一个化学诱变的小麦斑点叶突变体的生理和遗传分析

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杜丽芬, 李明飞, 刘录祥, 王超杰, 刘洋, 许喜堂, 邹淑芳, 谢彦周, 王成社. 一个化学诱变的小麦斑点叶突变体的生理和遗传分析. 作物学报, 2014, 40(6): 1020-1026[DU Li-Fen, LI Ming-Fei, LIU Lu-Xiang, WANG Chao-Jie, LIU Yang, XU Xi-Tang, ZOU Shu-Fang, XIE Yan-Zhou, WANG Cheng-She. Physiological Characteristics and Genetic Analysis on a Spotted-leaf Wheat Derived from Chemical Mutation. Acta Agron Sin, 2014, 40(6): 1020-1026] 复制到剪切板

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一个化学诱变的小麦斑点叶突变体的生理和遗传分析

杜丽芬^1,^*, 李明飞^1,^*, 刘录祥², 王超杰¹, 刘洋¹, 许喜堂¹, 邹淑芳¹, 谢彦周^1,^*, 王成社^1,^*

¹西北农林科技大学旱区作物逆境生物学国家重点实验室 / 农学院, 陕西杨凌 712100

²中国农业科学院作物科学研究所, 北京100081

^* 通讯作者(Corresponding authors): 谢彦周, E-mail:xieyanzhou@gmail.com; 王成社, E-mail:wangcs2008@126.com

第一作者联系方式: E-mail:dulifen617@163.com

^** 对本文有相同贡献.

收稿日期:2014-03-04

基金资助:本研究由国家自然科学基金项目(31101139), 国家高技术研究发展计划(863计划)项目(2012AA101202), 陕西省统筹项目(2011KTZB02-01-01)和西北农林科技大学博士科研启动经费(2010BSJJ035)资助;

摘要

通过EMS诱变普通小麦品系H261, 获得一个稳定遗传的斑点叶突变体LF2010。自然条件下, 该突变体在三叶期叶片基部第一片叶最先出现黄色斑点, 随着植株生长从基部老叶片到上部新生叶片依次出现黄斑,最后叶片上斑点逐渐增多并扩散到全片叶、叶鞘、颖壳和麦芒。斑点部位不存在细胞死亡, 斑点性状的表达受光照和温度诱导, 突变体的色素含量、光合速率随着斑点的出现而显著下降。突变体的株高、有效穗数、单株产量、穗长、结实率和旗叶长等农艺性状显著下降, 但是千粒重和旗叶宽却与野生型无差异。将突变体与正常绿色品系杂交, 对其F₁、F₂和BC₁代的遗传分析表明, LF2010的突变性状由1对隐性核基因控制。

关键词: 斑点叶突变; 光合色素; 光合特性; 农艺性状; 遗传分析

Physiological Characteristics and Genetic Analysis on a Spotted-leaf Wheat Derived from Chemical Mutation

DU Li-Fen^1,^**, LI Ming-Fei^1,^**, LIU Lu-Xiang², WANG Chao-Jie¹, LIU Yang¹, XU Xi-Tang¹, ZOU Shu-Fang¹, XIE Yan-Zhou^1,^*, WANG Cheng-She^1,^*

¹ State Key Laboratory of Crop Stress Biology in Arid Areas / College of Agronomy, Northwest A&F University, Yangling 712100, China

² Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081, China

Fund:

Abstract

A light and temperature affected spotted-leaf mutant LF2010 (Triticum aestivum L.) was obtained from a mutagenic treatment with EMS on a wheat line H261. The bright yellow spots first appear on the primary leaf at three leaves stage, then covered later leaves and leaf sheaths of the whole plant including tissues of spike. The mutant plants differed from normal plants in terms of total chlorophyll content and net photosynthetic rate once symptom appeared on the leaves. The plant height, spike length, spike number per plant, grain weight per plant, grain number per spike, seed setting rate, and flag leaf length were lower in the mutant than in the wild type. However, the 1000-grain weight and flag leaf width of the mutant had no significant difference with those of the wild type. Genetic analysis based on F₁, F₂, and BC₁ populations revealed that the yellow spot trait was controlled by a single recessive gene.

Keyword: Spotted-leaf mutant; Photosynthetic pigments; Photosynthetic characteristics; Agronomic traits; Inheritance

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0 引言

叶色突变是普遍存在于各种植物的一类突变类型。根据不同表型的叶色特点将其划分成3类, 一是单色突变, 即失绿的叶片只表现黄、白或淡绿等其中的一种颜色; 二是杂色突变, 即叶片上有2~3种颜色, 如黄白、白绿、绿白和斑点等; 三是阶段性失绿突变体, 即在个体发育的某个阶段叶色不正常^{[ 1]}。植物成色过程十分复杂, 这就决定了叶色突变原因的多变性和突变类型的多样性^{[ 2, 3, 4, 5]}, 因此, 收集和研究种类更多或独特的叶色突变体不仅对高等植物光合作用、色素合成、基因表达调控等基础研究有重要理论意义, 还可为作物高光效育种提供有利依据。

常见的叶色突变类型主要为单色突变, 如黄化、白化、浅绿等, 通常色素含量表现不同程度降低^{[ 6, 7, 8, 9]}。斑点突变是一种特殊的杂色突变类型, 受到广泛关注, 自20世纪90年代以来, 已在玉米、水稻、大麦、大豆、花生、拟南芥中报道了大量斑点突变, 并相继进行了深入研究^{[ 10, 11, 12, 13, 14]}。斑点突变大致可以分为两类, 一类是斑点部位伴随着细胞坏死, 称为类病变突变; 另一类是普通斑点突变, 不存在坏死症状而纯粹为叶片颜色的变化^{[ 15]}。多数斑点突变为类病变突变体, 因此常用于抗病性研究^{[ 15, 16]}, 而对于植物色素代谢和光合特性相关的斑点突变鲜有报道, Les22是已较深入研究的玉米突变体, 其突变基因编码卟啉代谢途径中合成叶绿素的关键酶尿卟啉原脱氢酶(UROD)^{[ 17]}。

在小麦中也相继报道了一些斑点突变体的生理和遗传等研究。Nair等^{[ 18]}利用化学诱变得到的显性斑点突变体C591(M8), 在斑点出现初期, 突变体的叶绿素含量与野生型差异不显著, 而当斑点布满整个叶片时, 突变体的色素含量较野生型明显下降, 但该突变性状对产量性状影响不显著。Kamlofski等^{[ 19, 20]}对HLP (hypersensitive-like phenotype)斑点突变体的研究发现, 斑点对产量性状同样没有显著影响, 并且HLP突变体成株期抗病性增强。Luo等^{[ 21]}研究发现, 小麦斑点突变体AIM9受1对隐性基因控制, 其色素含量、净光合速率以及单株产量较野生型材料明显下降。最近, 个别小麦斑点突变基因已被定位到染色体上^{[ 22, 23]}。然而, 我们对小麦斑点突变体的研究仍然不深入, 只有更多地鉴定和研究斑点突变体, 才能全面解析小麦斑点突变形成的机制。本文拟初步研究普通斑点突变体LF2010, 以揭示其生理特性和遗传规律, 为进一步研究斑点叶突变体及其基因定位打下基础。

1 材料与方法

1.1 材料

斑点叶突变体LF2010是小麦品系H261经过EMS (ethane methyl sulfonate)诱变获得的, 经过连续5代自交后已遗传稳定。斑点叶突变体LF2010、正常叶色小麦材料中国春和H261作为杂交亲本, 用于构建遗传分析群体。

1.2 表型鉴定

2012年10月突变体LF2010和野生型H261播种于西北农林科技大学试验田, 4行区, 行长1 m, 株距5.0 cm, 种植管理与一般大田相同。自种子出苗起, 全生育期观察突变体的叶色变化过程, 同时随机选取10株突变体LF2010和野生型H261, 考察株高、单株有效穗数、单株产量、千粒重、穗粒数、穗长、结实率和旗叶长宽等重要农艺性状。

1.3 突变体表型的环境敏感性鉴定

抽穗期选取突变体植株上尚未出现黄色斑点的旗叶, 将其中部用1 cm宽的锡箔纸遮光处理, 2周后观察叶色变化。为测定影响斑点出现的温度值, 2013年7月10日于人工气候箱进行盆栽, 温度梯度为20、15、10和5℃; 光周期12 h光/12 h暗, 光照强度150 μmol m^-2 s^-1; 相对湿度65%。随机选取LF2010和原始亲本种子各40粒, 每盆10粒, 室温催芽。将幼苗移至20℃使其生长一致, 至第2片叶长出后, 选4盆幼苗分别放置于不同温度的人工气候箱, 其他培养条件同上。当第4片叶长出后观察叶片颜色变化。

1.4 锥虫蓝染色法检测突变体叶片斑点部位的细胞活性

锥虫蓝染色是检测植物细胞死亡的常用方法之一, 坏死的细胞由于细胞膜遭到破坏会被染成深蓝色^{[ 24]}。参考马健阳等^{[ 25]}的方法检测突变体叶片的细胞死亡, 略有改动。当人工气候箱中幼苗长至五叶期的时, 随机选取已经表现斑点性状的突变体和野生型第1叶片, 浸泡在煮沸的锥虫蓝染液^{[ 26]}中, 染色7 min, 室温放置14 h, 再用2.5 mg mL^-1水合氯醛(10 mL水中加入25 g水合氯醛)中脱色3~4 d, 观察叶片是否被染成蓝色并照相。用50%甘油保存叶片。实验重复3次。

1.5 光合色素含量和光合参数测定方法

根据旗叶上斑点出现的时期分为出现斑点前、开始出现斑点和布满斑点3个阶段。孕穗期取旗叶, 制成直径5 mm叶圆片, 用80%丙酮提取色素。用分光光度计分别测定叶绿素a和叶绿素b及类胡萝卜素的含量^{[ 27]}。重复测定3次, 取平均值计算小麦单位叶面积的色素含量。

同一时期随机选取LF2010和H261长势一致的6个单株, 利用LI-6400型便携式光合作用测定仪测定旗叶的净光合速率( P_n)、气孔导度( G_s)、胞间二氧化碳浓度( C_i)和蒸腾速率( T_r)。

1.6 遗传分析

2011年5月至2012年11月, 将突变体LF2010分别与野生型H261和中国春杂交, 再自交或回交获得F₁、F₂和BC₁群体。观察苗期(三叶期之后)各群体性状的分离比。用SAS 8.0软件进行统计分析和卡方检验。

2 结果与分析

2.1 突变体的形态特征及主要农艺性状表现

田间观察发现, 三叶期前突变体与野生型无明显差异; 三叶期基部第1片叶开始出现黄色斑点后, 黄斑依次出现在第2和第3叶片上(图1-a, b); 越冬期随着温度进一步降低, 部分叶片保持正常绿色(图1-c); 次年春季随着温度升高, 冬前保持绿色的叶片和新生叶片逐渐出现黄斑, 并且黄斑随植株生长蔓延到每个叶片, 甚至到达叶鞘、颖壳和麦芒(图1-e, g, h), 整个蔓延过程具有从基部向顶部的方向性。

叶色失绿导致突变体植株矮小, 生长滞后(图1-d, e, f), 显著影响植株的大部分农艺性状, 如株高、有效穗数、单株产量、穗粒数、穗长、结实率、旗叶长比野生型下降19%~61%, 但千粒重和旗叶宽与野生型差异不大(表1)。

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图1 突变体LF2010与野生型小麦的表型比较a: 苗期野生型; b:苗期突变体; c: 突变体, 箭头示保持绿色叶片; d: 灌浆期野生型; e: 灌浆期突变体; f: 田间生长的野生型(左)和突变体(右); g: 突变体(上)和野生型(下)的叶鞘颜色; h: 突变体(上)和野生型(下)的叶片颜色; i: 突变体部分遮光后(上)和遮光前(下)叶片。Fig. 1 Phenotype of mutant LF2010 and its wild-type wheata: wild type seedling; b: mutant seedling; c: mutant. The arrow shows the keep-greening leaf; d: Wild type plant at grain-filling stage; e: mutant plant at grain-filling stage; f: wild type (left) and mutant (right) growing in the field; g: sheath color in mutant (upper) and wild type (lower); h: leaf color in mutant (upper) and wild type (lower); i: mutant leaf after partially shading (upper) and before shading (lower).

表1 突变体LF2010及其野生型H261的主要农艺性状差异 Table 1 Agronomic traits of wheat mutant LF2010 and its wild type H261

2.2 环境条件对突变体性状的影响

田间叶片遮光2周后揭开锡箔纸, 发现遮光部位的叶片几乎没有形成黄斑, 遮光处与未遮光处形成鲜明对比(图1-i), 说明LF2010突变体的黄色斑点受光照诱导。

室内温度试验表明, 20℃和15℃下生长至三叶期, 几乎全部单株在第1叶片上出现明显的黄斑; 而在10℃条件下, 只有少数单株叶片上有黄斑, 而且斑点明显少于在20℃和15℃培养条件下的叶片斑点; 在生长于5℃的幼苗上均未发现黄色斑点, 表明低温抑制斑点表达。进一步在5~10℃间设定温度梯度, 间隔为1℃。最终确定斑点出现的临界温度为8℃。

2.3 叶片斑点部位细胞死亡的鉴定

正常生长条件下, 野生型叶片和突变体斑点部位经锥虫蓝染色后均未出现深蓝色(图2), 表明突变体斑点部位不存在细胞坏死。

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	图2 突变体LF2010与野生型H261叶片锥虫蓝染色结果a: 染色前H261叶片; b: 染色后H261叶片; c: 染色前LF2010叶片; d: 染色后LF2010叶片。Fig. 2 Trypan blue staining results of mutant LF2010 and the wide typea: Leaf of H261 before staining; b: Leaf of H261 after staining; c: Leaf of LF2010 before staining; d: Leaf of the LF2010 after staining.

2.4 光合特性

2.4.1 光合色素含量斑点出现前, 突变体的单位叶面积叶绿素a、b和类萝卜素含量均与野生型无显著差异, 但总叶绿素含量低于野生型; 斑点出现初期, 突变体的叶绿素含量明显下降, 而类胡萝卜素含量与野生型无显著差异; 斑点布满整个旗叶后, 突变体的所有光合色素含量都与野生原始材料差异显著, 而且叶绿素a/b变大, 说明叶绿素b的下降程度比叶绿素a明显(表2)。

表2 不同时期突变体LF2010与野生型H261的叶绿素和类胡萝卜素含量及相对比值 Table 2 Contents of chlorophyll, carotenoid and chlorophyll a/b ratio in mutant LF2010 and the wild type H261 at different stages

2.4.2 光合参数测定旗叶出现斑点前, 突变体的 P_n、 G_s和 T_r均与野生型无显著性差异, 而 C_i显著高于野生型,说明该突变在斑点出现之前对旗叶的光合特性并无显著影响。在斑点出现初期和后期, 突变体的 P_n、 G_s和 T_r都明显下降, 且达到显著水平, 但 C_i始终显著大于野生型(表3)。

表3 不同时期突变体LF2010与野生型H261叶片的光合特性 Table 3 Photosynthetic characteristics of mutant LF2010 and the wild type H261 at different stages

2.5 遗传分析

斑点突变体LF2010与中国春和H261正反交F₁代单株全部表现正常绿色, 并且LF2010与中国春杂交F₂群体中正常植株与突变植株分离比, 经卡方检验符合3∶1 ( P= 0.8~0.9), BC₁群体中正常植株与突变植株分离比符合1∶1 ( P= 0.8~0.9), 表明LF2010的突变性状是由1对隐性核基因控制(表4)。

表4 斑点突变与H261和中国春杂交后代性状分离比 Table 4 Segregation for spotted mutation

3 讨论

已报道的小麦斑点突变包括类病变突变体和普通斑点突变体2种类型, 前者如隐性基因控制的突变体HLP^{[ 19]}、AIM9^{[ 21]}、Ning 7840^{[ 22]}和Yanzhan/ Zaosui 30^{[ 23]}, 后者如显性基因控制的突变体C591 (M8)^{[ 18]}。HLP、Ning 7840和Yanzhan/Zaosui 30及本试验中的LF2010都是隐性核基因突变体, 但与其他突变体不同的是, LF2010在斑点出现部位不存在坏死症状, 为普通斑点突变类型; 另外, 斑点的颜色和出现的时期及部位也不同。LF2010是黄色斑点, 在三叶期表达, 随后遍布全株; HLP为白色斑点, 于植株长出第5或第6叶期开始表达, 并且扩散到整株; Ning 7840是微黄色斑点, 在抽穗后只在叶片上表达; Yanzhan/Zaosui 30为褐色坏死斑点, 孕穗期表达, 斑点会变大且干枯, 遍布全株。LF2010与C591(M8)虽然属于同一类型的斑点突变, 但LF2010为隐性突变, 而C591(M8)为显性突变, 并且二者斑点出现的时期也不同。LF2010是在三叶期表达斑点, C591(M8) 在长出旗叶后才开始出现斑点。温度影响斑点的发生, 通过对水稻类病变 lrd^{[ 28]}和玉米类病变突变体 rp1^{[ 29]}研究发现, 低温明显促进病斑的发生, 然而, LF2010与此表现相反, 低温抑制了斑点发生。目前, 尚未见到关于温度对小麦斑点表达影响的报道, 因此推测, LF2010中的突变基因可能为新的斑点突变基因。

光合色素尤其是叶绿素在植物光合作用中对光能的吸收、传递和转化有着极为重要的作用, 类萝卜素作为捕光色素是色素蛋白复合体不可缺少的组分, 并且在光保护中起重要作用。多数叶色突变会导致植株色素含量和光合作用的改变^{[ 30, 31, 32]}。本研究表明, LF2010的叶绿素a、叶绿素b和类胡萝卜素含量在斑点出现前与野生型并无区别, 在黄斑出现初期叶绿素含量开始下降, 并随着斑点的增多而进一步下降, 到斑点布满整个叶片时, 已显著低于野生型; 而类胡萝卜素含量在黄斑刚出现后并不受影响, 只有在后期才明显下降, 说明斑点的出现主要影响的是叶绿素的代谢平衡。

光合速率的变化也随着斑点的出现与色素变化规律一致, 即斑点出现前LF2010与H261的 P_n值无显著差别, 斑点一旦出现, P_n值显著下降。我们还发现, 斑点出现前后 C_i值始终呈增加趋势。有研究表明, 胞间CO₂分压变化的方向(上升还是下降)是判断光合速率的降低是由于气孔限制还是非气孔因素引起的一个重要依据, 当 P_n和 C_i同时下降时, 其原因是气孔的部分关闭; 如果 C_i上升则是由于叶肉细胞光合活性的降低^{[ 33]}。由于ATP和NADPH供应不足, 即同化力不足会引起 P_n和 G_s下降, C_i上升, 而本试验中 P_n和 G_s下降正是黄斑出现后叶绿素减少, 在光照条件下叶绿体产生同化力不足叶肉细胞光合活性下降所造成的。

与突变体C591(M8)和HLP的农艺性状与其各自野生型无显著差异^{[ 18, 19]}不同, LF2010的株高、单株有效穗数、单株产量均显著低于野生型H261, 其原因可能与LF2010突变性状表达的时期较早, 斑点数量较多有关, 也说明该类突变对农艺性状的影响程度取决于斑点出现的时期、位置和扩展形式。Yao等^{[ 23]}认为, 在小麦产量性状中, 千粒重受突变类型的影响最明显。与此不同的是, LF2010的千粒重与原始亲本无差别, 这可能与本突变材料的穗粒数明显下降有关^{[ 34]}。

斑点突变体中多数属于类病斑坏死突变体, 其表达的斑点与植物在防卫过程中的过敏反应(hypersensitive response, HR)很相似, 多数属于类病斑坏死突变体在类病斑坏死形成之前或之后表现出对病原物局部或系统抗性的增强^{[ 35]}, 推测类病斑坏死突变体和防御反应可能具有一些共同的形成机制^{[ 36]}, 因此, 多数学者将其作为研究植物抗病性机制以及细胞程序性死亡(programmed cell death, PCD)途径的重要实验材料。LF2010不存在细胞死亡, 我们对LF2010和H261进行白粉病菌侵染实验, LF2010未表现出优于野生型的抗性(未发表结果), 目前正在验证该突变体对其他病原菌的抗性反应, 希望能对斑点形成机制有所解释。

4 结论

LF2010突变体的斑点性状由1对隐性基因控制, 斑点部位不存在细胞坏死, 但影响叶绿素的正常代谢, 导致部分农艺性状与野生型有显著差异。对斑点部位、发生和扩展、光照和温度反应等鉴定表明, 该突变体可能是一类新突变型。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.

参考文献

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1967

0.0

... 三是阶段性失绿突变体, 即在个体发育的某个阶段叶色不正常^[1] ...

1999

3.15

0.0

... 植物成色过程十分复杂, 这就决定了叶色突变原因的多变性和突变类型的多样性^[2,3,4,5], 因此, 收集和研究种类更多或独特的叶色突变体不仅对高等植物光合作用、色素合成、基因表达调控等基础研究有重要理论意义, 还可为作物高光效育种提供有利依据 ...

2005

1.44

0.0

1991

3.386

0.0

2006

0.0

2006

0.0

1.321

. 2006, 26:2083-2087

Photosynthesis and chlorophyll fluorescence characters of xantha wheat mutants

对小麦自然黄化突变体及其突变亲本(西农1718)的叶绿素含量、光合速率及叶绿素荧光动力学参数进行比较分析.结果显示:(1)突变体金黄株、绿黄株、黄绿株的叶绿素含量均显著低于突变亲本,总叶绿素含量分别为突变亲本的17%、24%和58%,表明该突变体为叶绿素缺乏突变体;3个突变体叶绿素a与叶绿素b的比值(Chl a/Chl b)均小于突变亲本,而且突变体叶绿素含量越低,Chl a/Chl b比值越小,说明该突变体Chl a下降幅度大于Chl b.(2)金黄株净光合速率在孕穗期、开花期仅为突变亲本的5.7%、2.4%;绿黄株净光合速率显著低于突变亲本,为突变亲本的57.7%、43.3%;而叶绿素含量仅为突变亲本一半的黄绿株,其净光合速率接近突变亲本,表明该黄化突变体叶绿素含量在一定范围内单位叶绿素含量的光合效率较高.(3)突变体Fo均显著低于突变亲本;金黄株、绿黄株的Fm,Fy,qP,qN显著低于突变亲本;金黄株Fv/Fm比值(0.671)显著低于突变亲本.研究表明,叶绿素含量在一定范围内减少,未引起突变体叶绿素荧光动力学参数(Fo除外)显著改变,而当叶绿素含量较大程度减少时,这些荧光参数会急剧降低.

... 常见的叶色突变类型主要为单色突变, 如黄化、白化、浅绿等, 通常色素含量表现不同程度降低^[6,7,8,9] ...

2008

0.0

1.237

. 2008, 22:131-135

Construction of mutant library for soybean ‘Nannong 94-16’ and analysis of some characters

The soybean mutant library was created by treating the seeds of soybean "Nannong 94-16" with NaN_3-60Co γ-rays irradiation combination treatment and EMS,inspectively.In M_3 generations,54 mutants of leaf,stem,flower,seed and cotyledon mutants were obtained by treatment of NaN_3-60Co γ-rays irradiation,and 66 mutants by EMS,inspectively.Water-logging tolerance mutants of 14 and the light green leaf color mutant were identified by 644 SSR primer pairs.The results showed that,among 14 water-logging tolerance mutants,more than 100 primer pairs displayed polymorphism in 2 mutants and one to seven SSR primer pairs displayed polymorphism in other plants;for the light green leaf color mutant,39 primer pairs displayed polymorphism.Genetic analysis to the light green leaf color mutant showed that this character was controlled by one pair of recessive nuclear genes.The mutant library created for "Nannong 94-16" will be helpful for the research in soybean functional genomics.

本研究对"南农94-16"大豆种子用NaN3-60Coγ射线复合诱变以及EMS诱变来构建大豆突变体库。两种方式诱变大豆分别获得54份和66份叶、茎、花、种子、子叶等性状变异的突变体。用644对SSR标记分别对其中的14株耐涝突变体和1株叶色浅绿突变体进行了鉴定。结果表明,14株耐涝突变体,有2株与对照有超过100个标记的差异,其余植株分别有1到7个标记的差异;叶色浅绿突变体与对照有39个标记的差异,遗传分析表明该性状由细胞核单基因隐性遗传控制。这些突变体可以作为新的种质资源,构建的突变体库也有助于大豆功能基因组研究的开展。

... 常见的叶色突变类型主要为单色突变, 如黄化、白化、浅绿等, 通常色素含量表现不同程度降低^[6,7,8,9] ...

2011

0.0

1.667

. 2011, 37:119-126 DOI:10.3724/SP.J.1006.2011.00119

Agronomic traits and photosynthetic characteristics of chlorophyll-deficient wheat mutant induced by spaceflight environment

Institute of Crop Sciences, Chinese Academy of Agricultural Sciences/National Key Facility for Crop Gene Resources and Genetic Improvement/ National Center of Space Mutagenesis for Crop Improvement, Beijing 100081, China

Chlorophyll deficiency mutants are ideal materials to study photosynthetic mechanism, pathways of chlorophyll biosynthesis and degradation, and genes related to photosynthesis. The chlorophyll-deficient wheat (Triticum aestivum L.) mutant Mt135, induced by space mutagenesis, was investigated on main agronomic traits and photosynthesis characteristics with a comparison to its wild type. The leaf color of the Mt135 showed albino, stripe, and green phenotypes. The albino plants with entire albino leaves died at seedling stage, while plants with green-and-white striped leaves usually matured 5–7 d later than the wild type, and the plant height, spike length, grains and grain weight per plant, and 1000-grain weight were lower. The green plants were not significantly different from the wild type. Preliminary genetic analysis showed that the inheritance model of Mt135 was nucleo-cytoplasmic interaction. When exposed to photosynthetic active radiation (PAR) of 110 μmol m^-2 s^-1, the green tissue of striped plants (S-G) and the wild plant had no significant difference on the value of maximum photosystem II quantum yield, but the potential activity of photosystem II was significantly lower in the S-G plants than in the wild plant. The changes of the photochemical quenching, non-photochemical quenching, effective quantum yield, regulated non-photochemical energy dissipation, and non-regulated energy dissipation were different at various growth stages. In addition, the differences of the electron transport rate, photochemical quenching, and effective quantum yield between S-G plant and wild plant varied under different PAR conditions. The white tissue of striped plant and the albino plant completely lose the photosynthetic function. As a result, the photosynthesis of Mt135 was greatly restricted. High photosynthetic active radiation had tremendous impact on the mutant at elongation stage and relatively weak impact at heading stage. The changes of photosynthetic function of striped plant are in accordance with the reduction of their plant height, spike length, and yield related traits.

叶绿素缺失突变体对于研究植物光合作用机制, 揭示叶绿素生物合成与降解途径, 发掘鉴定光合作用相关新基因以及了解基因间的相互作用有重要意义。空间诱变创制的小麦叶绿素缺失突变体Mt135的叶色表现为完全白化、条纹和绿3种类型, 其中完全白化株叶片完全白化, 于苗期死亡; 条纹株叶片为绿白相间的条纹, 能够正常成穗结实, 但其株高、穗长、株粒数、株粒重、千粒重都显著低于原始亲本, 生育期比原始亲本延长5~7 d; 绿株与原始亲本没有显著差异。初步遗传分析表明, Mt135是一个由核质基因共同作用的突变材料。对突变体及其原始亲本叶绿素荧光动力学参数分析表明, 当光照强度为110 μmol m^-2 s^-1时, 条纹株绿色组织光系统II的最大量子产量与原始亲本无显著差异, 光系统II的潜在活性显著低于原始亲本, 而光化学猝灭系数、非光化学猝灭系数、实际量子产量、调节性能量耗散的量子产量、非调节性能量耗散的量子产量在不同的生育期间变化不同。另外, 不同的光照强度下, 条纹株绿色组织的电子传递速率、光化学猝灭系数、实际量子产量的变化也不相同。条纹株白色组织和完全白化株则完全失去光合能力。上述结果证实, 小麦叶绿素缺失突变体Mt135的光合作用受到很大的影响, 光合特性发生了改变, 较高的光照强度在拔节期对突变体影响较大, 抽穗期影响相对较小。条纹株光合特性的改变与其株高、穗长和产量相关性状显著降低的结果相互印证。

... 常见的叶色突变类型主要为单色突变, 如黄化、白化、浅绿等, 通常色素含量表现不同程度降低^[6,7,8,9] ...

1980

1.513

0.0

... 常见的叶色突变类型主要为单色突变, 如黄化、白化、浅绿等, 通常色素含量表现不同程度降低^[6,7,8,9] ...

1999

6.582

0.0

... 斑点突变是一种特殊的杂色突变类型, 受到广泛关注, 自20世纪90年代以来, 已在玉米、水稻、大麦、大豆、花生、拟南芥中报道了大量斑点突变, 并相继进行了深入研究^{[10,11,12,13,14]} ...

2002

1.995

0.0

2012

0.0

1.494

. 2012, 26:297-301 DOI:10.3969/j.issn.10017216.2012.03.007

Genetic analysis and gene mapping of a light brown spotted leaf mutant in rice

¹ State Key Laboratory of Rice Biology/Chinese National Center for Rice Improvement, China National Rice Research Institute, Hangzhou 310006, China; ² College of Life and Environmental Sciences, Hangzhou Normal University, Hangzhou 310036, China; ³ College of Life Sciences, Anhui Normal University, Wuhu 241000， China; ⁴ International Rice Research Institute, DAPO Box 7777 Metro Manila, Philippines;

A light brown spottedleaf mutant of rice was isolated from an EMSinduced IR64 mutant bank. The mutant, designated as lbsl1（light brown spottedleaf 1）, displayed light brown spot in the whole growth period from the first leaf to the flag leaf under natural summer field conditions. Agronomic traits including plant height, growth duration, seedsetting rate and 1000grain weight of the mutant were significantly affected. Genetic analysis showed that the mutation was controlled by a single recessive gene, tentatively termed lbsl1(t), which was mapped to the short arm of chromosome 6. With developed SSR markers, the gene was finally delimited to an interval of 130 kb between markers RM586 and RM588. The lbsl1(t) gene is likely a novel rice spottedleaf gene since no other similar genes have been identified near the chromosomal region. The genetic data and recombination populations provided will facilitate further fine mapping and cloning of the gene.

通过EMS诱变籼稻品种IR64获得一个稳定遗传的淡褐色斑点叶突变体lbsl1（light brown spotted leaf 1）。在自然条件下，突变体播种后10~14 d，叶片上出现淡褐色斑点，随后逐渐扩散至全叶，第1叶至剑叶上均有淡褐色斑，为全生育期性状。斑点性状的表达对株高、生育期、结实率和千粒重等农艺性状具有显著的影响。遗传分析结果表明，该淡褐色斑点叶性状受一个隐性核基因控制。将突变体lbsl1与正常叶色水稻Morobereken杂交构建F2定位群体，利用SSR标记，最终将该淡褐叶基因lbsl1(t)定位在第6染色体短臂上一个约130 kb的区段上。定位的结果和发展的群体为该基因的进一步精细定位和克隆奠定了基础。

1997

31.957

0.0

2001

6.582

0.0

2010

0.0

1.494

. 2010, 24:108-115 DOI:10.3969/j.issn.1001-7216.2010.02.02

Recent advances in research on spotted-leaf mutants of rice (Oryza sativa)

1State Key Laboratory of Rice Biology/Chinese National Center for Rice Improvement, China National Rice Research Institute, Hangzhou 310006, China; 2College of Life and Environmental Sciences, Hangzhou Normal University, Hangzhou 310036, China; ＃These authors contributed equally to this paper; *Corresponding author, E-mail: beishangd@163.com

Many rice spottedleaf mutants are ideal sources for understanding the mechanisms involved in blast resistance, bacterial blight resistance and programmed cell death in plants. The genetic controls of 49 spottedleaf mutants in rice have been characterized and a few couple of spl/Spl genes have been isolated as well. This article reviews on the origin, genetic modes, isolation and characterization of spottedleaf genes responsible for their phenotypes, and their resistance responses to main rice diseases.

大多数水稻斑点叶突变体是研究稻瘟病抗性、白叶枯病抗性以及植物程序性细胞死亡机理的理想材料。已经阐明了49份水稻斑点叶突变体的遗传模式,并成功克隆分离了若干斑点叶基因。综述了这些突变体的来源、遗传模式、基因定位和克隆，基因的功能以及它们对水稻主要病害的反应。

... 另一类是普通斑点突变, 不存在坏死症状而纯粹为叶片颜色的变化^[15] ...

... 多数斑点突变为类病变突变体, 因此常用于抗病性研究^[15,16], 而对于植物色素代谢和光合特性相关的斑点突变鲜有报道, Les22是已较深入研究的玉米突变体, 其突变基因编码卟啉代谢途径中合成叶绿素的关键酶尿卟啉原脱氢酶(UROD)^[17] ...

1975

1.995

0.0

1998

9.251

0.0

2001

1.643

0.0

. 2001, 121:53-58 DOI:10.1023/A:1012061016783

Genetical and anatomical analyses of a leaf flecking mutant in Triticum aestivum L

1.Division of Genetics, Indian Agricultural Research Institute, New Delhi, 110012 India

Flecking trait in the mutant C591 (M8) Triticum aestivum L. is a stable,developmentally programmed, dominant mutation under monogenic control resembling pathogenic attack and starts appearing only from boot leaf stage of the plant. Mutant plants differ significantly from normal plants in terms of total chlorphyll contents only at later stages of symptom spread when the flecks fully cover the leaf sheath and leaves. However, total grain weight per main spike of mutant did not differ significantly from the normal plants. Microscopic studies of the mutant leaves did not reveal any damaging effect of the mutation on leaf anatomy per se, even though differences were observed in chlorophyll filling in mesophyll cells. Considering the peculiar characteristics of the mutation, many of which resembling the disease lesion mimic mutations in other crops, this is suspected to be such a mutation in wheat.

... Nair等^[18]利用化学诱变得到的显性斑点突变体C591(M8), 在斑点出现初期, 突变体的叶绿素含量与野生型差异不显著, 而当斑点布满整个叶片时, 突变体的色素含量较野生型明显下降, 但该突变性状对产量性状影响不显著 ...

... 3 讨论已报道的小麦斑点突变包括类病变突变体和普通斑点突变体2种类型, 前者如隐性基因控制的突变体HLP ^[19]、AIM9 ^[21]、Ning 7840 ^[22]和Yanzhan/ Zaosui 30 ^[23], 后者如显性基因控制的突变体C591 (M8) ^[18] ...

... 与突变体C591(M8)和HLP的农艺性状与其各自野生型无显著差异^[18,19]不同, LF2010的株高、单株有效穗数、单株产量均显著低于野生型H261, 其原因可能与LF2010突变性状表达的时期较早, 斑点数量较多有关, 也说明该类突变对农艺性状的影响程度取决于斑点出现的时期、位置和扩展形式 ...

2007

2.729

0.0

... Kamlofski等^[19,20]对HLP (hypersensitive-like phenotype)斑点突变体的研究发现, 斑点对产量性状同样没有显著影响, 并且HLP突变体成株期抗病性增强 ...

2010

0.796

0.0

... Kamlofski等^[19,20]对HLP (hypersensitive-like phenotype)斑点突变体的研究发现, 斑点对产量性状同样没有显著影响, 并且HLP突变体成株期抗病性增强 ...

2006

0.0

1.344

... Luo等^[21]研究发现, 小麦斑点突变体AIM9受1对隐性基因控制, 其色素含量、净光合速率以及单株产量较野生型材料明显下降 ...

2009

3.658

0.0

. 2009, 119:13-21 DOI:10.1007/s00122-009-1012-7

Lesion mimic associates with adult plant resistance to leaf rust infection in wheat

1.Kansas State University Department of Plant Pathology Manhattan KS 66506 USA
2.USDA-ARS Plant Science and Entomology Research Unit Manhattan KS 66506 USA

Lesion mimics (LM) that resemble plant disease symptoms in the absence of plant pathogens may confer enhanced plant disease resistance to a wide range of pathogens. Wheat line Ning7840 has adult plant resistance (APR) to leaf rust (Puccinia triticina) and shows LM symptoms at heading. A recessive gene (lm) was found to be responsible for LM in Ning7840 and located near the proximal region of chromosome 1BL using a population of 179 recombinant inbred lines (RIL) derived from the cross Ning7840/Chokwang. Genomic in situ hybridization showed that Ning7840 carries the short arm of 1R chromosome from rye (Secale cereale L.), on which the race-specific gene Lr26 resides. The RILs were infected with the isolate PRTUS 55, an isolate virulent to Lr26, at anthesis in two greenhouse experiments. The result showed that the lines with LM phenotype had a significantly higher rust resistance than the non-LM lines. Composite interval mapping consistently detected a QTL, Qlr.pser.1BL, for APR on chromosome 1BL. Qlr.pser.1BL peaked at lm and explained up to 60.8% of phenotypic variation for leaf rust resistance in two greenhouse experiments, therefore, lm from Ning7840 may have pleiotropic effects on APR to leaf rust.

... 最近, 个别小麦斑点突变基因已被定位到染色体上^[22,23] ...

2009

3.658

0.0

. 2009, 119:1005-1012 DOI:10.1007/s00122-009-1104-4

Characterization and mapping of complementary lesion-mimic genes lm1 and lm2 in common wheat

1.Sichuan Agricultural University Agronomy College 625014 Yaan People’s Republic of China
2.Chinese Academy of Agricultural Sciences Key Laboratory of Crop Germplasm and Biotechnology, Ministry of Agriculture, National Key Facility for Crop Gene Resources and Genetic Improvement, Institute of Crop Science 100081 Beijing People’s Republic of China
3.Zhejiang University College of Agriculture and Biotechnology 310029 Hangzhou People’s Republic of China

A lesion-mimic phenotype appeared in a segregating population of common wheat cross Yanzhan 1/Zaosui 30. The parents had non-lesion normal phenotypes. Shading treatment and histochemical analyses showed that the lesions were caused by light-dependent cell death and were not associated with pathogens. Studies over two cropping seasons showed that some lines with more highly expressed lesion-mimic phenotypes exhibited significantly lower grain yields than those with the normal phenotype, but there were no significant effects in the lines with weakly expressed lesion-mimic phenotypes. Among yield traits, one-thousand grain weight was the most affected by lesion-mimic phenotypes. Genetic analysis indicated that this was a novel type of lesion mimic, which was caused by interaction of recessive genes derived from each parent. The lm1 (lesion mimic 1) locus from Zaosui 30 was flanked by microsatellite markers Xwmc674 and Xbarc133/Xbarc147 on chromosome 3BS, at genetic distances of 1.2 and 3.8 cM, respectively, whereas lm2 from Yanzhan 1 was mapped between microsatellite markers Xgwm513 and Xksum154 on chromosome 4BL, at genetic distances of 1.5 and 3 cM, respectively. The linked microsatellite makers identified in this study might be useful for evaluating whether potential parents with normal phenotype are carriers of lesion-mimic alleles.

... 最近, 个别小麦斑点突变基因已被定位到染色体上^[22,23] ...

... Yao等^[23]认为, 在小麦产量性状中, 千粒重受突变类型的影响最明显 ...

2005

0.0

1.494

. 2005, 19:111-116

Disease resistance and cytological analyses on lesion resembling disease mutant lrd40 in Oryza sativa

Lesion resembling disease mutant lrd40, obtained from gamma ray mutagenesis of a japonica rice variety Zhonghua 11, expressed whole life lesion mimic in different environments. Genetic analysis showed that this mutant is controlled by a single recessive locus nonallelic to the loci of lrd38, lrd39 and lrd42. Repeated inoculation with Xanthomonas oryzae pv. oryzae (Xoo) showed that lrd40 conferred non race specific resistance to Xoo, and resistance to Xoo was associated with the lesion expression of lrd40 in the F₂ generations tested, indicating that disease resistance is controlled by the lrd40 locus. Trypan blue and DAB staining methods were used to detect programmed like cell death and deposition of H₂O₂ during lesion development, all these observations resemble the early events of plant defense responses, including cell death and oxidative burst during the hypersensitive response.

由中花11经γ射线诱变获得的水稻类病变突变体lrd40，是一个对环境不敏感的全生育期表达的突变体。遗传分析表明，lrd40的类病变性状为1个隐性位点控制，与已报道的类病变突变体lrd38、lrd39、lrd42是非等位的。经多点接种试验，lrd40对水稻白叶枯病2个菌系PXO99A和DY89031都表现抗病。lrd40与野生型杂交F2接种结果证实，lrd40的抗白叶枯病与类病变的表现是一致的。锥虫蓝（trypan blue）染色观察到lrd40的类病变发生可能是一个程序性细胞死亡过程，经二氨基联苯胺（diamino benzidine，DAB）染色检测到lrd40的类病变表达时，叶片上产生H2O2沉淀。细胞学观察结果说明，lrd40的类病变发生类似于植物防卫反应的早期特征，如细胞死亡和过敏反应导致的氧迸发（oxidative burst）。

... 4 锥虫蓝染色法检测突变体叶片斑点部位的细胞活性锥虫蓝染色是检测植物细胞死亡的常用方法之一, 坏死的细胞由于细胞膜遭到破坏会被染成深蓝色^[24] ...

2011

0.0

1.494

. 2011, 25:150-156 DOI:10.3969/j.issn.1001-7216.2011.02.005

Identification and genetic mapping of a lesion mimic strip mutant in rice

(1Laboratory of Plant Genetics and Functional Gene, College of Life and Environment Science, Shanghai Normal University, Shanghai 200234, China; 2Institute of Plant Physiology & Ecology, Shanghai Institutes for Biological Sciences, Chinese Academy of Sciences, Shanghai 200032, China; 3National Key Laboratory of Crop Genetics and Germplasm Enhancement, Nanjing Agricultural University, Nanjing 210095, China; *Corresponding author, E-mail: dong@shnu.edu.cn; steng@sippe.ac.cn)

A lesion mimic stripe mutant was obtained from the M2 progeny of a japonica variety Jiahua 1 by 60Co γ radiation,temporarily designated as lms1 (lesion mimic stripe 1). The mutant displayed lesion mimic in its whole life, and belonged to propagation type. Physiological and histochemistry analysis suggested that the mutant displayed a phenotype of white stripe at a high temperature (30℃) and lesion mimic caused by cell initiative death at a low temperature(20℃)，respectively. The genetic analysis indicated that the mutant trait was controlled by a single recessive nuclear gene. In addition, using SSR markers and an F2 segregating population derived from two crosses(lms1/9311 and Pei’ai 64S/lms1), the lms1 gene was mapped between Indel1 and MM0112-4 with a physical distance of 400 kb on chromosome 6 in rice.

粳稻品种嘉花1号经60Co γ射线辐射诱变后获得一个水稻类病斑突变体MR07, 暂命名为lms1。该突变体的病斑在全生育期均表现，属于扩散型类病斑突变体。生理和组织化学分析表明，该突变体在高温条件下（30℃）培养时只表现为白色条斑，低温条件下（20℃）培养时表现出细胞自主性死亡的坏死病斑。遗传分析表明，该突变体受1对隐性核基因控制。以lms1突变体与籼稻9311、培矮64S杂交的两个F2分离群体作为定位群体，利用SSR标记和Indel标记将该基因定位在第6染色体上，位于标记InDel1和MM0112-4之间，其物理距离为400 kb。

... 参考马健阳等^[25]的方法检测突变体叶片的细胞死亡, 略有改动 ...

1990

9.251

0.0

... 当人工气候箱中幼苗长至五叶期的时, 随机选取已经表现斑点性状的突变体和野生型第1叶片, 浸泡在煮沸的锥虫蓝染液^[26]中, 染色7 min, 室温放置14 h, 再用2 ...

1985

15.333

0.0

... 用分光光度计分别测定叶绿素a和叶绿素b及类胡萝卜素的含量^[27] ...

2010

0.0

1.889

. 2010, 43:2039-2044

Response to illumination induction and effect of temperature on lesion formation of lrd (lesion resembling disease) in rice

(浙江省农业科学院作物与核技术利用研究所)

【Objective】 To determine the spectral range of light required for lesion formation and the effect of temperature on the expression of lrd lesions, the response of lrd expression to light and temperatures was tested for certain mutants in Oryza sativa L. 【Method】 Six lrd mutants derived from Indica and Japonica by gamma-ray mutagenesis simultaneously were illuminated by Light Emitting Diode (LED) light and were cultured in two different growth chambers set at three different temperature regimes. Lesions occurred on leave were observed and photographed for comparison the effect of different treatments. 【Result】 The expression of lrd lesions for lrd31 was induced by UV light, that of lrd36 was induced by yellow light, that of lrd37and lrd35 was induced by blue light, and lrd32 or lrd40 expressed lesion under all illuminated light. lrd32, lrd36 and lrd37 formed lesion under all culturing temperatures, and cooler environment accelerated their lesion expression, lrd35 and lrd40 were suppressed at high temperature and expressed at intermediate temperature, also accelerated under lower temperature. But lrd31 formed lesion only under intermediate temperature, and no lesion formation at high or lower culturing temperature. 【Conclusion】 The expressions of lrd lesions are light-dependent, different genotypes of lrd mutants in rice need specially spectral range of light. Temperature is another factor influencing lrd expression. The lower temperature seemed to enhance the expression of lrd phenotype.

【目的】对水稻lrd表达的光温诱导反应进行研究,明确水稻lrd表达光诱导波段和温度的抑制或促进作用。【方法】利用自有的6个水稻lrd突变体,用LED单色光源诱激和人工气候箱中不同温度梯度处理,观察类病斑的发生情况和统计病斑个数,比较诱导的效果。【结果】lrd31类病斑表达受紫外光诱导,lrd36的表达受黄光诱导,lrd37和 lrd35的表达受蓝光诱导,而lrd32 和lrd40 在所有处理的光源下均表达类病斑;lrd32、lrd36和lrd37等3种温度下都正常产生病斑,低温明显促进病斑的发生。lrd35和lrd40在高温下不表达,在中温下可正常表达,低温下也表现促进作用,但lrd31只有在中温下正常表达,低温和高温抑制病斑出现。【结论】不同的lrd基因型所需的光诱导反应波段是不相同的。温度是抑制或促进lrd表达的另一个重要环境因子,一般是较低的温度促进lrd表达,高温抑制lrd表达。

... 温度影响斑点的发生, 通过对水稻类病变lrd^[28]和玉米类病变突变体rp1^[29]研究发现, 低温明显促进病斑的发生, 然而, LF2010与此表现相反, 低温抑制了斑点发生 ...

1996

9.251

0.0

1979

9.737

0.0

... 多数叶色突变会导致植株色素含量和光合作用的改变^[30,31,32] ...

2003

0.0

. 2003, 30:1127-1132

Mapping of QTLs controlling chlorophyll content in rice

1.College of Life Science,Fujian Agriculture and Forestry University,Fuzhou 350002, China 2.College of Crop Science,Fujian Agriculture and Forestry University,Fuzhou 350002, China

Mapping of QTLs controlling chrolophyll content was conducted with the method of multiple-trait composite interval mapping, based on a recombinant inbred (RI) population consisting of 131 lines (F₁₀) derived from a cross between two indica rice varieties, Acc8558 and H359, and on a corresponding genetic map comprising 147 RFLP and 78 AFLP markers. Six QTLs for contents of chloropghyll a and chlorophyll b were detected, respectively. Among them, five QTLs were the same between the two characters. These QTLs were mainly distributed on chromosomes 1 and 4,indicating that the two chromosomes were more important for chlorophyll content. qChlA1c and qChlB1b (these two QTLs were mapped at the same location) showed relatively large effects at all the four observation times and showed the largest effects at the sword leaf period. Another two QTLs (qChlA4a/qChlB4a and qChlA4b/qChlB4b) exhibited significant effects only at the second observation time, suggesting that they might act only at specific developmental stages.

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（F10）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在两性状间是相同的。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 qChlA1c/qChlB1b（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（qChlA4a/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。

... 多数叶色突变会导致植株色素含量和光合作用的改变^[30,31,32] ...

2005

9.737

0.0

... 多数叶色突变会导致植株色素含量和光合作用的改变^[30,31,32] ...

1997

0.0

0.849

... 如果C_i上升则是由于叶肉细胞光合活性的降低^[33] ...

2002

0.0

0.3268

. 2002, (2):13-15

Study on the coordinating activity among major agromomic traits in wheat

选用40个小麦品种(系),运用方差分析、产量构成分析、生物产量分析等方法,对供试品种的11个主要农艺性状进行比较分析,以探讨小麦主要农艺性状协调性对单株产量的影响.结果表明,小麦品种(系)的这些性状与产量因素之间是相互制约、相互促进、协调发展的,不能片面追求单一目标性状的提高,而忽略品种其它性状的协调性.在育种工作中,既要注重某一优良性状,又要兼顾其它性状,考虑各优良性状的内在联系,只有各性状协调发展才是提高产量的最佳途径.

... 与此不同的是, LF2010的千粒重与原始亲本无差别, 这可能与本突变材料的穗粒数明显下降有关^[34] ...

1996

9.251

0.0

... 斑点突变体中多数属于类病斑坏死突变体, 其表达的斑点与植物在防卫过程中的过敏反应(hypersensitive response, HR)很相似, 多数属于类病斑坏死突变体在类病斑坏死形成之前或之后表现出对病原物局部或系统抗性的增强^[35], 推测类病斑坏死突变体和防御反应可能具有一些共同的形成机制^[36], 因此, 多数学者将其作为研究植物抗病性机制以及细胞程序性死亡(programmed cell death, PCD)途径的重要实验材料 ...

2008

2.277

0.0