响叶杨三倍体和四倍体诱导技术研究
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摘要
鉴于多倍体育种在杨树遗传改良方面的优势,本研究选取我国重要的乡土树种响叶杨(Populus adenopoda Maxim.)为对象,针对南方低山丘陵地区对纸浆材新品种的需求问题,综合细胞遗传学与多倍体诱导技术,系统开展关于响叶杨天然未减数花粉发生机制、人工诱导雌配子及合子染色体加倍技术研究,为响叶杨多倍体育种奠定基础。主要研究结果如下：
1.在我国首次发现响叶杨可自然产生未减数2n花粉,并揭示了天然2n花粉的发生机制。花粉观察表明,来自四个省份的9个响叶杨无性系2n花粉发生频率介于0.49-12.56%。利用醋酸洋红涂片结合免疫荧光技术证明,小孢子母细胞减数分裂后期Ⅱ的纺锤体微管缺失导致姊妹染色单体不分离,以及减数分裂末期Ⅱ成膜体微管缺失导致胞质分裂异常,是响叶杨2n花粉发生的细胞学机制。
2.在建立响叶杨雌花芽外部形态与大孢子发生进程对应关系的基础上,以雌花芽发育形态为参照适时施加理化处理,从这一途径诱导获得响叶杨三倍体19株,并初步确定了施加处理的有效时期与处理强度等技术条件。其中,采用高温处理诱导响叶杨大孢子染色体加倍时,以雌花芽1/4花序露出芽鳞,即大部分细胞处于减数分裂双线期-终变期时,施加38-41℃持续处理4-6h的效果较好,最高诱导效率达40%；利用秋水仙碱溶液处理时,当雌花芽花序微露出芽鳞,即大孢子母细胞减数分裂主要处于粗线期附近,施加0.3%浓度的秋水仙碱持续处理24h可取得较好的加倍效果。
3.在确定响叶杨最佳授粉时期的基础上,通过施加理化处理抑制胚囊有丝分裂过程中的染色体分离,诱导获得胚囊染色体加倍的响叶杨三倍体子代381株,建立了高效的胚囊染色体加倍选育响叶杨三倍体技术,并明确了处理温度、浓度以及处理时间等条件。其中,施加高温处理诱导胚囊染色体加倍时,以授粉后18-24h雌花序施加38℃持续处理4-6h或41℃持续处理2-4h的条件为宜,三倍体诱导率最高达83.33%。细胞学观察显示,三倍体得率与单核胚囊,二核胚囊及四核胚囊期细胞所占的比例均呈正相关性,表明高温处理对响叶杨胚囊发育的三次有丝分裂具有加倍机会。施加秋水仙碱处理诱导响叶杨胚囊染色体加倍,以授粉后12-18h的雌花序施加0.3%浓度秋水仙碱持续处理24h的效果最好,结合细胞学进行相关分析显示,三倍体诱导率与单核胚囊期及二核胚囊期细胞比例存在正相关。
4.根据响叶杨雌花序授粉时间及子房内絮状纤维发育状态的综合指标,对合子发育进程进行即时判别,并在此基础上对合子第一次有丝分裂施加高温和秋水仙碱处理,诱导获得响叶杨四倍体子代43株。其中,施加高温处理诱导四倍体时,以絮状纤维开始发生时施加44℃高温持续处理2h的诱导效果较佳,最高诱导率可达14.12%。采用秋水仙碱浸泡处理诱导四倍体时,在雌花序授粉5d后,且絮状纤维尚未发生时施加0.3%的秋水仙碱持续处理30h的效果最好。
5.响叶杨三倍体气孔属性、叶片特征、光合指数及内源激素的分析结果显示,三倍体由于染色体数目的增加,导致其在气孔大小、叶片大小、光合速率和生长刺激型激素含量平均值均高于同一杂交组合的二倍体对照植株,而胞间CO2浓度及生长抑制型激素ABA的含量低于二倍体对照。
6.对响叶杨三倍体子代2年生实生苗生长情况的调查结果显示,同一生长条件下的三倍体植株在生长方面与二倍体对照平均值相比优势明显。其中,在贵阳与湖南的响叶杨杂交组合中,平均苗高与地径生长量分别超出二倍体子代平均值的17%和38%：在贵阳的响叶杨杂交组合中,在平均苗高和地径生长分别超出二倍体子代平均值的44%和47%。
Populus adenopoda Maxim, is a native tree species that is often used as timber sources to the mountainous regions of southern China. Polyploid breeding has been proven to be a valuable approach in populus improvement. In the present paper, P. adenopoda was employed as polyploid breeding materials, in order to provide new populus varieties of puplpwood plantation for those regions. The occurrence of spontaneous2n pollen and its cytological mechanisms, chromosome doubling of female gametes and zygote of P. adenopoda were studied by combining the cytogenetics and techniques of polyploid breeding. And these investigations provide a platform for polyploid breeding of P. adenopoda. The main results are described as follows:
1. The production of spontaneous2n pollen in P. adenopoda was firstly investigated in China, and its cytological mechanisms were revealed. Nine clones of P. adenopoda collected from four provinces could produce2n pollen at different frequency, and varying from0.49%to12.56%. The meiosis analysis of anthers by squashed technique and indirect immunofluorescence showed that the non-disjunction of sister chromatid resulted from lack of spindles microtubule in anaphase Ⅱ, and the partial failure of cytokinesis caused by the absence of phragmoplasts in telophase Ⅱ, were the main cytological mechanism for2n pollen formation in P. adenopoda.
2. Based on the relationship between flower bud morphological characteristics and the female meiotic stage of P. adenopoda, the female bud at different morphological characteristics were treated with heat and colchicine to induce megaspore chromosome doubling. In the resulting progeny,19triploids were obtained. The optimal periods and treatment intensity with high temperature and colchicine in P. adenopoda were determined respectively. When approximately one fourth of the catkins protruded from the bract scales, cytological observation revealed that most megaspore mother cells were at the period from diplotene to diakinesis, was the optimal period for megaspore chromosome doubling induced by high temperature. The treatments with38-41℃for4-6h were efficient, and the highest efficiency of triploid production was40.0%. When the catkins emerged from bract scales slightly, cytological observation showed that major cells were in pachytene stage, which was the optimal period for megaspore chromosome doubling induced by colchicine. The appropriate treatment condition was immersed the flower buds with0.3%colchicine solution for24h.
3. On the basis of the optimal pollinated time was determined in P. adenopoda, the pollinated female catkins during embryo sac development were treated with high temperature and colchicine to induce embryo sac chromosome doubling. In the offspring seedlings,381triploids were detected. The optimal treatment periods and treatment conditions for embryo sac chromosome doubling by high temperature and colchicine in P. adenopoda were investigated respectively. When high temperature was employed to induce embryo sac chromosome doubling, the treatments with38℃for4-6h or with41℃for2-4h during18-24h after pollination were both effective, and the highest frequency of triploid production was83.33%. Furthermore, correlation analyses combined with cytological observation showed that the percentage of triploid production was positively correlated with the rate of uni-, two-and four-nucleate embryo sacs, revealed that three mitotic divisions in embryo sac development provide chances for embryo sac chromosome doubling induced by high temperature. When the colchicine solution was used, the optimal treatment combination was immersed the female catkins during12-18h after pollination with0.3%colchicine solution for24h. Further correlation analyses combined with cytological observation showed that the percentage of triploid production was positively correlated with the rate of uni-and two-nucleate embryo sacs.
4. The combination between pollination time of female catkins and developmental changes of the cottonlike fiber in ovaries were established to mark the zygote division stage of P. adenopoda, and43tetraploids were produced by inhibiting the first zygote division with high temperature exposure and colchicine. When treated by high temperature, the cottonlike fibers in the ovary start to enclose the funiculus was the efficient state for zygote chromosome doubling, and the highest rate of tetraploid production was14.12%by treating the catkins with44℃for2h. When treated with colchicine, the female catkin5d after pollination and the cottonlike fiber in the ovary did not occur was optimal stage for zygote chromosome doubling, and soaking the pollinated catkins with0.3%colchicine solution for30h was effective.
5. The comparative study on the stomatal features, leaf characteristic, photosynthetic index and endogenous hormone between triploids and diploids of P. adenopoda was conducted. The results showed that the stomatal size, leaf size, photosynthetic rate and the content of stimulated hormones in triploid offsprings were higher than that of the diploids in average. However, the concentration of intercellular CO2and the content of inhibited hormones in triploids were lower than that of diploid controls.
6. The investigation of stem height and basal diameter of two-year-old diploids and triploids of P. adenopoda revealed that triploids had significant advantages in growth compared with diploid plants. Cross combination between Guiyang and Hunan, the average values of stem height and basal diameter in triploids exceeded the diploids by17%and38%, respectively. In Guiyang cross combination, the average values of stem height and basal diameter in triploids exceeded the diploids by44%and47%, respectively.
1.在我国首次发现响叶杨可自然产生未减数2n花粉,并揭示了天然2n花粉的发生机制。花粉观察表明,来自四个省份的9个响叶杨无性系2n花粉发生频率介于0.49-12.56%。利用醋酸洋红涂片结合免疫荧光技术证明,小孢子母细胞减数分裂后期Ⅱ的纺锤体微管缺失导致姊妹染色单体不分离,以及减数分裂末期Ⅱ成膜体微管缺失导致胞质分裂异常,是响叶杨2n花粉发生的细胞学机制。
2.在建立响叶杨雌花芽外部形态与大孢子发生进程对应关系的基础上,以雌花芽发育形态为参照适时施加理化处理,从这一途径诱导获得响叶杨三倍体19株,并初步确定了施加处理的有效时期与处理强度等技术条件。其中,采用高温处理诱导响叶杨大孢子染色体加倍时,以雌花芽1/4花序露出芽鳞,即大部分细胞处于减数分裂双线期-终变期时,施加38-41℃持续处理4-6h的效果较好,最高诱导效率达40%；利用秋水仙碱溶液处理时,当雌花芽花序微露出芽鳞,即大孢子母细胞减数分裂主要处于粗线期附近,施加0.3%浓度的秋水仙碱持续处理24h可取得较好的加倍效果。
3.在确定响叶杨最佳授粉时期的基础上,通过施加理化处理抑制胚囊有丝分裂过程中的染色体分离,诱导获得胚囊染色体加倍的响叶杨三倍体子代381株,建立了高效的胚囊染色体加倍选育响叶杨三倍体技术,并明确了处理温度、浓度以及处理时间等条件。其中,施加高温处理诱导胚囊染色体加倍时,以授粉后18-24h雌花序施加38℃持续处理4-6h或41℃持续处理2-4h的条件为宜,三倍体诱导率最高达83.33%。细胞学观察显示,三倍体得率与单核胚囊,二核胚囊及四核胚囊期细胞所占的比例均呈正相关性,表明高温处理对响叶杨胚囊发育的三次有丝分裂具有加倍机会。施加秋水仙碱处理诱导响叶杨胚囊染色体加倍,以授粉后12-18h的雌花序施加0.3%浓度秋水仙碱持续处理24h的效果最好,结合细胞学进行相关分析显示,三倍体诱导率与单核胚囊期及二核胚囊期细胞比例存在正相关。
4.根据响叶杨雌花序授粉时间及子房内絮状纤维发育状态的综合指标,对合子发育进程进行即时判别,并在此基础上对合子第一次有丝分裂施加高温和秋水仙碱处理,诱导获得响叶杨四倍体子代43株。其中,施加高温处理诱导四倍体时,以絮状纤维开始发生时施加44℃高温持续处理2h的诱导效果较佳,最高诱导率可达14.12%。采用秋水仙碱浸泡处理诱导四倍体时,在雌花序授粉5d后,且絮状纤维尚未发生时施加0.3%的秋水仙碱持续处理30h的效果最好。
5.响叶杨三倍体气孔属性、叶片特征、光合指数及内源激素的分析结果显示,三倍体由于染色体数目的增加,导致其在气孔大小、叶片大小、光合速率和生长刺激型激素含量平均值均高于同一杂交组合的二倍体对照植株,而胞间CO2浓度及生长抑制型激素ABA的含量低于二倍体对照。
6.对响叶杨三倍体子代2年生实生苗生长情况的调查结果显示,同一生长条件下的三倍体植株在生长方面与二倍体对照平均值相比优势明显。其中,在贵阳与湖南的响叶杨杂交组合中,平均苗高与地径生长量分别超出二倍体子代平均值的17%和38%：在贵阳的响叶杨杂交组合中,在平均苗高和地径生长分别超出二倍体子代平均值的44%和47%。
Populus adenopoda Maxim, is a native tree species that is often used as timber sources to the mountainous regions of southern China. Polyploid breeding has been proven to be a valuable approach in populus improvement. In the present paper, P. adenopoda was employed as polyploid breeding materials, in order to provide new populus varieties of puplpwood plantation for those regions. The occurrence of spontaneous2n pollen and its cytological mechanisms, chromosome doubling of female gametes and zygote of P. adenopoda were studied by combining the cytogenetics and techniques of polyploid breeding. And these investigations provide a platform for polyploid breeding of P. adenopoda. The main results are described as follows:
1. The production of spontaneous2n pollen in P. adenopoda was firstly investigated in China, and its cytological mechanisms were revealed. Nine clones of P. adenopoda collected from four provinces could produce2n pollen at different frequency, and varying from0.49%to12.56%. The meiosis analysis of anthers by squashed technique and indirect immunofluorescence showed that the non-disjunction of sister chromatid resulted from lack of spindles microtubule in anaphase Ⅱ, and the partial failure of cytokinesis caused by the absence of phragmoplasts in telophase Ⅱ, were the main cytological mechanism for2n pollen formation in P. adenopoda.
2. Based on the relationship between flower bud morphological characteristics and the female meiotic stage of P. adenopoda, the female bud at different morphological characteristics were treated with heat and colchicine to induce megaspore chromosome doubling. In the resulting progeny,19triploids were obtained. The optimal periods and treatment intensity with high temperature and colchicine in P. adenopoda were determined respectively. When approximately one fourth of the catkins protruded from the bract scales, cytological observation revealed that most megaspore mother cells were at the period from diplotene to diakinesis, was the optimal period for megaspore chromosome doubling induced by high temperature. The treatments with38-41℃for4-6h were efficient, and the highest efficiency of triploid production was40.0%. When the catkins emerged from bract scales slightly, cytological observation showed that major cells were in pachytene stage, which was the optimal period for megaspore chromosome doubling induced by colchicine. The appropriate treatment condition was immersed the flower buds with0.3%colchicine solution for24h.
3. On the basis of the optimal pollinated time was determined in P. adenopoda, the pollinated female catkins during embryo sac development were treated with high temperature and colchicine to induce embryo sac chromosome doubling. In the offspring seedlings,381triploids were detected. The optimal treatment periods and treatment conditions for embryo sac chromosome doubling by high temperature and colchicine in P. adenopoda were investigated respectively. When high temperature was employed to induce embryo sac chromosome doubling, the treatments with38℃for4-6h or with41℃for2-4h during18-24h after pollination were both effective, and the highest frequency of triploid production was83.33%. Furthermore, correlation analyses combined with cytological observation showed that the percentage of triploid production was positively correlated with the rate of uni-, two-and four-nucleate embryo sacs, revealed that three mitotic divisions in embryo sac development provide chances for embryo sac chromosome doubling induced by high temperature. When the colchicine solution was used, the optimal treatment combination was immersed the female catkins during12-18h after pollination with0.3%colchicine solution for24h. Further correlation analyses combined with cytological observation showed that the percentage of triploid production was positively correlated with the rate of uni-and two-nucleate embryo sacs.
4. The combination between pollination time of female catkins and developmental changes of the cottonlike fiber in ovaries were established to mark the zygote division stage of P. adenopoda, and43tetraploids were produced by inhibiting the first zygote division with high temperature exposure and colchicine. When treated by high temperature, the cottonlike fibers in the ovary start to enclose the funiculus was the efficient state for zygote chromosome doubling, and the highest rate of tetraploid production was14.12%by treating the catkins with44℃for2h. When treated with colchicine, the female catkin5d after pollination and the cottonlike fiber in the ovary did not occur was optimal stage for zygote chromosome doubling, and soaking the pollinated catkins with0.3%colchicine solution for30h was effective.
5. The comparative study on the stomatal features, leaf characteristic, photosynthetic index and endogenous hormone between triploids and diploids of P. adenopoda was conducted. The results showed that the stomatal size, leaf size, photosynthetic rate and the content of stimulated hormones in triploid offsprings were higher than that of the diploids in average. However, the concentration of intercellular CO2and the content of inhibited hormones in triploids were lower than that of diploid controls.
6. The investigation of stem height and basal diameter of two-year-old diploids and triploids of P. adenopoda revealed that triploids had significant advantages in growth compared with diploid plants. Cross combination between Guiyang and Hunan, the average values of stem height and basal diameter in triploids exceeded the diploids by17%and38%, respectively. In Guiyang cross combination, the average values of stem height and basal diameter in triploids exceeded the diploids by44%and47%, respectively.
引文
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