陆地棉花粉氮离子注入诱变效应及机理研究
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摘要
离子束生物工程技术是一种新的生物技术,它通过离子束注入生物细胞,来研究其生物学效应和作用机理,用于诱变育种和基因工程等方面。1986年首次将离子束注入技术应用于诱变育种,通过离子束注入诱变,在多种植物、微生物中已获得各种不同类型的突变体。但是,离子注入对细胞的作用机理尚不清楚,离子的穿透能力很弱,离子能否进入生物细胞及诱发基因发生突变仍然是有争议的问题。本文主要研究氮离子束注入棉花花粉引起的生物学效应。
本研究采用能量为20keV的低能氮离子,分别以不同剂量(100单位(0.26×10~(16)N~+/cm~2)、150单位(0.39×10~(16)N~+/cm~2)、200单位(0.52×10~(16)N~+/cm~2)、300单位(0.78×10~(16)N~+/cm~2))的氮离子注入陆地棉(Gossypium hirsutum)品种苏棉12号的成熟花粉,用处理花粉授粉,收获种子。对处理花粉的活力、花粉表面结构、内部结构的变化、花粉萌发率、花粉管在花柱中的生长速度、受精情况等进行系统分析。用SSR分子标记的方法检测了离子注入花粉对于授粉后胚珠DNA变异的程度,用抑制性消减杂交法分析离子注入花粉授粉后代的基因表达差异。并分析离子注入花粉授粉后的农艺形状变异。主要结果如下。
用花粉原位萌发法、联苯胺-甲萘酚染色法、花粉管快速萌发法、FDA法(荧光素二醋酸脂法)、TTC(2,3,5-氯代三苯基四氮唑)染色法、离体萌发法等6种方法分别测定对照花粉、抽真空处理花粉及N~+注入后的棉花花粉活力。TTC染色法不能有效检测自然花粉及N~+注入后的花粉的活力,测定结果均为0%。除了TTC染色法外,其它5种方法均能测定自然花粉的活力,测定结果均为98%左右。花粉管快速萌发法不能准确测定经抽真空处理后的花粉的活力。抽真空处理对花粉的活力无明显影响。联苯胺-甲萘酚染色法、花粉管快速萌发法、TTC染色法、离体萌发法等4种方法均不能用来准确测定N~+注入后的棉花花粉活力。FDA荧光染色法测定的花粉活力结果稳定,重复性好,与原位萌发法测定的结果一致,是快速测定经N~+注入处理后的棉花花粉活力的简便方法。N~+注入花粉后,随着离子注入剂量的增加,花粉活力呈明显下降趋势。
通过荧光显微镜观察棉花花粉活体原位萌发(柱头上萌发)和离体条件下的原位萌发状况,建立准确的观察花粉粒在柱头上的萌发和花柱中花粉管的生长状况的简便的方法,并找出棉花花粉萌发的最适培养温度。结果表明:花粉在原位和离体情况下的原位萌发时间是基本一致的。活体原位萌发在授粉1 h后极个别的花粉粒开始萌发,4 h后柱头上的花粉粒全部萌发。12 h后大量的花粉管生长至花柱基部;离体原位萌发在授粉2 h后开始有花粉粒萌发,13 h后大量的花粉管长至花柱基部。授粉后,活体原位萌发较原位萌发晚1b。此外,在花粉管生长的同时,花柱也在生长。花粉萌发的合适培养温度为25-35℃。而在培养温度为30℃时,花柱中花粉管数量最多。
通过扫描电镜、透射电镜、原位萌发和石蜡切片等方法观察了N~+注入棉花花粉后产生的各种变化,初步研究了不同剂量的N~+注入棉花花粉的诱变机理及诱变后产生的一系列生物学效应。结果表明:随着离子注入剂量的增大,离子注入使表面结构破损的花粉粒的数量逐渐增加,离子注入对于花粉表面有明显的刻蚀作用,可产生不同大小和深度的孔洞和裂缝;花粉粒内涵物的量逐渐减少,排列致密性降低;离子注入剂量越高,花粉粒内部出现的空隙越多,造粉质体的体积变大,数量也越多;离子注入会明显降低花粉的萌发率;N~+注入花粉授粉后,花柱中花粉管的数量明显下降,注入剂量愈大,花柱中花粉管的数量愈少,对照花粉授粉后花粉管的数量为:320±42条,剂量为100单位、150单位、200单位和300单位的N离子注入花粉后,花粉管的数量分别为:145±19.8、114±12.7、94±11.4和51.3±11条。
离子注入不影响授粉后雌雄配子受精。不同处理的花粉管生长速度是一致的,授粉后13小时,花粉管都能生长至花柱基部。不同剂量处理的花粉粒萌发后的受精时间是一致的。N~+注入花粉授粉后,对子房的生长有明显的影响,子房重量和直径都随注入剂的增加而降低。剂量为100单位、200单位和300单位的N~+注入花粉授粉后,SSR分子标记结果表明,胚珠的DNA多态性发生一定程度的改变。剂量为300单位的N~+注入花粉授粉后,SSH分析获得1个与已知功能基因同源序列,与EST库中的序列同源的4条EST序列。离子注入花粉授粉后成铃率降低,对M_1代农艺形状也产生变异,但变异程度是随机的,与离子注入剂量之间无一定的规律。本研究结果充分说明N~+注入花粉,会通过刻蚀作用进入细胞使细胞结构发生改变,改变细胞内的遗传物质,降低花粉的活力,降低子房的生长速度,使后代农艺形状发生变异,产生明显的生物学效应。
Ion beam bioengineering technology has been considered as a new method for mutation breeding.With low-energy ions impanted organisms, the biological effects and the function mechanisms of the technology were investigated by some researchers,and the technology has been used into genentic breeding and gene engineering. The researchers have observed some biological effects and revealed some basic mechanisms for the past decade,and the great development and application have been obtained .In this paper ,the biological effects of the cotton pollon implanted by nitrogen ions were studied and discussed.
The biological effect of Sumian 12 pollen implanted by N ion beams were studied in present thesis . The viability of Sumian 12 pollen were implanted by nitrogen ion beam (the energy was 20KeV, the doses were 100 units( 0.26×10~(16) N~+/cm~2), 150 uints(0.39×10~(16) N~+/cm~2), 200 units(.52×10~(16) N~+/cm~2)、300 units(0.78×10~(16) N~+/cm~2) respectively,no-treated was control.) . The effects on viability of cotton pollen , the exterior and interior structure of pollen, pollen germination rate, the growth rate of the pollen tube, the situation of insemination, the mutation aroused by ions implantation in ovule using SSR and diversity gene of the offspring after ions implantation using SSHwre studied . The result showed that:
The viability of Sumian 12 pollen before and after nitrogen ion implantation were measured by six methods:In situ pollen germination, Benzidine-α-Naphthol method, pollen tube quick germination, FDA fluorescence method, TTC method and pollen in vitro germination. Both the results of pollen viability before and after N~+ injecting were zero, so TTC method was not suitable. We found that five methods, except TTC method, be suitable for measuring pollen viability before N~+ implantation, their results were all about 98%. Pollen tube quick germination couldn't measure the viability of pollen in vacuum, but we know that vacuum not influence pollen viability. Benzidine-α-Naphthol method, pollen tube quick germination and pollen in vitro germination were also incompetent for measuring the viability after N~+ injecting. The results of FDA fluorescence method were repeatable and consonant with the results of in situ pollen germination. In conclusion, FDA fluorescence method were competent for measuring the viability of cotton pollen before and after N~+ injecting. Our results showed a significant negative correlation between ion implantation and pollen viability.
The pollen germination on the stigma and pollen tube grow in the style by decolorized analine blue methed with fluorescence microscope were observed. A simple methed for truly research the pollen germination on the stigma and pollen tube growth and the suitable temperature of pollen germination were found .The result showed that: The beginning time of in situ pollen germination were consistent with the time of pollen in vitro germination. At 1 hour after pollination of in situ pollen germination, only few pollen grain germinated in stigma. At 4 hour after pollination of in situ pollen germination, all of the pollen grains on stigma were germination. At 12 hour after pollination of in situ pollen germination, large number of pollen tubes have grown to the end of the style. At 2 hour after pollination of in vitro pollen germination, few pollen grains begin germination. At 13 hour after pollination of in vitro pollen germination, a lot of pollen tubes have grown to the end of the style. Otherwise, the style waas growing while the pollen tubes growing. The suitable temperature for incubating pollen germination was 25～35℃. The most suitable temperature was 30℃.
In the study , all of the changes after N~+ injecting cotton pollen were observed by scanning electron microscope(SEM), transmission electron microscope(TEM), decolorized analine blue methed and paraffin section method. The radiation mechanism and its biologic effect after ion implantation were research . The result showed that: The number of break pollen grain were increasing with the increase of the N ion implantation dose, ion beam could distinctly cut pollen wall and produce different holes. The inclusion in pollen was decreasing with the increase of ion implanting dose, the higher the dose was , the more the interspace had, the larger of the starch granules volume were. Ion beam could distinctly decrease pollen germination rate. The number of pollen tubes in style declined after N ion implantation, after pollination, the number of pollen tubes of control pollen were 320±42, after implanting with dosages 100 units,150 units,200 units,300 units, the number of pollen tubes were 145±19.8,114±12.7、94±11.4,51.3±11 respectively. But the pollen tubes growth rate were accordant after implanting different dose. At 13 hours after pollination, all pollen tubes have grown to the end of the style. The fecundation time and insemination time were coinciden with different ion dose. Ovary weight and diameter size were reduce with the increase of implantation doses. SSR (simple sequence repeat) analysis indicated that the molecular genetic polymorphism of ovules, which being pollinated by N ion treated pollens, were changed. This result suggested that ion beam might cause some genetic changes of pollens. The SSH result showed: The BLASTN homology search analysis revealed that 3 nonredundant clones were classified into the following groups : 1 known function genes , 2 cotton ESTps of unknown function . The ion beam implanted could bring on M_1 genetic variation of agronomic character, but there was no orderliness between genetic variation of agronomic character and ion dosage. The result suggested that ion beam could enter into the inner of the cell and changed the structure and the genetic materials, reduced the viability of pollen and the growth rate of ovary, lead M_1 genetic variation of agronomic character and produce obviously biological effects.
本研究采用能量为20keV的低能氮离子,分别以不同剂量(100单位(0.26×10~(16)N~+/cm~2)、150单位(0.39×10~(16)N~+/cm~2)、200单位(0.52×10~(16)N~+/cm~2)、300单位(0.78×10~(16)N~+/cm~2))的氮离子注入陆地棉(Gossypium hirsutum)品种苏棉12号的成熟花粉,用处理花粉授粉,收获种子。对处理花粉的活力、花粉表面结构、内部结构的变化、花粉萌发率、花粉管在花柱中的生长速度、受精情况等进行系统分析。用SSR分子标记的方法检测了离子注入花粉对于授粉后胚珠DNA变异的程度,用抑制性消减杂交法分析离子注入花粉授粉后代的基因表达差异。并分析离子注入花粉授粉后的农艺形状变异。主要结果如下。
用花粉原位萌发法、联苯胺-甲萘酚染色法、花粉管快速萌发法、FDA法(荧光素二醋酸脂法)、TTC(2,3,5-氯代三苯基四氮唑)染色法、离体萌发法等6种方法分别测定对照花粉、抽真空处理花粉及N~+注入后的棉花花粉活力。TTC染色法不能有效检测自然花粉及N~+注入后的花粉的活力,测定结果均为0%。除了TTC染色法外,其它5种方法均能测定自然花粉的活力,测定结果均为98%左右。花粉管快速萌发法不能准确测定经抽真空处理后的花粉的活力。抽真空处理对花粉的活力无明显影响。联苯胺-甲萘酚染色法、花粉管快速萌发法、TTC染色法、离体萌发法等4种方法均不能用来准确测定N~+注入后的棉花花粉活力。FDA荧光染色法测定的花粉活力结果稳定,重复性好,与原位萌发法测定的结果一致,是快速测定经N~+注入处理后的棉花花粉活力的简便方法。N~+注入花粉后,随着离子注入剂量的增加,花粉活力呈明显下降趋势。
通过荧光显微镜观察棉花花粉活体原位萌发(柱头上萌发)和离体条件下的原位萌发状况,建立准确的观察花粉粒在柱头上的萌发和花柱中花粉管的生长状况的简便的方法,并找出棉花花粉萌发的最适培养温度。结果表明:花粉在原位和离体情况下的原位萌发时间是基本一致的。活体原位萌发在授粉1 h后极个别的花粉粒开始萌发,4 h后柱头上的花粉粒全部萌发。12 h后大量的花粉管生长至花柱基部;离体原位萌发在授粉2 h后开始有花粉粒萌发,13 h后大量的花粉管长至花柱基部。授粉后,活体原位萌发较原位萌发晚1b。此外,在花粉管生长的同时,花柱也在生长。花粉萌发的合适培养温度为25-35℃。而在培养温度为30℃时,花柱中花粉管数量最多。
通过扫描电镜、透射电镜、原位萌发和石蜡切片等方法观察了N~+注入棉花花粉后产生的各种变化,初步研究了不同剂量的N~+注入棉花花粉的诱变机理及诱变后产生的一系列生物学效应。结果表明:随着离子注入剂量的增大,离子注入使表面结构破损的花粉粒的数量逐渐增加,离子注入对于花粉表面有明显的刻蚀作用,可产生不同大小和深度的孔洞和裂缝;花粉粒内涵物的量逐渐减少,排列致密性降低;离子注入剂量越高,花粉粒内部出现的空隙越多,造粉质体的体积变大,数量也越多;离子注入会明显降低花粉的萌发率;N~+注入花粉授粉后,花柱中花粉管的数量明显下降,注入剂量愈大,花柱中花粉管的数量愈少,对照花粉授粉后花粉管的数量为:320±42条,剂量为100单位、150单位、200单位和300单位的N离子注入花粉后,花粉管的数量分别为:145±19.8、114±12.7、94±11.4和51.3±11条。
离子注入不影响授粉后雌雄配子受精。不同处理的花粉管生长速度是一致的,授粉后13小时,花粉管都能生长至花柱基部。不同剂量处理的花粉粒萌发后的受精时间是一致的。N~+注入花粉授粉后,对子房的生长有明显的影响,子房重量和直径都随注入剂的增加而降低。剂量为100单位、200单位和300单位的N~+注入花粉授粉后,SSR分子标记结果表明,胚珠的DNA多态性发生一定程度的改变。剂量为300单位的N~+注入花粉授粉后,SSH分析获得1个与已知功能基因同源序列,与EST库中的序列同源的4条EST序列。离子注入花粉授粉后成铃率降低,对M_1代农艺形状也产生变异,但变异程度是随机的,与离子注入剂量之间无一定的规律。本研究结果充分说明N~+注入花粉,会通过刻蚀作用进入细胞使细胞结构发生改变,改变细胞内的遗传物质,降低花粉的活力,降低子房的生长速度,使后代农艺形状发生变异,产生明显的生物学效应。
Ion beam bioengineering technology has been considered as a new method for mutation breeding.With low-energy ions impanted organisms, the biological effects and the function mechanisms of the technology were investigated by some researchers,and the technology has been used into genentic breeding and gene engineering. The researchers have observed some biological effects and revealed some basic mechanisms for the past decade,and the great development and application have been obtained .In this paper ,the biological effects of the cotton pollon implanted by nitrogen ions were studied and discussed.
The biological effect of Sumian 12 pollen implanted by N ion beams were studied in present thesis . The viability of Sumian 12 pollen were implanted by nitrogen ion beam (the energy was 20KeV, the doses were 100 units( 0.26×10~(16) N~+/cm~2), 150 uints(0.39×10~(16) N~+/cm~2), 200 units(.52×10~(16) N~+/cm~2)、300 units(0.78×10~(16) N~+/cm~2) respectively,no-treated was control.) . The effects on viability of cotton pollen , the exterior and interior structure of pollen, pollen germination rate, the growth rate of the pollen tube, the situation of insemination, the mutation aroused by ions implantation in ovule using SSR and diversity gene of the offspring after ions implantation using SSHwre studied . The result showed that:
The viability of Sumian 12 pollen before and after nitrogen ion implantation were measured by six methods:In situ pollen germination, Benzidine-α-Naphthol method, pollen tube quick germination, FDA fluorescence method, TTC method and pollen in vitro germination. Both the results of pollen viability before and after N~+ injecting were zero, so TTC method was not suitable. We found that five methods, except TTC method, be suitable for measuring pollen viability before N~+ implantation, their results were all about 98%. Pollen tube quick germination couldn't measure the viability of pollen in vacuum, but we know that vacuum not influence pollen viability. Benzidine-α-Naphthol method, pollen tube quick germination and pollen in vitro germination were also incompetent for measuring the viability after N~+ injecting. The results of FDA fluorescence method were repeatable and consonant with the results of in situ pollen germination. In conclusion, FDA fluorescence method were competent for measuring the viability of cotton pollen before and after N~+ injecting. Our results showed a significant negative correlation between ion implantation and pollen viability.
The pollen germination on the stigma and pollen tube grow in the style by decolorized analine blue methed with fluorescence microscope were observed. A simple methed for truly research the pollen germination on the stigma and pollen tube growth and the suitable temperature of pollen germination were found .The result showed that: The beginning time of in situ pollen germination were consistent with the time of pollen in vitro germination. At 1 hour after pollination of in situ pollen germination, only few pollen grain germinated in stigma. At 4 hour after pollination of in situ pollen germination, all of the pollen grains on stigma were germination. At 12 hour after pollination of in situ pollen germination, large number of pollen tubes have grown to the end of the style. At 2 hour after pollination of in vitro pollen germination, few pollen grains begin germination. At 13 hour after pollination of in vitro pollen germination, a lot of pollen tubes have grown to the end of the style. Otherwise, the style waas growing while the pollen tubes growing. The suitable temperature for incubating pollen germination was 25～35℃. The most suitable temperature was 30℃.
In the study , all of the changes after N~+ injecting cotton pollen were observed by scanning electron microscope(SEM), transmission electron microscope(TEM), decolorized analine blue methed and paraffin section method. The radiation mechanism and its biologic effect after ion implantation were research . The result showed that: The number of break pollen grain were increasing with the increase of the N ion implantation dose, ion beam could distinctly cut pollen wall and produce different holes. The inclusion in pollen was decreasing with the increase of ion implanting dose, the higher the dose was , the more the interspace had, the larger of the starch granules volume were. Ion beam could distinctly decrease pollen germination rate. The number of pollen tubes in style declined after N ion implantation, after pollination, the number of pollen tubes of control pollen were 320±42, after implanting with dosages 100 units,150 units,200 units,300 units, the number of pollen tubes were 145±19.8,114±12.7、94±11.4,51.3±11 respectively. But the pollen tubes growth rate were accordant after implanting different dose. At 13 hours after pollination, all pollen tubes have grown to the end of the style. The fecundation time and insemination time were coinciden with different ion dose. Ovary weight and diameter size were reduce with the increase of implantation doses. SSR (simple sequence repeat) analysis indicated that the molecular genetic polymorphism of ovules, which being pollinated by N ion treated pollens, were changed. This result suggested that ion beam might cause some genetic changes of pollens. The SSH result showed: The BLASTN homology search analysis revealed that 3 nonredundant clones were classified into the following groups : 1 known function genes , 2 cotton ESTps of unknown function . The ion beam implanted could bring on M_1 genetic variation of agronomic character, but there was no orderliness between genetic variation of agronomic character and ion dosage. The result suggested that ion beam could enter into the inner of the cell and changed the structure and the genetic materials, reduced the viability of pollen and the growth rate of ovary, lead M_1 genetic variation of agronomic character and produce obviously biological effects.
引文
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