甘蓝型油菜品种细胞壁特性与硼效率关系研究
摘要
甘蓝型油菜缺硼是普遍存在的生产制约因子。但其品种间的硼效率存在显著差异。因此通过对品种筛选获得硼高效品种，进而对其进行生理生化和遗传学研究，最终通过育种和分子生物学方法获得理想的硼高效品种，是缓解我国硼矿资源缺乏和减少环境污染、提高作物产量和品质的重要措施。本论文是在总结了本研究室多年来有关甘蓝型油菜品种硼效率基因型差异机理研究成果的基础上，利用王运华等筛选出的甘蓝型油菜硼高效品种(95090和9118)和硼低效品种(95105和9141)，运用营养液培养和细胞培养技术，研究甘蓝型油菜品种细胞壁特性与硼效率的关系及硼与植物细胞壁的关系。主要研究结果如下：
1．研究发现在Hoagland营养液培养体系中，0.001mg／L～0.005mg／L的硼浓度可作为在该体系中进行油菜硼效率苗期品种筛选和生理生化机制研究的硼临界浓度。同时发现油菜根系比地上部对缺硼更敏感，可作为鉴定品种硼效率的主要器官。
2．探明了不同油菜品种及其不同器官细胞壁组成的差异：根系细胞壁纤维素含量高于叶片，果胶含量低于叶片；愈伤细胞细胞壁纤维素含量低于根系和叶片，半纤维素含量高于根系和叶片，果胶含量与根系差异不大。不同硼效率油菜品种在细胞壁果胶、半纤维素和纤维素含量上差异不大；但硼高效品种新生叶片螫合剂可溶性果胶含量显著高于低效品种，碱溶性果胶含量显著低于低效品种；缺硼处理一周对根和叶细胞壁组成的影响很小；但使愈伤细胞细胞壁纤维素和半纤维素含量显著降低，SDS(十二烷基磺酸钠)可提取物显著升高。
3．证明了细胞壁及其果胶是硼、钙在油菜体内的主要结合位点，镁在细胞壁中的含量很低。这一结论丰富了前人的研究结果。
3．1．正常供硼条件下，不同油菜品种根系中均有约50％以上的硼结合在细胞壁中，叶片中的比例较低；胞壁硼含量受外界硼浓度变化的影响均小于相应的器官。缺硼时细胞壁硼含量占相应器官总硼量之比显著升高。
3．2．同一油菜品种不同器官细胞壁硼含量与其细胞壁果胶含量呈显著正相关。根系细胞壁果胶含量及其硼含量显著低于叶片。不论缺硼与否，根系细胞壁中85％以上、叶片细胞壁中90％以上的硼结合在果胶中。
3．3．正常供硼条件下，不同油菜品种根系中均有约30％以上的钙结合在细胞壁中。但细胞壁钙含量及其在细胞壁不同组分的分配受品种、生育期和外界硼浓度的影响较大，在细胞壁及其果胶中结合的专一性比硼弱。
3．4．硼、钙含量在根系和叶片细胞壁中呈负相关。根系细胞壁硼含量低，钙含量高；叶片细胞壁硼含量高，钙含量低。
4．引用先进的酶解法和DEAE-琼脂糖色谱柱法，首次在油菜不同品种细胞壁中分离出了含硼化合物，并初步证明该化合物为B-鼠李半乳糖醛酸聚糖-Ⅱ(B-RG-Ⅱ)；硼在细胞壁中专一地和该化合物结合，果胶的硼含量与其RG-Ⅱ含量呈正相关；低效品种果胶结构比高效品种复杂，但果胶中B-RG-Ⅱ的结构基本相同；缺硼影响果胶和RG-Ⅱ的结构。
5．首次运用原生质体培养方法，证实了硼是油菜不同品种细胞壁结构形成的必需元素。
6．在油菜硼效率差异的生理生化机制方面，提出了如下新见解：
6．1．从理论上证明了高效品种硼向上运转能力强是由于其根系和老叶细胞壁占器官鲜重之比及其硼含量较低，在正常供硼和缺硼条件下细胞壁硼量占器官总硼量之比均低于硼低效品种，即高效品种根系和老叶中自由态硼酸所占比例较高，因此在缺硼条件下硼向上运转的能力较强。
6．2．高效品种新生叶细胞壁硼含量较低，细胞壁形成所需的硼量显著低于低效品种，因而在一定的低硼浓度条件下，高效品种新生部位细胞壁的形成和细胞的生长必然好于低效品种。
6．3．高效品种新生叶片果胶的聚合度较低，可能有利于其适应外界环境条件的变化。
B deficiency in rape(Brassica Napus L.) is a common micronutrient deficiency phenomenon in field. But the cultivate response to low B varied widely. So it is possible to get ideal B efficient rape cultivars through methods of breeding and molecular biology. To achieve this goal, it is essential to make clear the mechanisms of efficiency. Recently studies showed that pectin content of plant cell wall may be the main factor to determine plant B efficient. In this stud), two B efficient rape cultivars and two B inefficient rape cultivars, which were screened by Wang Yunhua et al(1995) in our laboratory, were used to study B functions and distribution laws in the wall, and the differences of these respects among the cultivars. The main results as follow:
1. 0.001Bmg/L~0.005Bmg/L B content range can be used as critical B content to screen different B efficient rape cultivars in seedling stage in Hoagland nutrition culture system. The growing conditions and root activities in this range were distinguished difference in different B-efficient cultivars. In addition, the root was more sensible to B deficiency than the shoot, which can looked on as a main index to judge B efficient cultivars.
2. The constitution of wall were significantly different in different organs and cultivars and it was controlled by genetic and organ differentiation. The cellulose content in root is higher than that in leaf, pectin content is lower than that in leaf, and little difference exists in hetnicellulose. Cultured cells had the lowest cellulose content and the highest hemicellulose content The content of total pectin, cellulose and hemicellulose had little differences between B-efficient and B-inefficient cultivars. However, the dictation soluble pectin content was much higher and the alkaline soluble pectin content was much lower in B-efficient cultivars than in B-inefficient cultivars.
3. B had little effect on the constitution of root and leaf wall, but B deficiency could significantly decrease the cellulose content in cultured cell and significantly increases the SDS extractants.
4. It is proved that cell wall and pectin were the main sites combining B and Ca, and the Mg concentration in wall was very low, those were in agree with the other researcher's result.
4.1. Under normal B condition, about more than 50% of root B was found in the wall; about 30% in upper leaf and about 20% in lower leaf. And the wall B content was more stable in different B conditions than the organs. So the ratio of wall B was significantly increased in B deficient conditions.
4.2. More than 80% B in root wall and more than 90% B in leaf wall was found in pectin's.
4.3.The B concentration in root wall and in the pectin was distinguished lower than that in leaf. 4.4.The Ca concentration in cell wall and its distribution in cell wall constitutions were influenced by growth stages and vitro boron concentration, its capability of combining with cell wall pectin is not strong.
5. Through pectinase decomposition and DEAE- Sepharose chromalography methods, a B-complexe was separated for the first time in rape. And proved that this complexes may be B-RG-II.
6. Boron is proved to be an essential element in cell wall formation through the method of protoplast breeding.
7. New suggestions were put forward on mechanism of boron efficiency in rape cultivars.
7.1. B-efficient cultivars had lower B content in root and old leaf wall and the ratio of free boron was higher under B deficiency condition, which meaned stronger capability of transportation of B upward.
7.2. The B content needed in the protoplast wall rebuilding in B-efficient cultivars was much lower that that in B-inefficient cultivars. Therefore, the wall of B-efficient cultivars can form and grow normally under certain lower boron concentration.
7.3. Lower polymerization of the pectin in upper leaves was founded in B-efficient cultivars, which might be advantageous for the cultivator to adapt to the change of outside environment.
1．研究发现在Hoagland营养液培养体系中，0.001mg／L～0.005mg／L的硼浓度可作为在该体系中进行油菜硼效率苗期品种筛选和生理生化机制研究的硼临界浓度。同时发现油菜根系比地上部对缺硼更敏感，可作为鉴定品种硼效率的主要器官。
2．探明了不同油菜品种及其不同器官细胞壁组成的差异：根系细胞壁纤维素含量高于叶片，果胶含量低于叶片；愈伤细胞细胞壁纤维素含量低于根系和叶片，半纤维素含量高于根系和叶片，果胶含量与根系差异不大。不同硼效率油菜品种在细胞壁果胶、半纤维素和纤维素含量上差异不大；但硼高效品种新生叶片螫合剂可溶性果胶含量显著高于低效品种，碱溶性果胶含量显著低于低效品种；缺硼处理一周对根和叶细胞壁组成的影响很小；但使愈伤细胞细胞壁纤维素和半纤维素含量显著降低，SDS(十二烷基磺酸钠)可提取物显著升高。
3．证明了细胞壁及其果胶是硼、钙在油菜体内的主要结合位点，镁在细胞壁中的含量很低。这一结论丰富了前人的研究结果。
3．1．正常供硼条件下，不同油菜品种根系中均有约50％以上的硼结合在细胞壁中，叶片中的比例较低；胞壁硼含量受外界硼浓度变化的影响均小于相应的器官。缺硼时细胞壁硼含量占相应器官总硼量之比显著升高。
3．2．同一油菜品种不同器官细胞壁硼含量与其细胞壁果胶含量呈显著正相关。根系细胞壁果胶含量及其硼含量显著低于叶片。不论缺硼与否，根系细胞壁中85％以上、叶片细胞壁中90％以上的硼结合在果胶中。
3．3．正常供硼条件下，不同油菜品种根系中均有约30％以上的钙结合在细胞壁中。但细胞壁钙含量及其在细胞壁不同组分的分配受品种、生育期和外界硼浓度的影响较大，在细胞壁及其果胶中结合的专一性比硼弱。
3．4．硼、钙含量在根系和叶片细胞壁中呈负相关。根系细胞壁硼含量低，钙含量高；叶片细胞壁硼含量高，钙含量低。
4．引用先进的酶解法和DEAE-琼脂糖色谱柱法，首次在油菜不同品种细胞壁中分离出了含硼化合物，并初步证明该化合物为B-鼠李半乳糖醛酸聚糖-Ⅱ(B-RG-Ⅱ)；硼在细胞壁中专一地和该化合物结合，果胶的硼含量与其RG-Ⅱ含量呈正相关；低效品种果胶结构比高效品种复杂，但果胶中B-RG-Ⅱ的结构基本相同；缺硼影响果胶和RG-Ⅱ的结构。
5．首次运用原生质体培养方法，证实了硼是油菜不同品种细胞壁结构形成的必需元素。
6．在油菜硼效率差异的生理生化机制方面，提出了如下新见解：
6．1．从理论上证明了高效品种硼向上运转能力强是由于其根系和老叶细胞壁占器官鲜重之比及其硼含量较低，在正常供硼和缺硼条件下细胞壁硼量占器官总硼量之比均低于硼低效品种，即高效品种根系和老叶中自由态硼酸所占比例较高，因此在缺硼条件下硼向上运转的能力较强。
6．2．高效品种新生叶细胞壁硼含量较低，细胞壁形成所需的硼量显著低于低效品种，因而在一定的低硼浓度条件下，高效品种新生部位细胞壁的形成和细胞的生长必然好于低效品种。
6．3．高效品种新生叶片果胶的聚合度较低，可能有利于其适应外界环境条件的变化。
B deficiency in rape(Brassica Napus L.) is a common micronutrient deficiency phenomenon in field. But the cultivate response to low B varied widely. So it is possible to get ideal B efficient rape cultivars through methods of breeding and molecular biology. To achieve this goal, it is essential to make clear the mechanisms of efficiency. Recently studies showed that pectin content of plant cell wall may be the main factor to determine plant B efficient. In this stud), two B efficient rape cultivars and two B inefficient rape cultivars, which were screened by Wang Yunhua et al(1995) in our laboratory, were used to study B functions and distribution laws in the wall, and the differences of these respects among the cultivars. The main results as follow:
1. 0.001Bmg/L~0.005Bmg/L B content range can be used as critical B content to screen different B efficient rape cultivars in seedling stage in Hoagland nutrition culture system. The growing conditions and root activities in this range were distinguished difference in different B-efficient cultivars. In addition, the root was more sensible to B deficiency than the shoot, which can looked on as a main index to judge B efficient cultivars.
2. The constitution of wall were significantly different in different organs and cultivars and it was controlled by genetic and organ differentiation. The cellulose content in root is higher than that in leaf, pectin content is lower than that in leaf, and little difference exists in hetnicellulose. Cultured cells had the lowest cellulose content and the highest hemicellulose content The content of total pectin, cellulose and hemicellulose had little differences between B-efficient and B-inefficient cultivars. However, the dictation soluble pectin content was much higher and the alkaline soluble pectin content was much lower in B-efficient cultivars than in B-inefficient cultivars.
3. B had little effect on the constitution of root and leaf wall, but B deficiency could significantly decrease the cellulose content in cultured cell and significantly increases the SDS extractants.
4. It is proved that cell wall and pectin were the main sites combining B and Ca, and the Mg concentration in wall was very low, those were in agree with the other researcher's result.
4.1. Under normal B condition, about more than 50% of root B was found in the wall; about 30% in upper leaf and about 20% in lower leaf. And the wall B content was more stable in different B conditions than the organs. So the ratio of wall B was significantly increased in B deficient conditions.
4.2. More than 80% B in root wall and more than 90% B in leaf wall was found in pectin's.
4.3.The B concentration in root wall and in the pectin was distinguished lower than that in leaf. 4.4.The Ca concentration in cell wall and its distribution in cell wall constitutions were influenced by growth stages and vitro boron concentration, its capability of combining with cell wall pectin is not strong.
5. Through pectinase decomposition and DEAE- Sepharose chromalography methods, a B-complexe was separated for the first time in rape. And proved that this complexes may be B-RG-II.
6. Boron is proved to be an essential element in cell wall formation through the method of protoplast breeding.
7. New suggestions were put forward on mechanism of boron efficiency in rape cultivars.
7.1. B-efficient cultivars had lower B content in root and old leaf wall and the ratio of free boron was higher under B deficiency condition, which meaned stronger capability of transportation of B upward.
7.2. The B content needed in the protoplast wall rebuilding in B-efficient cultivars was much lower that that in B-inefficient cultivars. Therefore, the wall of B-efficient cultivars can form and grow normally under certain lower boron concentration.
7.3. Lower polymerization of the pectin in upper leaves was founded in B-efficient cultivars, which might be advantageous for the cultivator to adapt to the change of outside environment.
引文
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