葡萄四倍体诱导及其生物学研究
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摘要
本文以二倍体青岛早红、鞍山早红、力扎马特和诱变获得的同源四倍体玫瑰香葡萄为试材,系统研究了提高秋水仙素诱变葡萄茎尖成功率的方法、不同世代的变异规律、同源四倍体玫瑰香嫩枝扦插生根特性变异、光合特性变异以及部分多倍体鉴别指标的评价方法。主要结果如下:
1、进一步完善了秋水仙素茎尖诱变技术
在系统研究的基础上,秋水仙素诱变葡萄茎尖的成活率较对照提高了3.9%～13.3%,其中“秋水仙素处理+保湿袋+报纸袋+遮阳网”以及处理后逐渐解除保护的方法,诱变节位的成活率达到了90.6%;系统完善了VM1代的初次选择技术,以及VM1代和VM2代的快速分离技术;提出了秋水仙素诱变获得的葡萄多倍体材料,在无性繁殖过程中存在较大幅度的变异,在后代筛选时应增加VM2代的群体数量。创新了13份鞍山早红和力扎马特葡萄同源多倍体种质资源。
2、系统研究了秋水仙素茎尖诱变不同世代的变异规律
同源四倍体葡萄的叶脉结构发生了显著的变异,其主脉的主导地位被消弱,主脉上分生的侧脉对称性降低,叶片形状变为楔形或肾形比率显著增加,叶脉数量明显减少。利用秋水仙素诱变葡萄茎尖后,其VM1代叶片的宏观结构和微观结构均发生了广泛的变异。其中叶片形状、大小、叶脉、叶缘等13个宏观性状的变异幅度增加,出现了一些新的叶片类型。
3、探明了同源多倍体葡萄生根特性的变异
(1)在形态学方面,同源四倍体葡萄嫩枝扦插生根能力较母株显著下降,扦插不定根发生率、平均每个插条的不定根数量和不定根总长度均显著低于其二倍体母株。
(2)在解剖学方面,同源四倍体玫瑰香嫩枝插条的髓部面积增大、髓射线数量减少;在根原基诱导阶段,其表皮细胞和皮层细胞过早、过量分化,根原基诱导期延长了1d;在根原基发育期,其根原基体积变大、顶部变平。上述形态与解剖性状的变异可能是导致同源四倍体玫瑰香生根能力下降的原因。并提出,将同源四倍体葡萄插条不定根发育可划分为根原基诱导期、根原基发育期和不定根发育期三个阶段。
(3)在生理学方面,同源四倍体玫瑰香嫩枝插条在扦插生根过程中的碳素营养、酚类和黄酮物质、IAAO酶活性、POD酶活性、PPO酶活性和内源激素代谢发生了显著的变异:①同源四倍体玫瑰香在扦插初期的可溶性糖含量、淀粉含量显著低于二倍体玫瑰香。同源四倍体玫瑰香的淀粉含量在根原基诱导期与二倍体玫瑰香品种的变化态势相反,且在扦插第3～12d其淀粉含量始终低于二倍体玫瑰香。②同源四倍体玫瑰香嫩枝插条基部皮层的黄酮含量在根原基诱导初期显著高于二倍体玫瑰香、后期又显著低于二倍体玫瑰香,两个不同倍性品种的变化态势相反。同源四倍体玫瑰香的多酚含量在根原基诱导初期高于二倍体玫瑰香,第3d以后低于二倍体玫瑰香。③在根原基诱导期,同源四倍体玫瑰香的IAAO酶活性显著高于二倍体玫瑰香,POD酶活性显著低于二倍体玫瑰香,PPO酶活性高于二倍体玫瑰香,而在此期以后PPO酶活性又低于二倍体玫瑰香。④玫瑰香葡萄同源加倍以后,不仅改变了不同时期插条生根部位皮层的内源激素水平,也改变了该部位对外施NAA的反应。在不定根诱导期(0～3 d),促进生根的生长素水平下降,抑制生根的脱落酸和赤霉素含量上升,生长素与脱落酸和赤霉素的比值下降;应用NAA处理以后,在根原基诱导期(0～3 d),赤霉素、玉米素和玉米素核苷含量不降反升,改变了内源激素的整体变化态势。
4、探讨了同源多倍体葡萄光合特性的变异
同源四倍体玫瑰香较二倍体玫瑰香单位面积的叶绿素含量增大,栅栏组织细胞长度与宽度增加,单位叶片解剖长度内栅栏组织细胞层数减少。其光合特性发生了显著变异:①同源四倍体玫瑰香最适的光合温度为20℃左右。在温度诱导条件下,同源四倍体玫瑰香2号对温度的敏感性最强,表现具有较强的耐高温能力。同源四倍体玫瑰香1号和二倍体玫瑰香对温度的反应较迟钝。②在自然条件下,同源四倍体玫瑰香与二倍体玫瑰香的光合作用曲线均为双峰曲线。不同倍性玫瑰香葡萄的蒸腾速率、气孔导度和净光合速率为:M4x-1﹥M2x﹥M4x-2。同源四倍体玫瑰香1号和二倍体玫瑰香光合作用曲线相似,第一个峰值均出现在上午9:00时,第二个峰值均出现在14:00时,第一个峰值大于第二个峰值,且9:00～11:00时的净光合速率下降不明显;同源四倍体玫瑰香2号的第一个峰值也出现在上午9:00时,但第二个峰值出现在13:00,第一个峰值与第二个峰值相当,且9:00～11:00净光合速率明显下降。
5、探索改进了部分性状的评价方法
简化了观测气孔的制片方法;该方法简便易行,效果良好,提高了气孔性状观测效率。提出了一种测量花蕾大小的新方法;应用这种方法可以准确测量花蕾的大小,消除由于测量误差而导致的评判失误。探索了叶面平展度量化的评价方法。该方法以充分展开叶片背面在30度入射光角度下垂直拍摄的数码照片为材料,以“100-标准偏差”作为叶片平展度的量化指标,该数值越接近100,叶面越平展;该数值越小,叶面平展度越差。提出以充分展开叶片的亮度标准差与母株该部位叶片的亮度标准差的比值大于1.3作为判别变异发生的临界值,是否合适有待研究。
The protocal for increasing the ployploid inducing rates were studied by using diploid grapes of Qingdao Zaohong, Anshan Zaohong, and Rizamat, and their VM1 and VM2 progeny were studied on patterns of variation. The autotetraploid Muscat Hamburg, which was obtained by inducement of Colchicine, was studied on variation of rooting ability of green cuttings and leave photosynthesis, and on some indexes for polyploid evaluation. The main results were as followed.
1. Improving of protocal in shoot tips inducement of Colchicine
The shoot tips survival rates after treated with Colchicine were increased by using the combination of moisture keeping bags, newspaper bags, and shading net, and removing these materials gradually. The surving rate could reach higher than 90%. The primary selection technic for VM1 progeny was systematically improved, and the technic for fast seclection for VM1 and VM2 was as also. It was found that the polyploid plants obtained from inducement of Colchicine had a big varivation after asexual propagation. The amount of VM2 plants should be increased for selection. There were 13 new polyploid strains of Anshan Zaohong and Rizamat grapes obtained in this experiment.
2. Systematical studing in the variation patterns after shoot tips inducement of Colchicine
Significant variation was found in vein macro-structure of leaves in autotetraploid grapes. The dominant feature of main vein was abreaction. The symmetry of side veins along the main veins was weak. More leaves became cuniform or nephroid in shape, and had less veins.
A broad variation was found among the VM1 progeny in terms of the leaves macro-structure or micro-structure after the inducement of Colchicine, including thirteen aspects in macro-structure such as leaf shape, size, vein, margin, etc. New patterns of leaves occurred.
3. Variation of autotetraploid grapes in rooting characteristics
(1) In the aspect of morphology, the green cuttings of autotetraploid grapes had poor rooting ability compared to their mother plants in terms of their adventitious rooting rate, adventitious roots number per cuttings, and lengh of total adventitious roots.
(2) In the aspect of anatomy, the green cuttings of autotetraploid grapes had larger medullary area and fewer medullary rays. During the phase of primary root inducement, its epidermal cells and cortex cells differentiation too early and too much, that led this phase last for one day longer; and during the phase of primary root development, the top of the primary roots became flat, smooth, and big in size. That could be the reason to lead the poor rooting ability of the autotetraploid Muscat Hamburg. The development of the rooting of green cuttings in autotetraploid grapes could be divided into three phases as primary root inducement, primary root development, and adventitious root development.
(3) In the aspect of physiology, the carbon nutrition, phenolics, flavone, enzyme activities of IAAO, POD, and PPO, and hormone changed significantly in the autotetraploid Muscat Hamburg during the process of the rooting of green cuttings.①The content of soluble solid sugar in the autotetraploid Muscat Hamburg was lower than that of the diploid Muscat Hamburg all the time during the rooting process. The content of starch in the autotetraploid Muscat Hamburg was higher than that of the diploid Muscat Hamburg at the beginning of the primary roots inducement, but lower than that during the late of the primary roots inducement with different changing tendency. The content was lower for the autotetraploid during the 3rd day to 15th day period.
②The content of flavone in cortex at basal part of the green cuttings in autotetraploid was significant higher at the beginning of the primary root inducement phase than that of the diploid, and lower at the late period with different changing tendency. The content in the autotetraploid was lower than that of the diploid after the 3rd day. The content of phenolics in the autotetraploid was higher in the beginning of the primary root inducement phase than that of the diploid and lower after the 3rd.
③During the primary root inducement phase, the activity of IAAO in the autotetraploid was significant higher than that of the diploid; the activity of POD in the autotetraploid was significant lower than that of the diploid; and the activity of PPO in the autotetraploid was significant higher than that of the diploid and lower than that at the late period.
④It was not only the hormone content in cortex of cuttings was changed during the different phases of cutting’s rooting, but also the reaction of that part to NAA applied was changed after the double of its chromosome. During the adventitious inducement period (0~3 d), the content of Auxin that supposed to promote the rooting decreased and the content of ABA and GA that supposed to inhibit the rooting increased, and the ratio of Auxin to ABA and GA dropped. With application of NAA, the content of GA, ZT, and ZR did not decrease as normal but increase, and the tendency of hormone was changed.
4. Variation of photosynthesis characteristics in autopolyploidy
The content of chlorophyll of the autotetraploid Muscat Hamburg increased, the lengh and width of cells in palisade tissue increased, and the layers of cells in palisade tissue decreased compared to the diploid plants. Its photosynthesis characteristics had a significant variation.①The optimal temperature for photosynthesis of autotetraploid Muscat Hamburg was 20℃. Under controlled temperature condition, the autotetraploid Muscat Hamburg No.2 showed highest sensitivity to the changing of temperature with highest ability of tolerating to the high temperature. The No. 1 and diploid plants showed less sensitivity to the changing of temperature.
②Under natural condition, the patterns of photosynthesis of both autotetraploid Muscat Hamburg and diploid Muscat Hamburg were double peak curves. For the autotetraploid Muscat Hamburg No.1 and diploid Muscat Hamburg, the first peak occurred at 9:00 and the second peak occurred at 14:00, and the value of the first peak was higher than that of the second one. During 9:00 to 11:00, the net photosynthetic rate did not decrease significantly. The first peak of the autotetraploid Muscat Hamburg No.2 occurred at 9:00 and the second peak occurred at 13:00, and the value of the first peak was about the same as the second. During 9:00 to 11:00, the net photosynthetic rate decreased significantly. Then the net photosynthetic rate of the autotetraploid Muscat Hamburg No.2 was low. The transpiration rate, stomatal conductance, and net photosynthetic rate for differnet ploidy plants were as M4x-1﹥M2x﹥M4x-2.
5. Modification of some evaluation methods for polyploid selection
The slide making process for the stoma observation was simplified for easy-to-do and having better result, which could increase the efficiency. A new method for mesuring alabastrum size was developed with higher precision. Leaf flat grade was applied to the variation evaluation. The procedure was that: taking picture of lower surface of leaf from 30 degree incident light angle by using digital camera, caculating its leaf flat grade as“100-standard deviation”, and it was considered as variation occurred when the ratio of offsprings to its mother plants of the some position leaves reached 1.3.
1、进一步完善了秋水仙素茎尖诱变技术
在系统研究的基础上,秋水仙素诱变葡萄茎尖的成活率较对照提高了3.9%～13.3%,其中“秋水仙素处理+保湿袋+报纸袋+遮阳网”以及处理后逐渐解除保护的方法,诱变节位的成活率达到了90.6%;系统完善了VM1代的初次选择技术,以及VM1代和VM2代的快速分离技术;提出了秋水仙素诱变获得的葡萄多倍体材料,在无性繁殖过程中存在较大幅度的变异,在后代筛选时应增加VM2代的群体数量。创新了13份鞍山早红和力扎马特葡萄同源多倍体种质资源。
2、系统研究了秋水仙素茎尖诱变不同世代的变异规律
同源四倍体葡萄的叶脉结构发生了显著的变异,其主脉的主导地位被消弱,主脉上分生的侧脉对称性降低,叶片形状变为楔形或肾形比率显著增加,叶脉数量明显减少。利用秋水仙素诱变葡萄茎尖后,其VM1代叶片的宏观结构和微观结构均发生了广泛的变异。其中叶片形状、大小、叶脉、叶缘等13个宏观性状的变异幅度增加,出现了一些新的叶片类型。
3、探明了同源多倍体葡萄生根特性的变异
(1)在形态学方面,同源四倍体葡萄嫩枝扦插生根能力较母株显著下降,扦插不定根发生率、平均每个插条的不定根数量和不定根总长度均显著低于其二倍体母株。
(2)在解剖学方面,同源四倍体玫瑰香嫩枝插条的髓部面积增大、髓射线数量减少;在根原基诱导阶段,其表皮细胞和皮层细胞过早、过量分化,根原基诱导期延长了1d;在根原基发育期,其根原基体积变大、顶部变平。上述形态与解剖性状的变异可能是导致同源四倍体玫瑰香生根能力下降的原因。并提出,将同源四倍体葡萄插条不定根发育可划分为根原基诱导期、根原基发育期和不定根发育期三个阶段。
(3)在生理学方面,同源四倍体玫瑰香嫩枝插条在扦插生根过程中的碳素营养、酚类和黄酮物质、IAAO酶活性、POD酶活性、PPO酶活性和内源激素代谢发生了显著的变异:①同源四倍体玫瑰香在扦插初期的可溶性糖含量、淀粉含量显著低于二倍体玫瑰香。同源四倍体玫瑰香的淀粉含量在根原基诱导期与二倍体玫瑰香品种的变化态势相反,且在扦插第3～12d其淀粉含量始终低于二倍体玫瑰香。②同源四倍体玫瑰香嫩枝插条基部皮层的黄酮含量在根原基诱导初期显著高于二倍体玫瑰香、后期又显著低于二倍体玫瑰香,两个不同倍性品种的变化态势相反。同源四倍体玫瑰香的多酚含量在根原基诱导初期高于二倍体玫瑰香,第3d以后低于二倍体玫瑰香。③在根原基诱导期,同源四倍体玫瑰香的IAAO酶活性显著高于二倍体玫瑰香,POD酶活性显著低于二倍体玫瑰香,PPO酶活性高于二倍体玫瑰香,而在此期以后PPO酶活性又低于二倍体玫瑰香。④玫瑰香葡萄同源加倍以后,不仅改变了不同时期插条生根部位皮层的内源激素水平,也改变了该部位对外施NAA的反应。在不定根诱导期(0～3 d),促进生根的生长素水平下降,抑制生根的脱落酸和赤霉素含量上升,生长素与脱落酸和赤霉素的比值下降;应用NAA处理以后,在根原基诱导期(0～3 d),赤霉素、玉米素和玉米素核苷含量不降反升,改变了内源激素的整体变化态势。
4、探讨了同源多倍体葡萄光合特性的变异
同源四倍体玫瑰香较二倍体玫瑰香单位面积的叶绿素含量增大,栅栏组织细胞长度与宽度增加,单位叶片解剖长度内栅栏组织细胞层数减少。其光合特性发生了显著变异:①同源四倍体玫瑰香最适的光合温度为20℃左右。在温度诱导条件下,同源四倍体玫瑰香2号对温度的敏感性最强,表现具有较强的耐高温能力。同源四倍体玫瑰香1号和二倍体玫瑰香对温度的反应较迟钝。②在自然条件下,同源四倍体玫瑰香与二倍体玫瑰香的光合作用曲线均为双峰曲线。不同倍性玫瑰香葡萄的蒸腾速率、气孔导度和净光合速率为:M4x-1﹥M2x﹥M4x-2。同源四倍体玫瑰香1号和二倍体玫瑰香光合作用曲线相似,第一个峰值均出现在上午9:00时,第二个峰值均出现在14:00时,第一个峰值大于第二个峰值,且9:00～11:00时的净光合速率下降不明显;同源四倍体玫瑰香2号的第一个峰值也出现在上午9:00时,但第二个峰值出现在13:00,第一个峰值与第二个峰值相当,且9:00～11:00净光合速率明显下降。
5、探索改进了部分性状的评价方法
简化了观测气孔的制片方法;该方法简便易行,效果良好,提高了气孔性状观测效率。提出了一种测量花蕾大小的新方法;应用这种方法可以准确测量花蕾的大小,消除由于测量误差而导致的评判失误。探索了叶面平展度量化的评价方法。该方法以充分展开叶片背面在30度入射光角度下垂直拍摄的数码照片为材料,以“100-标准偏差”作为叶片平展度的量化指标,该数值越接近100,叶面越平展;该数值越小,叶面平展度越差。提出以充分展开叶片的亮度标准差与母株该部位叶片的亮度标准差的比值大于1.3作为判别变异发生的临界值,是否合适有待研究。
The protocal for increasing the ployploid inducing rates were studied by using diploid grapes of Qingdao Zaohong, Anshan Zaohong, and Rizamat, and their VM1 and VM2 progeny were studied on patterns of variation. The autotetraploid Muscat Hamburg, which was obtained by inducement of Colchicine, was studied on variation of rooting ability of green cuttings and leave photosynthesis, and on some indexes for polyploid evaluation. The main results were as followed.
1. Improving of protocal in shoot tips inducement of Colchicine
The shoot tips survival rates after treated with Colchicine were increased by using the combination of moisture keeping bags, newspaper bags, and shading net, and removing these materials gradually. The surving rate could reach higher than 90%. The primary selection technic for VM1 progeny was systematically improved, and the technic for fast seclection for VM1 and VM2 was as also. It was found that the polyploid plants obtained from inducement of Colchicine had a big varivation after asexual propagation. The amount of VM2 plants should be increased for selection. There were 13 new polyploid strains of Anshan Zaohong and Rizamat grapes obtained in this experiment.
2. Systematical studing in the variation patterns after shoot tips inducement of Colchicine
Significant variation was found in vein macro-structure of leaves in autotetraploid grapes. The dominant feature of main vein was abreaction. The symmetry of side veins along the main veins was weak. More leaves became cuniform or nephroid in shape, and had less veins.
A broad variation was found among the VM1 progeny in terms of the leaves macro-structure or micro-structure after the inducement of Colchicine, including thirteen aspects in macro-structure such as leaf shape, size, vein, margin, etc. New patterns of leaves occurred.
3. Variation of autotetraploid grapes in rooting characteristics
(1) In the aspect of morphology, the green cuttings of autotetraploid grapes had poor rooting ability compared to their mother plants in terms of their adventitious rooting rate, adventitious roots number per cuttings, and lengh of total adventitious roots.
(2) In the aspect of anatomy, the green cuttings of autotetraploid grapes had larger medullary area and fewer medullary rays. During the phase of primary root inducement, its epidermal cells and cortex cells differentiation too early and too much, that led this phase last for one day longer; and during the phase of primary root development, the top of the primary roots became flat, smooth, and big in size. That could be the reason to lead the poor rooting ability of the autotetraploid Muscat Hamburg. The development of the rooting of green cuttings in autotetraploid grapes could be divided into three phases as primary root inducement, primary root development, and adventitious root development.
(3) In the aspect of physiology, the carbon nutrition, phenolics, flavone, enzyme activities of IAAO, POD, and PPO, and hormone changed significantly in the autotetraploid Muscat Hamburg during the process of the rooting of green cuttings.①The content of soluble solid sugar in the autotetraploid Muscat Hamburg was lower than that of the diploid Muscat Hamburg all the time during the rooting process. The content of starch in the autotetraploid Muscat Hamburg was higher than that of the diploid Muscat Hamburg at the beginning of the primary roots inducement, but lower than that during the late of the primary roots inducement with different changing tendency. The content was lower for the autotetraploid during the 3rd day to 15th day period.
②The content of flavone in cortex at basal part of the green cuttings in autotetraploid was significant higher at the beginning of the primary root inducement phase than that of the diploid, and lower at the late period with different changing tendency. The content in the autotetraploid was lower than that of the diploid after the 3rd day. The content of phenolics in the autotetraploid was higher in the beginning of the primary root inducement phase than that of the diploid and lower after the 3rd.
③During the primary root inducement phase, the activity of IAAO in the autotetraploid was significant higher than that of the diploid; the activity of POD in the autotetraploid was significant lower than that of the diploid; and the activity of PPO in the autotetraploid was significant higher than that of the diploid and lower than that at the late period.
④It was not only the hormone content in cortex of cuttings was changed during the different phases of cutting’s rooting, but also the reaction of that part to NAA applied was changed after the double of its chromosome. During the adventitious inducement period (0~3 d), the content of Auxin that supposed to promote the rooting decreased and the content of ABA and GA that supposed to inhibit the rooting increased, and the ratio of Auxin to ABA and GA dropped. With application of NAA, the content of GA, ZT, and ZR did not decrease as normal but increase, and the tendency of hormone was changed.
4. Variation of photosynthesis characteristics in autopolyploidy
The content of chlorophyll of the autotetraploid Muscat Hamburg increased, the lengh and width of cells in palisade tissue increased, and the layers of cells in palisade tissue decreased compared to the diploid plants. Its photosynthesis characteristics had a significant variation.①The optimal temperature for photosynthesis of autotetraploid Muscat Hamburg was 20℃. Under controlled temperature condition, the autotetraploid Muscat Hamburg No.2 showed highest sensitivity to the changing of temperature with highest ability of tolerating to the high temperature. The No. 1 and diploid plants showed less sensitivity to the changing of temperature.
②Under natural condition, the patterns of photosynthesis of both autotetraploid Muscat Hamburg and diploid Muscat Hamburg were double peak curves. For the autotetraploid Muscat Hamburg No.1 and diploid Muscat Hamburg, the first peak occurred at 9:00 and the second peak occurred at 14:00, and the value of the first peak was higher than that of the second one. During 9:00 to 11:00, the net photosynthetic rate did not decrease significantly. The first peak of the autotetraploid Muscat Hamburg No.2 occurred at 9:00 and the second peak occurred at 13:00, and the value of the first peak was about the same as the second. During 9:00 to 11:00, the net photosynthetic rate decreased significantly. Then the net photosynthetic rate of the autotetraploid Muscat Hamburg No.2 was low. The transpiration rate, stomatal conductance, and net photosynthetic rate for differnet ploidy plants were as M4x-1﹥M2x﹥M4x-2.
5. Modification of some evaluation methods for polyploid selection
The slide making process for the stoma observation was simplified for easy-to-do and having better result, which could increase the efficiency. A new method for mesuring alabastrum size was developed with higher precision. Leaf flat grade was applied to the variation evaluation. The procedure was that: taking picture of lower surface of leaf from 30 degree incident light angle by using digital camera, caculating its leaf flat grade as“100-standard deviation”, and it was considered as variation occurred when the ratio of offsprings to its mother plants of the some position leaves reached 1.3.
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