摘要
本研究选用我国50年代至90年代育成的小麦品种,研究了不同基因型小麦品种籽粒淀粉粒粒度分布特征及其差异；再以扬麦158为材料,研究了强势籽粒(小穗基部籽粒)和弱势籽粒(小穗顶部籽粒)间淀粉粒粒度分布特征的差异与机理；最后以扬麦158和扬麦11(轮回亲本是扬麦158)为材料,进行遮光处理,设置自然光照(S0)和遮去33%自然光照(S1)两个处理,研究遮光对小麦籽粒淀粉粒度分布特征的影响及其机理。研究结果表明,淀粉合成酶活性及其编码基因表达模式对小麦籽粒灌浆期淀粉粒粒度分布特征具有重要的调控作用。获得的主要结果如下：
1.小麦籽粒淀粉粒发育及其粒度分布动态
在花后4天,小麦籽粒中只有一种尺寸的淀粉粒群体,呈单峰分布,这些早期生成的淀粉粒将发育成为A型淀粉粒；在花后8天,淀粉粒粒度分布呈现出双峰的分布特征,最大粒径达到了20μm以上,新淀粉粒群体的出现表明了B型淀粉粒在此时期开始形成；在花后20天,另一种淀粉粒群体—C型淀粉粒开始形成,由此至小麦籽粒成熟期,淀粉粒粒度分布一直保持着三峰分布特征。因此,小麦籽粒成熟期淀粉粒呈现三峰的分布特征：>10μm的A型淀粉粒,2～10μm的B型淀粉粒以及<2μm的C型淀粉粒。
2.不同基因型小麦品种籽粒淀粉粒粒度分布差异
研究测定了我国50年代至90年代普通小麦品种籽粒淀粉粒粒度分布和组分含量,并比较了它们之间的差异和相关性。供试小麦品种成熟期的籽粒淀粉粒粒度分布呈现典型的三峰分布特征。不同年代的小麦品种籽粒中大小不同的淀粉粒群体体积存在显著差异,其中A型淀粉粒体积百分比随着年代的演化逐渐升高,而B型和C型淀粉粒体积百分比则不断下降。与此同时,小麦籽粒淀粉粒的表面积和数目在不同年代的小麦品种之间亦存在较大的差异。其中>10μm的淀粉粒群体体积百分比与直链淀粉和支链淀粉含量呈现出显著或极显著的正相关性,而支链淀粉与籽粒淀粉粒表面积和数目呈现显著的负相关,与直链淀粉含量相关性未达到显著水平。因此,研究结果表明不同年代小麦品种籽粒淀粉粒粒度分布特征和淀粉组分含量存在显著差异,同时这些差异在90年代与50年代品种的对比中均达到了显著水平。
3.小麦强弱势籽粒淀粉粒粒度分布差异
与弱势籽粒比较,小麦强势籽粒的直链淀粉、支链淀粉以及总淀粉含量更高。基于体积分布,A型、B型和C型淀粉粒的形成时间分别为花后4天、8天和20天,淀粉粒的平均粒径在花后16天达到最大,随后缓慢下降,而弱势籽粒淀粉粒的平均粒径在花后16天以后大于强势籽粒淀粉粒,众数粒径随着灌浆进程逐渐上升,最大粒径在花后16天达到了与成熟期相近的尺寸；基于数目分布,0.5～2μm之间的小淀粉粒群体在强势籽粒和弱势籽粒中均是主体部分,占总数目的90%以上。基于表面积分布,其灌浆期的变化动态趋势与体积分布相似,但量化趋势却与之相反,B型和C型淀粉粒在表面积分布中占主导地位。尽管如此,小麦强势籽粒淀粉粒的粒度分布演化进程要提前于弱势籽粒,弱势籽粒中淀粉粒的形成时期、淀粉粒的增大进程以及小淀粉粒在灌浆后期的发育等方面在时态性上表现相对滞后。
4.长期遮光对小麦籽粒淀粉粒粒度分布的影响
长期遮光降低了小麦支链淀粉的含量而对直链淀粉含量影响不明显,导致小麦籽粒总淀粉含量和支/直显著下降。长期遮光导致基于体积、表面积和数目的平均粒径下降。遮光处理条件下,在灌浆中期和后期小麦籽粒淀粉粒合成受到了抑制,造成了籽粒淀粉峰值粒径后移程度下降、最大粒径范围缩小以及灌浆后期小淀粉粒增殖与增大受抑制等。对比不同小麦品种的籽粒淀粉粒粒度分布对遮光的响应程度,扬麦158对遮光的抗性要好于扬麦11。
5.小麦籽粒淀粉粒粒度分布调控的生理机制
淀粉合成酶活性以及编码淀粉合成酶基因的表达水平在小麦灌浆期具有很强的时态性。可溶性淀粉合成酶和束缚态淀粉合成酶活性分别在花后20天和24天达到最大值,强势籽粒的淀粉合成酶活性要高于弱势籽粒。同时,SSS和GBSS活性的互作效应对大小淀粉粒的体积比例也表现出了明显的调控作用。编码淀粉合成酶亚型基因在小麦灌浆各时期的表达水平差异较大。SSⅠ在整个灌浆期均有所表达；SSⅡ和SSⅢ主要集中在灌浆早期至中期表达；GBSS I主要集中在灌浆中后期表达。淀粉合成酶编码基因的表达模式与淀粉酶活性在灌浆期的变化趋势相符,可见基因表达对淀粉合成的影响表现为其对淀粉合成酶活性的调控。因此,编码淀粉合成酶基因的表达水平对小麦籽粒淀粉粒粒度分布的调控是通过对淀粉合成酶活性的调控来实现的。
6.小麦籽粒淀粉粒粒度分布的调控途径
通过对比不同供试小麦品种和不同遮光处理下小麦籽粒淀粉粒粒度的差异,我们发现了可在目标育种和调优栽培应用中发挥重要作用的几个关键指标：1)调控小麦籽粒淀粉粒度分布的重要灌浆时期；2)大小淀粉粒在优质小麦籽粒中的粒径分界值和体积比例；3)大小淀粉粒因小麦品种不同可能对环境条件的变化产生不同的响应趋势；4)10μm的粒径分界线和40%的累积率分界线受到基因型的控制程度比环境条件的影响大。
综上所述,小麦籽粒淀粉粒粒度分布在不同基因型小麦品种、不同环境条件下以及不同粒位之间均存在显著差异,并受到淀粉合成酶活性及其编码基因表达模式的调控。可在大小淀粉粒群体(10μm)和灌浆中后期对小麦籽粒淀粉粒粒度分布实现有效调控。
Wheat cultivars released in the years from 1950s to 1990s were selected to evaluate the difference on starch granule size among diverse genotypes in the field experiment. The representative cultivars were used for the subsequent experiment. In addition, the most basal grain and the most distal grain on each spikelet of wheat cultivar Yangmai 158 were detached and identified as superior grain and inferior grain, respectively. The difference in starch granule size distribution was then compared between superior and inferior grains, the corresponding mechanism was also studied. Finally, wheat cultivar Yangmai 158 and Yangmai 11 (with Yangmai 158 as recurrent parent) were subjected to 33% less intercept radiation (S1, non-shading as control, SO) from jointing to maturity. Effect of shading on starch granule size distribution and its mechanism were then investigated. Based on the results here, the starch synthase activities and their coded gene expression were found to play important regulation roles in starch granule size distribution formation in wheat grain. Here are the main results.
1. Observed the development progress of starch granule size distribution in wheat endosperm during grain filling
Analysis of 4 DAA endosperm revealed a single population (destined to become the large A granules). At 8 DAA starch granule size exhibited a bimodal distribution with a maximum diameter more than 20μm. The new class of starch population suggested the initial formation of the small B-type granules. At 20 DAA, a new different class of starch granule population, defined as the C-type granules, was synthesized. From 24 to maturity, starch granules exhibited three distinct populations. At maturity three distinct size groups of starch granules were present: large type A granules with diameters greater than 10μm, type B granules with diameters between 2 and 10μm, and small type C granules with diameters less than 2μm.
2. Characterized the difference in starch granule size distribution in wheat endosperm among diverse genotypic cultivars Starch samples isolated from the wheat cultivars released in the years from 1950s to 1990s were examined. A clear trimodal distribution of granule sizes was shown by all starch samples. Highly significant differences were seen among the cultivars from different years for volume percentage of starch granules. Following with the years evolution, volume percentage of A-type granules increased and that of B- and C-type granules decreased. Starch granule surface area and number also differed within cultivars. Mean amylose and amylopectin content varied significantly among cultivars. Amylose and amylopectin content were positively correlated with the volume percentage of granules with diameters greater than 10μm. Amylopectin content was negatively correlated with surface area and number of starch granules. Results suggested that significant differences existed in granules size distribution and starch content between wheat cultivars from different years, especially between 1950s and 1990s.
3. Revealed the difference in starch granule size distribution in wheat endosperm between superior and inferior grain Compared with inferior grains, superior grains showed higher starch content as well as amylose and amylopectin. Based on volume, the formation of A-, B-, C-type starch granules initiated at 4,8,20 DAA, respectively. The mean diameter of starch granules achieved at 16 DAA in superior and inferior grains, and it was higher in inferior grains during later grain filling. The mode diameter of starch granules increased with grain filling process in superior grain, as well as inferior grains. The maximum diameter of starch granules increased slightly after 16 DAA, even becoming close to the terminative diameter at 20 DAA and not varying after this. Based on number, the small starch granules, between 0.5-2μm, were in the predominant position, composing above 90% of total number. Based on surface area, dynamic variation tendency of size distribution was similar to that based on volume, and the population of B-type and C-type starch granules was major component. However, the process of starch granule size distribution was faster in superior grain, and inferior grain had temporal delay in initiation and enlargement of granule at early stage and development of small granules at late stage during grain filling.
4. Evaluated the effect long-term shading on starch granule size distribution in wheat endosperm Long-term shading showed no effect on amylose content while significantly reduced amylopectin content, which resulted in the significant decrease in the ratio of amylopectin to amylase content in grain. Long-term shading also reduced the mean diameters by volume, number and surface area. Under the long-term shading conditions, the development of starch granule was suppressed during the mid to late grain filling stage, as a result of decline in peak diameter, reduced range of max diameter and enlargement of small granules. Based on the response of starch granule size distribution to shading, Yangmai 158 showed the better tolerance than Yangmai 11.
5. Explored the physiological mechanism for development of starch granule size distribution in wheat endosperm The activities of soluble and granule-bound starch synthases (designated SSS and GBSS) peaked on 20 and 24 DAA. Our results showed that the ratio of large to small starch granules in volume frequency percentage, at least partially, was related to the interplay between the SSS and GBSS activities. Genes encoding isoforms of starch synthases expressed at different grain filling period. SS I was generally expressed over the grain filling stage; the SS II and SS III were expressed over the early and mid grain filling stage, and the GBSS I was expressed during the mid to late grain filling stage. In addition, the activities of starch synthases and expression of starch synthase encoding genes temporally shared the dynamics of the starch granule size distribution. It was concluded that the starch granule size distribution in the grains of wheat was associated with the activities of the starch synthases, which were regulated at the transcription levels of the genes encoding the starch synthases.
6. Debated the regulation approached for starch granule size distribution in wheat endosperm In this study, depending on the variances in starch size distribution from cultivars and treatments, some key indices for accelerating the progress of target breeding and better cultivation were found as below:1) the fitting time for regulation of starch size distribution during grain filling; 2) the bound of diameter and volume ratio between the large and small starch granules in high quality wheat grains; 3) the details of changes in size distribution of the populations of large and small starch granules under controlled conditions; 4) more effects of genotype than growth environment on the indices which the bound of diameter in 10μm and volume accumulation rate in 40%.
In conclusion, the variances in starch granule size distribution follow with diverse genotype, ecological environment and grain position. However, size distribution was regulated by starch synthase activities and their coding gene expression. It was possible that large and small starch granule population (cutoff at 10μm) could be regulated during mid and late grain filling stage.
引文
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