玉米叶片叶绿素含量的全基因组关联性分析

植物叶片的叶绿素含量与叶片的光合作用效率和产量潜力紧密相关,因此是作物的一个重要生理指标。但是,目前已克隆的控制叶绿素含量的基因大多来自拟南芥和水稻,尚不清楚玉米自然群体中哪些基因控制叶片叶绿素含量的变异。本研究发现玉米苗期第一片叶片的叶绿素含量和吐丝期穗位叶的叶绿素含量高度相关,且与后者相比具有更高的遗传力。进一步分析了287份玉米自交系的第一片叶的叶绿素含量,利用558 269个单核苷酸多态性分子标记进行全基因组关联性分析,获得9个显著关联SNP位点和16个候选基因。通过候选基因的序列分析和功能预测,发现两个可能和叶绿素含量相关的基因,包括拟南芥Tic22基因的同源基因和水稻衰老相关基因SAG12的同源基因。     

杂交水稻开花结实期间叶片衰老

在杂交水稻开花期间,通过剪去穗部部分枝梗和叶片等处理,形成不同的库源比,叶片衰老的生理生化指标变化和结实率的测定表明,杂交水稻组合的库源矛盾大,叶片衰老快;但谷粒数与旗叶面积比不一定高,降低库源比,能明显减缓杂交水稻叶片中蛋白质、叶绿素含量及倒3、4、5片叶叶绿素含量的平均值与旗叶叶绿素含量的比率的下降和丙二醇含量的升高。这些都说明杂交水稻叶片功能早衰与其库源矛盾较大有关。     

水稻叶片衰老过程中氨肽酶活性的变化

水稻叶片中存在着氨肽酶,其最适反应pH和最适反应温度分别为8.2℃和40℃,酶促反应的产物量在最初30min内与时间呈直线相关。 水稻叶片衰老过程中叶绿素和蛋白质含量下降,而氨肽酶比活上升;用植物激素延缓或促进叶片衰老蛋白质降解的同时也抑制或促进了氨肽酶比活的上升,说明氨肽酶在水稻叶片衰老蛋白质降解过程中起一定的作用。根据水稻叶片衰老过程中大分子化合物和叶片外部形态的变化,可将叶片衰老过程划分为缓衰期、急衰期和竭衰期。     

杂交水稻及其亲本光合特性的研究 Ⅰ．功能叶片叶绿素含量、叶绿素－蛋白复合物及诱导荧光动力学

研究了杂交水稻青优１５９和广优四号及其亲本功能叶片的光合速率、叶绿素含量、叶绿素－蛋白复合物及诱导荧光动力学特性．这二个杂交水稻的光合速率分别高于其亲本,其超亲优势分别为１８．７２％和１８．２％,平均优势分别为２９．６％和２６．２％．杂交水稻功能叶片的叶绿素－蛋白复合物在６５０ｎｍ和６７５ｎｍ处光密度扫描峰面积具有明显的杂种优势,并与光合速率之间有较密切的正相关关系;叶绿素诱导荧光动力学特征参数Ｆｖ／Ｆｍ和Ｆｖ／Ｆｏ比值超过其高值亲本,具有杂种优势;杂交水稻功能叶片的叶绿素含量没有明显的杂种优势,光合速率与叶绿素含量之间没有明显的相关关系。而杂交水稻功能叶片的叶绿素ａ／ｂ比值均低于其亲本,并且叶绿素ａ／ｂ比值与光合速率呈较为密切负相关．     

杂交水稻开花结实期间叶片衰老

在杂交水秀开花期间,通过剪却穗部部分枝梗和叶片等处理,形成不同的库源比,叶片衰老的生理生化指标变化和结实率的测定表明,杂交水稻组合的库源矛盾大,叶片衰老快;但谷粒数与旗叶面积比不一定高,降低库源比,能明显减缓杂交水稻叶片中蛋白质,叶绿素含量及倒３,４,５片叶叶绿素含量的平均值和旗叶叶绿素含量的比率的下降和丙二醛含量的升高。     

水稻冠层高光谱特征变量与叶片叶绿素含量的相关性研究

为了探讨水稻冠层光谱对叶片叶绿素含量的响应规律,以双季早稻为材料,设置不同施氮量处理的田间试验,测定水稻冠层光谱和叶片叶绿素含量,计算基于冠层反射光谱的特征变量,研究水稻冠层高光谱特征变量与叶片叶绿素含量之间的关系。结果表明:施用氮肥对反射光谱有明显的影响,在可见光范围内,不施氮处理的反射率高于施氮处理,尤其在波长550 nm左右的绿峰处显著增加,在近红外区反射率随施氮量的增加而增加;与叶绿素含量相关性较好的光谱位置参数是红边位置和红谷反射率,随着叶绿素含量的增加,红谷反射率降低,红边位置向长波方向移动;比值植被指数R800/R550、R750/R553和R990/R553,以及色素比值指数PSSRa、PSSRb与chla、chlb、chl(a+b)呈极显著正相关,可以作为水稻冠层叶片叶绿素监测的特征变量。     

水稻叶片衰老相关基因的研究进展

水稻叶片的衰老是制约杂交稻产量提高的主要因素之一,有数据表明水稻籽粒灌浆所需营养物质的60%～80%来自叶片的光合作用,实践证明叶片每推迟1天衰老,产量可提高产1%左右.因此,对叶片衰老的形态、生理生化及其相关分子机理等进行研究具有重要的现实意义.近年来水稻叶片衰老的相关研究表明,叶片的衰老是一个受众多因素影响的复杂过程,在这个过程中叶片发生了巨大的形态与生理生化变化,而这些变化均离不开基因的调控作用.大量实验结果表明:在衰老过程中,叶片细胞有选择地启动或增强某些基因(叶片衰老相关基因)的表达,而关闭或减弱另一些基因(衰老下调基因)的表达,由此来调控叶片衰老的进程.目前研究者已在研究衰老突变体等相关的材料中发现了许多与水稻叶片衰老有关的基因.本文重点概述了近年来水稻叶片衰老相关基因的研究状况,并对未来研究方向等问题做了思考与探讨,以期能为开展进一步的研究工作提供参考.     

测定水稻叶片叶绿素含量的混合液提取法

水稻叶片叶绿素含量与光合速率、作物生长率有一定相关。科研上常测定叶绿素含量以表征水稻生长状态,生产上也往往根据叶色变化作为看苗诊断和采取肥水措施的重要指标。     

4PU-30对水稻叶片衰老与内源激素的调控

4PU－30能显著地延缓水稻叶片衰老。根据叶片衰老过程中内源激素含量的变化,可明确减缓水稻叶片衰老期间内源ABA含量的增加和内源ZRs、GAs和IAA含量的减少,使叶片中保持有较低水平的ABA与较高水平的ZRs、GAs和IAA,是4PU－30延缓水稻叶片衰老的主要调控机理。     

苗期玉米叶片碳氮平衡与干旱诱导的叶片衰老之关系

为了探究干旱诱导的碳氮平衡破坏与干旱诱导的叶片衰老之间的关系,该实验以8个在干旱胁迫下叶片衰老进程有明显差异的玉米品种为实验材料,采用PEG模拟干旱处理,通过测定光合速率、叶绿素含量和叶绿素荧光参数等叶片衰老指标以及非结构性碳水化合物(可溶性糖、淀粉)和全氮含量等变化,分析玉米中干旱诱导的叶片衰老与叶片中碳氮平衡(碳氮比)之间的关系。结果显示:(1)干旱胁迫下,8个玉米品种叶片净光合速率受到严重抑制,Fv/Fm大幅下降,叶绿素含量显著降低,说明干旱诱导了玉米叶片的衰老;(2)干旱诱导玉米叶片衰老的同时,8个玉米品种的叶片中可溶性糖含量显著升高,淀粉含量小幅上升,全氮含量大幅降低,碳氮比显著升高,碳氮平衡遭到了破坏;(3)8个玉米品种叶片的叶绿素含量与非结构性碳水化合物含量以及碳氮比呈极显著负相关关系,与全氮含量呈极显著正相关关系。因此,碳氮代谢与干旱诱导的叶片衰老紧密联系,碳氮平衡可能参与了干旱诱导的叶片衰老调控。     

Are contents of Rubisco, soluble protein and nitrogen in flag leaves of rice controlled by the same genetics?

Genetic relations among the contents of Rubisco, soluble protein and total leaf nitrogen (N) in leaves of rice (Oryza sativa L.) were studied by quantitative trait loci (QTL) analysis with a population of backcross inbred lines (BILs) of japonica Nipponbarexindica Kasalath. The ratio of Rubisco to total leaf N in leaves is the main target in improving photosynthetic N-use efficiency in plants. QTLs controlling Rubisco content were not detected near QTLs for total leaf N content. These results indicate that contents of Rubisco and total leaf N are controlled by different genetics. QTLs that controlled the ratio of Rubisco to total leaf N (CORNs) were detected. These results suggest that some mechanism(s) may be involved in determining this ratio, while the contents of Rubisco and total leaf N are controlled in other ways. In elite BILs, the ratios of Rubisco to total leaf N were higher than those of both parents. These results suggest a good possibility of improving N-use efficiency by CORNs in cultivated rice. A QTL controlling Rubisco content was mapped near a QTL for soluble protein content on chromosome 8 at 5 d after heading and on chromosome 9 at 25 d. In each chromosome region, the peaks of both QTLs overlapped accurately, giving a high possibility of pleiotropic effects by the same genes. Different QTLs controlling soluble protein or Rubisco were detected from those detected at 5 d or 25 d after heading. This suggests that these traits are genetically controlled depending on the growth stages of leaves.     

QTL analysis to study the association between leaf size and abscisic acid accumulation in droughted rice leaves and comparisons across cereals

Plants accumulate abscisic acid (ABA) under droughted conditions. Genetic variation in the accumulation of ABA in detached and partially dehydrated leaves of rice has previously been reported, and this was found to be associated with variation in leaf size (smaller leaves made more ABA). Correlation analysis failed to distinguish clearly between a causal relationship between the two traits and close genetic linkage between loci controlling the traits. Here we present a detailed genetic analysis of ABA accumulation in detached and partially dehydrated rice leaves, using a population of F₂ plants generated from the lowland × upland cross IR20 (high-ABA) × 63-83 (low-ABA) which was mapped with RFLP and AFLP markers. Several highly significant quantitative trait loci (QTLs) for ABA accumulation and leaf weight were identified. Only one of the minor QTLs for ABA accumulation (accounting for only 4% of the phenotypic variance) was coincident with any QTLs for leaf size such that the high-ABA allele was associated with smaller leaves. This analysis, therefore, showed that the association previously found between ABA accumulation and leaf size was probably largely due to genetic linkage and not to a direct effect of leaf size on ABA accumulation or vice versa. Because of the importance of ABA accumulation in regulating responses of plants to drought stress and the effects of plant size on the rate of development of stress, QTLs for drought-induced ABA accumulation, leaf size and tiller number were compared between rice and wheat. In particular, a possible location in rice was sought for a homoeologue of the major wheat vernalization responsive gene, Vrn1, as this gene is also associated with major effects on leaf size, tiller number and ABA accumulation in wheat. The likelihood of homoeologous loci regulating ABA accumulation, leaf size and tiller number in the two crops is discussed.     

利用粳稻染色体片段置换系群体检测水稻抗亚铁毒胁迫有关性状QTL

潜育性水稻田广泛分布于中国、斯里兰卡、印度、印度尼西亚、塞拉里昂、利比亚、尼日利亚、哥伦比亚和菲律宾等国,其中我国南方稻区就有近700万公顷低产潜育性水稻田。该类水稻田还原性强,矿质营养失调,尤以Fe^2 过量积累,对水稻生长发育产生不良的逆境胁迫作用。培育抗亚铁毒的水稻品种是简便、经济有效地提高稻谷产量的重要途径之一。该文利用由粳稻品种Asominori与籼稻品种IR24杂交衍生的Asominori染色体片段置换系(Chromosome Segment Substitution Lines,CSSLs)群体为材料,检测与抗亚铁毒胁迫有关性状QTL。共检测到与抗亚铁毒胁迫有关性状QTL14个,各QTL的LOD值为2．72～6．63。其中检测到与抗亚铁毒胁迫直接有关的性状叶片棕色斑点指数QTL3个,分别位于第3、9、11染色体C515～XNpb279、R2638～C1263和G1465～C950之间,对应的贡献率分别为16．45％、11．16％和28．02％;与其他已发表的定位结果比较发现,位于第三染色体C515～XNpb279间控制叶片棕色斑点指数的QTL与水稻功能图谱上控制叶绿素含量的QTL的位置一致;表明在亚铁毒胁迫条件下,水稻在其叶片表面出现棕色斑点,叶片衰老,产生一些叶绿素降解物或衍生物,以提高叶片细胞对亚铁等重金属毒害的耐受力。另外,在第11染色体G1465～C950之间检测到了控制叶片棕色斑点指数、茎干重和根干重QTL1个,为主效QTL。在第6染色体XNpb386～XNpb342之间检测到控制茎干重、株高、根长和根干重QTL1个,是否与水稻抗亚铁毒有关需要进一步研究。本研究旨在通过定位与抗亚铁毒有关的QTL,借助与之紧密连锁的分子标记有效地聚合这些QTL,培育出抗亚铁毒性强的水稻新种质材料。     

Mapping QTLs associated with drought avoidance in upland rice

The identification of molecular markers linked to genes controlling drought resistance factors in rice is a necessary step to improve breeding efficiency for this complex trait. QTLs controlling drought avoidance mechanisms were analyzed in a doubled-haploid population of rice. Three trials with different drought stress intensities were carried out in two sites. Leaf rolling, leaf drying, relative water content of leaves and relative growth rate under water stress were measured on 105 doubled haploid lines in two trials and on a sub-sample of 85 lines in the third one. Using composite interval mapping with a LOD threshold of 2.5, the total number of QTLs detected in all trials combined was 11 for leaf rolling, 10 for leaf drying, 11 for relative water content and 10 for relative growth rate under stress. Some of these QTLs were common across traits. Among the eleven possible QTLs for leaf rolling, three QTLs (on chromosomes 1, 5 and 9) were common across the three trials and four additional QTLs (on chromosomes 3, 4 and 9) were common across two trials. One QTL on chromosome 4 for leaf drying and one QTL on chromosome 1 for relative water content were common across two trials while no common QTL was identified for relative growth rate under stress. Some of the QTLs detected for leaf rolling, leaf drying and relative water content mapped in the same places as QTLs controlling root morphology, which were identified in a previous study involving the same population. Some QTL identified here were also located similarly with other QTLs for leaf rolling as reported from other populations. This study may help to chose the best segments for introgression into rice varieties and improvement of their drought resistance.     

Role of ethylene in the senescence of detached rice leaves

The role of ethylene in the senescence of detached rice leaves in relation to their changes in 1-aminocyclopropane-1-carboxylic acid (ACC) content and ethylene production was studied. In freshly excised rice leaf segments, ACC level and ethylene production rates were very low. Following incubation, the rates of ethylene production increased and reached a maximum in 12 h, and subsequently declined. The rise of ethylene production was associated with a 20- to 30-fold increase in ACC level.

Ethylene seems to be involved in the regulation of the senescence of detached rice leaves. This conclusion was based on the observations that (a) maximum ethylene production preceded chlorophyll degradation, (b) ACC application promoted chlorophyll degradation, (c) inhibitors of ethylene production and ethylene action retarded chlorophyll degradation, and (d) various treatments such as light, cycloheximide, α,α-dipyridyl, Ni²⁺, and cold temperature, which retarded chlorophyll degradation, also inhibited ethylene production.

Abscisic acid promoted senescence but significantly decreased ethylene production, whereas benzyladenine retarded senescence but promoted ethylene production. This is interpreted to indicate that abscisic acid treatment increased the tissue sensitivity to ethylene, whereas benzyladenine treatment decreased it.

     

Dissecting a wheat QTL for yield present in a range of environments: from the QTL to candidate genes

Previous studies with 95 bread wheat doubled haploid lines (DHLs) from the cross Chinese Spring (CS)xSQ1 trialled over 24 yearxtreatmentxlocations identified major yield quantitative trait loci (QTLs) in homoeologous locations on 7AL and 7BL, expressed mainly under stressed and non-stressed conditions, respectively. SQ1 and CS contributed alleles increasing yield on 7AL and 7BL, respectively. The yield component most strongly associated with these QTLs was grains per ear. Additional results which focus on the 7AL yield QTL are presented here. Trials monitoring agronomic, morphological, physiological, and anatomical traits revealed that the 7AL yield QTL was not associated with differences in flowering time or plant height, but with significant differences in biomass at maturity and anthesis, biomass per tiller, and biomass during tillering. In some trials, flag leaf chlorophyll content and leaf width at tillering were also associated with the QTL. Thus, it is likely that the yield gene(s) on 7AL affects plant productivity. Near-isogenic lines (NILs) for the 7AL yield QTL with CS or SQ1 alleles in an SQ1 background showed the SQ1 allele to be associated with >20% higher yield per ear, significantly higher flag leaf chlorophyll content, and wider flag leaves. Epidermal cell width and distance between leaf vascular bundles did not differ significantly between NILs, so the yield-associated gene may influence the number of cell files across the leaf through effects on cell division. Interestingly, comparative mapping with rice identified AINTEGUMENTA and G-protein subunit genes affecting lateral cell division at locations homologous to the wheat 7AL yield QTL.     

Analysis of quantitative trait loci affecting chlorophyll content of rice leaves in a double haploid population and two backcross populations

Chlorophyll content, one of the most important physiological parameters related to plant photosynthesis, is usually used to predict yield potential. To map the quantitative trait loci (QTLs) underlying the chlorophyll content of rice leaves, a double haploid (DH) population was developed from an indica/japonica (Zhenshan 97/Wuyujing 2) crossing and two backcross populations were established subsequently by backcrossing DH lines with each of their parents. The contents of chlorophyll a and chlorophyll b were determined by using a spectrophotometer to directly measure the leaf chlorophyll extracts. To determine the leaf chlorophyll retention along with maturation, all measurements were performed on the day of heading and were repeated 30 days later. A total of 60 QTLs were resolved for all the traits using these three populations. These QTLs were distributed on 10 rice chromosomes, except chromosomes 5 and 10; the closer the traits, the more clustering of the QTLs residing on common rice chromosomal regions. In general, the majority of QTLs that specify chlorophyll a content also play a role in determining chlorophyll b content. Strangely, chlorophyll content in this study was found mostly to be lacking or to have a negative correlation with yield. In both backcross F₁ populations, overdominant (or underdominant) loci were more important than complete or partially dominant loci for main-effect QTLs and epistatic QTLs, thereby supporting previous findings that overdominant effects are the primary genetic basis for depression in inbreeding and heterosis in rice.     

水稻叶绿素含量的QTL定位

利用由两个籼稻品种Acc8558和H359杂交构建的一个包含131个株系(F19)的重组自交系群体,及其相应的包含147个RFLP和78个AFLP标记的遗传图谱,采用多性状复合区间定位方法,对控制水稻叶绿素含量的QTL进行了定位分析。对叶绿素a和叶绿素b含量各检测到6个QTL,其中5个QTL在两性状问是相同的。这些QTL主要分布在第1和第4染色体上,因此这两条染色体对叶绿素含量是重要的。QTL qChlAlc／qChlBlb(二者位置相同)在4个观测时期均表现较大效应,且在最后的剑叶期贡献最大,因此对叶绿素含量最为重要。另两个QTL(qCh-LA4a／qChlB4a和qChlA4b／qChlB4b)只在第2次观测时期效应显著,表明它们只在特定发育阶段发挥作用。     

水稻叶绿素含量动态QTL分析

为剖析水稻不同生育时期叶绿素含量的变化动态及遗传机制,以‘Sasanishiki’（粳稻）、‘Habataki’（籼稻）及其杂交衍生的85个回交重组自交系（BILs）群体为材料,对控制水稻叶片叶绿素含量的数量性状基因位点（QTL）变化动态进行了分析。共检测到39个QTL,包括26个非条件QTL和13个条件QTL,分布在除第7和第11号染色体以外的10条染色体上,平均每个时期检测到3．25个非条件QTL。其中生育前期和生育中后期检测到的QTL位点较少,仅为1—3个;在生育中期（盛期）检测到的QTL位点相对较多,一般为4～5个。在生育期的中期和后期均能在第1和2号染色体上检测到控制叶绿素含量的QTL,并且这些QTL位点在1和2号染色体上呈现聚集现象。本研究同时发现了一些新的QTL位点,这些QTL将有助于我们更全面地了解叶绿素在不同发育时期的遗传基础。     

Analysis of quantitative trait loci underlying the traits related to chlorophyll content of the flag leaf in rice

A population of 117 doubled haploid (DH) lines derived from the cross of Zhaiyeqing 8 (indica) x Jingxi 17 (japonica) was employed to map quantitative trait loci (QTL) underlying four physiological traits related to chlorophyll contents of the flag leaf. There were significantly positive correlations among chlorophyll a, chlorophyll b and chlorophyll a+ b content. Chlorophyll a/b ratio was significantly negatively correlated with chlorophyll b content. These four traits were normally distributed with transgressive segregation, suggesting that they were controlled by multiple minor genes. A total of 11 QTLs were detected for the four traits and they lay on six chromosomes. Each of them explained 9.2%-19.6% of the phenotypic variations, respectively. Of these, two QTLs controlling chlorophyll a content were mapped on chromosomes 2 and 5; four QTLs underlying chlorophyll b content were mapped on chromosomes 2, 3, 5 and 9; three QTLs underlying chlorophyll a+b amount were mapped on chromosomes 3, 5 and 9; two QTLs under-lying chlorophyll a/b ratio were mapped on chromosomes 6 and 1 1. The intrinsic relationship among the four traits and the practical implication in rice breeding are discussed.