链淀粉与直支链淀粉的比率。遮荫处理AGPase活性在两个品种中均有降低；GBSS活性增加(SC8)或无显著变化(SC124)；SSS和SBE活性在两品种表现完全不同,SSS在SC124活性增加,但在SC8品种中降低,SBE活性则相反,即在SC8活性增加,但在SC124品种中降低。我们推测总淀粉含量及支链淀粉含量的下降可能与AGPase活性有关；GBSS活性增加与否与直链淀粉含量相对应；SSS和SBE活性任一酶的下降都有可能降低支链含量。从组织结构看,遮荫处理后的两品种淀粉充实度低,淀粉结构疏松,SC8单粒淀粉增多,结合淀粉含量和淀粉粒大小分析表明,遮荫处理使贮藏淀粉中有更多直径较小的淀粉粒,遮荫影响淀粉粒充实程度从而造成淀粉含量降低。
The accumulation, transportation, and conversion of sucrose affect the development of cassava "storage root" and starch accumulation from" source", and starch synthesis, which directly relates to the yield of starch. The period from 100 d to 200 d after planting, three high yield and far related cassava cultivars (SC124, SC8, and Arg7) were used for researches,' which contributed in the following aspects:the changes of root starch and its component accumulation characteristics; the changes of leaf sucrose content, reducing sugar content and enzyme activity involved in sucrose metabolism; the relation between root starch content and leaf growth, sucrose accumulation as well as key enzymes activity involved in sucrose metabolism; the changes of root sucrose content, reducing sugar content and root enzyme activity including in sucrose metabolism and starch synthesis; the relation between root starch content and sugar content along with enzymes activity including in sucrose metabolism and starch synthesis.
1. The characteristics of starch accumulation in cassava storage roots have been studied in the three cassava cultivars (SC124、SC8、Arg7) during root development from 100 d to 200 d after planting. The quantity of total starch per plant was continuously increased, but the starch content (%) was increased at intervals. The starch content (%) was significant higher in Arg7, among which it was Arg7>SC124>SC8. The content and rate of starch accumulation among the three cultivars was changed in the pattern of a single-peak curve during root mature processing, in which SC124 and SC8 was much earlier to reach the peak in content, and in the peak of rate was SC124>SC8>Arg7. Thus, the starch accumulation in SC124 and SC8 was faster than Arg7, but it lasted longer period in Arg7.The total quantity, content, accumulation rate of amylopectin was similar as those of total starch accumulation in the three cultivars. Between the cultivars, the average value of amylopectin content (%)in SC8 was much lower than other two cultivars, but the peak of accumulation rate and quantity of amylopectin was much slowly appeared in Arg7 than SC124 and SC8. At the level of P<0.05, amylose content(%) was not significantly different, but it was significantly different in amylopectin and total starch content (%) among SC124、SC8 and Arg7.
2. The enzymes involved in sucrose metabolism and starch synthesis have been investigated from "source" to" storage roots", and the relationship between enzymes and starch content of storage roots have been discussed. The results are as follows:AI activity was declined during root mature processing, and the activity of SPS and NI was changed in the pattern of a single-peak curve; The AI activity in Arg7 was much lower than that in SC8 or Arg7, and SPS and NI activity in SCI 24 was much higher than that in SC8 or Arg7; The activity of SPS was significantly correlated with the activity of NI, showing the coordinate reduation in controlling sucrose content of leaf; The activity of AI was significantly correlated with the activity of the reducing sugar content in leaves, but was negatively significantly correlated with the root starch content (%), it can be deduced that the lower activity of AI could produce less reducing sugar, which was propitious to sucrose transportation and root development.
The changes of AI and NI activity in storage roots are almost similar among three cassava cultivars, and both of them showed high activity in early developing stage. The peak of SS was much slowly appeared, but kept high level in the later developing stage. The activity of NI was significantly correlated with the AI activity in the storage roots, but was negatively significantly correlated with the root starch content, it can be deduced that the reducing sugar deprived from sucrose not only was used for substrate in starch synthase, but also play an important role in the root growth regulation, so that the starch content may declined slightly as a result of rapid swelling of root. The higher activity of AGPase was identified at two periods, and the first peak of AGPase activity arrived much earlier in SC8 than that in SC124 and Arg7, but the second peak arrived at the same period. The activity of soluble starch synthase (SSS), granule-bound starch synthase (GBSS) was changed in the pattern of a single-peak curve during root mature processing. The activity of AGPase and SBE in Arg7 cultivar with higher starch content was higher than that in SC124 and SC8. The content of total starch and amylopectin was significantly correlated with the activity of AGPase, but was not correlated with the activity of other enzymes (GBSS, SSS and SBE). The activity of AGPase may be used as standard to estimate the capability of starch accumulation in cassava storage roots.
3. The three cassava cultivars (SC124、SC8、Arg7) with far relationship and high-yielding were used for analysis of the isozyme electrophoresis. The correlation between isozyme locus, enzyme activities of SBE and AGPase, and starch content was investigated. The results showed that there were at least six isozyme loci in AGPase (AGPa、AGPb、AGPc、AGPd、AGPe、AGPf), which manifested genetic diversity among three cultivars. The locus of AGPe only appeared in Arg7, and it may closely relate with higher AGPase activity and starch content. SBE I manifested genetic diversity among three cultivars, while SBEⅡdid not exhibit genetic polymorphism, but showed different in activity among different varieties and root developing stages. The loci of SBEIc and SBEⅡmay provide much great contribution to SBE activity and amylopectin content.
4. By studying the starch development of the induced cassava storage roots in vitro of SC124, it indicated that in the early stage of starch grain development, a number of starch grains were developing in a closed amyloplast with membrane. With the maturation of amyloplast, the starch granules were released and many single granules were observed in induces cassava storage roots.
The storage roots of the three cassava cultivars (SC124、SC8、Arg7) in 120 and 160 d after planting have been scanned under SEM and TEM. Our observations by TEM confirm that the images sanned by SEM at 120 d were the amyloplasts with membrance envelope, and the morphology of starch such as momultidimensional were the starch granules at one amyloplast. On the 160 days after planting, the membrane envelop of mature amyloplast was disaggregated, and mature starch granules have been released from the amyloplasts. Both amyloplasts and starch granules can be observed in parenchyma. The starch granules in cassava mainly consist of many simple starch grains, few of them are compound starch granule. The frequency of the starch multigrain varies in different cultivars may be interrelated to starch quanlity. Our observations by SEM show that the size (the average diameters), pattern and arrangement are diversity among the three cassava cultivars:The average size of amyloplasts in SC124 are smallest, but their coefficient of variation and interspaces are larger; the granule size of amyloplasts in SC8 are relative uniform with smaller coefficient of variation and tight arrangement; however in Arg7, the granule size and coefficient of variation are smaller, while the interspaces are larger. SEM image showed that of the amyloplasts are stratiformly arranged in zylem parenchyma in SC124, sarciniformly arranged in SC8; and granularly arranged in Arg7. On the 160 days after planting, the starch granules were increased remarkablely.
Under TEM, two patterns of proliferation were observed:1, envelope dilates and invaginates to form new amyloplasts; 2, the amyloplast envelope buds to form the double membrane vesicle. The size, arrangement, the pattern and efficient of proliferation may maybe the immanent cause of character difference.
5. Two cassava cultivars SC124 and SC8 were used for the study of starch synthesis and accumulation in storage roots in response to shading light to reduce light intensity from the early stage (70 d after planting)to storage root filling stages (160 d after planting). The starch accumulation, starch structures and activities of the enzymes involved in starch synthesis have been examined at the developing stages of cassava storage roots. The results showed that the shading treatment (ST) has changed the distribution of the product from photosynthesis between the "source" and "storage" organs. The content of total starch and amylpectin was remarkably reduced, but the amylose content was significantly increased in comparison to the control (CK, no shading light). The ratio of amylose to amylopectin in ST conditions was significantly higher than that of CK. The activity of adenosine diphosphate glucose pyrophosphorylase (AGPase) in shading conditions was significantly lower than that . in control plants. The activity of GBSS was increased in SC8, but no obviously changed in SC124. The activity of SSS was increased in SC124, but was decreased in SC8, which was contrast to SBE; It was increased in SC8, but was decreased in SC124. It suggests that the low activity of AGPase resulted from shading light may relate to the lower total starch and amylopectin content. The GBSS activity was in response to the change of amylose content; and the any reduced activity of SSS, or SBE may result in lower amylose content. When the cassava was under shading conditions, the organization of starch structure became looser, the size of starch granule was significantly smaller, but the number of the single starch granule was obviously increased.
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