唐菖蒲体细胞无性系变异与种质资源创新的研究
摘要
唐菖蒲(Gladiolus hybridus Hort)是鸢尾科(Iridaceae)唐菖蒲属(Gladiolus)多年生球茎类植物,是世界上著名的四大切花之一,唐菖蒲切花生产是我国花卉产业的重要组成部分。我国唐菖蒲种球主要从国外引进,国内培育的较少,目前的主栽品种仅为少数几个夏花型大花品种,品种间存在着遗传资源狭窄的现象,有关唐菖蒲的遗传基础研究也鲜有报道。利用体细胞无性系变异结合化学诱变及分子标记辅助育种,通过筛选,构建唐菖蒲突变体库,可为选育优良形态学和生理生化特点的突变体,培育具有自主知识产权的新品系或品种,扩大可利用种质资源的范围奠定基础。
本研究以唐菖蒲主栽品种“超级玫瑰”(G. Rose Supreme)为试材,通过对不同器官来源的外植体进行愈伤组织诱导,建立了唐菖蒲高频再生体系,同时通过化学诱变剂处理愈伤组织,扩大变异谱；利用形态学、细胞学观察结合ISSR分子标记技术,筛选唐菖蒲体细胞无性系突变株,为唐菖蒲遗传图谱的构建及育种奠定基础,同时也为以无性繁殖、获得纯系年限长且有性杂交困难的花卉定向育种提供新途径。研究的主要结果如下：
1.以唐菖蒲主栽品种“超级玫瑰”不同器官为外植体进行了诱导,结果表明用子球、花瓣及花梗作外植体均可诱导形成再生植株,用叶片、花丝做外植体没有得到再生植株。诱导唐菖蒲愈伤组织的最佳培养基为MS+2,4-D4.0mg·L-1+6-BA0.5mg·L1,诱导体细胞胚的最佳培养基为MS+2,4-D1.0mg·L-1+TDZ0.3mg·L-1,确定了TDZ是决定唐菖蒲体细胞胚产生的重要因子；首次对唐菖蒲各器官来源的体细胞胚的诱导及发育过程进行了研究。
2.用花瓣作外植体建立再生体系过程中,再生植株出现了色素重新表达现象,即由花瓣诱导的再生植株在幼苗发育过程中明显携带花瓣中原有色素现象；由花瓣愈伤组织可直接再发育成花瓣；由花梗做外植体也出现部分色素表达现象,且再生植株易出现嵌合体；而用子球茎作外植体则没有出现色素表达现象。自然情况下,花瓣无性系变异的频率高于花梗,揭示了唐菖蒲不同器官来源再生植株器官发生及形态建成的特点。
3.用秋水仙素不同浓度、时间处理唐菖蒲愈伤组织,确定了秋水仙素的半致死剂量为0.5mg·L-1,用半致死剂量的秋水仙素处理各器官来源的愈伤组织,共获得124株变异株,其中花瓣无性系的变异率最高,为56.95%,花梗次之,为21.1%,子球茎最低,为12.6%,变异株形态上比对照明显粗壮。本研究根据细胞学检测过程中的重要环节,对秋水仙素、8-羟基喹啉不同预处理时间、不同酶解时间进行探索,确定了0.1%的秋水仙素预处理8h、37℃酶解60min后细胞学检测的效果最好,检测结果确定唐菖蒲“超级玫瑰”为二倍体,其染色体数2n=2x=30；变异株为四倍体,2n=4x=60。
4.用不同浓度EMS对唐菖蒲愈伤组织诱变处理,确定了EMS的半致死剂量为9mg·L-1,EMS诱变M0代出现了8种形态上的变异,花瓣无性系的变异率为5.15%,高于花梗(1.02%),子球茎再生体系未出现变异；从诱变无性系中找到了单一性状的突变体,为突变系的建立奠定了基础。
5.对不同器官来源及半致死剂量诱变处理的各组无性系M0代共1134株在第一生长周期进行了ISSR检测,共检测出136株变异单株。对这些检测出的变异株编号种植后,在第二、第三生长周期观察到花瓣无性系组中有2株花色变异株、EMS处理的无性系组中7株出苗变异株；而这些变异株在第一生长周期的ISSR分子标记检测中均已检测到,即ISSR检测与性状观察的符合率为100%；秋水仙素处理的无性系第一生长周期R共检测出的124株变异株,在第二、第三生长周期中实际观测到97株多倍体变异株,ISSR检测与性状观察的符合率为78.2%。
创新点：
(1)突破了传统的用电子束或钴60辐射诱变唐菖蒲种球的方法,首次用唐菖蒲不同器官来源的愈伤组织为材料,进行秋水仙素与EMS离体诱导,建立了不同的体细胞无性系,扩大了变异谱,实现了诱变方法的创新。
(2)采用ISSR分子标记技术与生物学性状观察相结合的方法,从体细胞无性系中筛选出了变异株,建立了无性系变异群体,获得了2株花色变异株、7株出苗变异株及6株无子球变异株,实现了种质资源的创新。
Gladiolus hybridus Hort is a perennial corm plant which belongs to Iridaceae Gladiolus family. It is one of four famous cutting-flowers all over the world. It plays an important role in the field of flower production and consumption. At present, the bulbs are mainly introduced from abroad. Few are bred in our country. There are only a few summer-flower-type varieties in current cultivars which resulted in narrow genetic resources among the cultivars. Little has been reported relating to Gladiolus genetic basic research. To establish mutant library by using somaclonal variation, chemical mutagenesis and molecular marker-assisted breeding is important to breed some new varieties or lines with independent intellectual property rights or lay the foundation for expanding the available germplasm resources.
In this study Gladiolus cultivar'Rose Supreme'was used as material and differen organs were used as explants to establish high frequency regeneration system. Callus derived from different organs were treated by chemical mutagen to expand variation spectrum at the same time. Mutants were screened from somatic clones by morphological, cytological observation and ISSR molecular marker assisted method. It laid the foundation for genetic map construction and breeding in Gladiolus. It also provided a new way for landscape plants which are difficult to hybrid and need asexual ways to propagate, long time to obtain pure lines.
1. GRose Supreme was used to establish regeneration systems from different organs such as cormels, leaves, petals, pedicels and filaments. The results indicated that regeneration systems can be established by using cormels, petals and pedicels as explants, while using leaves and filaments didn't succeed.The best medium for induction of callus is MS+2,4-D4.0mg-L-1+6-BA0.5mg-L-1. The best medium for somatic embryos induction is MS+2,4-D1.0mg·L-1+TDZ0.3mg-L-1.TDZ is the most important factors for somatic embryo induction. This is the first report to reveal the process of induction of somatic embryogenesis and their development.
2. When petal was used as explants to establish regeneration system, pigment of explants can be passed down to new organs with cell division. And the petal callus also can develop into petals directly. Plantlets induced by the petals carried the original pigment of peals in the process of plant architecture apparently. The phenomenon of the pigment expression in plantlets and chimerism also happens when pedicels were used as explants.The pigment expression does not appear when cormels were used as explants. The frequency of the petals somaclonal variation is higher than the pedicel under natural condition.The research revealed characteristics of plant architechture of regenerated plantlets derived from different organs.
3. Petal callus was treated by different concentration of colchicines for3w. Half fatal dose of colchicines was0.5mg·L-1.56.95%plantlets derived from callus treated by colchicines appeared sturdy, leaf hypertrophy. The stomata of mutants are larger than the control's. The chromosome number of control is2n=4x=30, while the chromosome number of mutants is4n=4x=60.
4. Petal callus was treated by different concentration of EMS for3w. Half lethal dose of EMS was9.0mg·L-1. Plantlets in Mo generation appeared8types of malformation. Contents of total soluble protein, soluble sugar and SOD, POD activity varied according to concentration and duration of EMS treatment.
5.136mutants were detected from1134somatic lines derived from different organs and callus treated by fatal dose of chemical mutagenesis. The mutation rate of different somatic lines was analyzed. Two flower-color mutants derived from petals clones were observed in third growth period,7geminating mutants derived from petal callus treated by EMS were observed. These mutants have been detected in the first growth period by1SSR molecular markers assisted method. The coincidence rate of1SSR detection and traits observation is100%; ISSR detected a total of124mutants were detected by ISSR molecular marker in the first growth period from somatic lines treated by colchicines and97mutants were actually observed in the second and third growth period, The coincidence rate of ISSR detection and traits observation is78.2%.
Innovation:
(1)The traditional methods of getting mutants were radiating bulbs with electronic beam or Co60. And callus of different organs of Gladiolus were first used as inducing materials for colchicine and EMS in vitro induction, and established different somaclonal, Expanding of the variation spectrum, and innovating the mutagenesis methods.
(1)The methods of combinating ISSR molecular markers and biological observation were used in the reserch.Somatic clones were established.Two mutants in flower,7mutants in germinating and6mutants in forming cormels were obtained in terms of innovation of germplasm resources.
本研究以唐菖蒲主栽品种“超级玫瑰”(G. Rose Supreme)为试材,通过对不同器官来源的外植体进行愈伤组织诱导,建立了唐菖蒲高频再生体系,同时通过化学诱变剂处理愈伤组织,扩大变异谱；利用形态学、细胞学观察结合ISSR分子标记技术,筛选唐菖蒲体细胞无性系突变株,为唐菖蒲遗传图谱的构建及育种奠定基础,同时也为以无性繁殖、获得纯系年限长且有性杂交困难的花卉定向育种提供新途径。研究的主要结果如下：
1.以唐菖蒲主栽品种“超级玫瑰”不同器官为外植体进行了诱导,结果表明用子球、花瓣及花梗作外植体均可诱导形成再生植株,用叶片、花丝做外植体没有得到再生植株。诱导唐菖蒲愈伤组织的最佳培养基为MS+2,4-D4.0mg·L-1+6-BA0.5mg·L1,诱导体细胞胚的最佳培养基为MS+2,4-D1.0mg·L-1+TDZ0.3mg·L-1,确定了TDZ是决定唐菖蒲体细胞胚产生的重要因子；首次对唐菖蒲各器官来源的体细胞胚的诱导及发育过程进行了研究。
2.用花瓣作外植体建立再生体系过程中,再生植株出现了色素重新表达现象,即由花瓣诱导的再生植株在幼苗发育过程中明显携带花瓣中原有色素现象；由花瓣愈伤组织可直接再发育成花瓣；由花梗做外植体也出现部分色素表达现象,且再生植株易出现嵌合体；而用子球茎作外植体则没有出现色素表达现象。自然情况下,花瓣无性系变异的频率高于花梗,揭示了唐菖蒲不同器官来源再生植株器官发生及形态建成的特点。
3.用秋水仙素不同浓度、时间处理唐菖蒲愈伤组织,确定了秋水仙素的半致死剂量为0.5mg·L-1,用半致死剂量的秋水仙素处理各器官来源的愈伤组织,共获得124株变异株,其中花瓣无性系的变异率最高,为56.95%,花梗次之,为21.1%,子球茎最低,为12.6%,变异株形态上比对照明显粗壮。本研究根据细胞学检测过程中的重要环节,对秋水仙素、8-羟基喹啉不同预处理时间、不同酶解时间进行探索,确定了0.1%的秋水仙素预处理8h、37℃酶解60min后细胞学检测的效果最好,检测结果确定唐菖蒲“超级玫瑰”为二倍体,其染色体数2n=2x=30；变异株为四倍体,2n=4x=60。
4.用不同浓度EMS对唐菖蒲愈伤组织诱变处理,确定了EMS的半致死剂量为9mg·L-1,EMS诱变M0代出现了8种形态上的变异,花瓣无性系的变异率为5.15%,高于花梗(1.02%),子球茎再生体系未出现变异；从诱变无性系中找到了单一性状的突变体,为突变系的建立奠定了基础。
5.对不同器官来源及半致死剂量诱变处理的各组无性系M0代共1134株在第一生长周期进行了ISSR检测,共检测出136株变异单株。对这些检测出的变异株编号种植后,在第二、第三生长周期观察到花瓣无性系组中有2株花色变异株、EMS处理的无性系组中7株出苗变异株；而这些变异株在第一生长周期的ISSR分子标记检测中均已检测到,即ISSR检测与性状观察的符合率为100%；秋水仙素处理的无性系第一生长周期R共检测出的124株变异株,在第二、第三生长周期中实际观测到97株多倍体变异株,ISSR检测与性状观察的符合率为78.2%。
创新点：
(1)突破了传统的用电子束或钴60辐射诱变唐菖蒲种球的方法,首次用唐菖蒲不同器官来源的愈伤组织为材料,进行秋水仙素与EMS离体诱导,建立了不同的体细胞无性系,扩大了变异谱,实现了诱变方法的创新。
(2)采用ISSR分子标记技术与生物学性状观察相结合的方法,从体细胞无性系中筛选出了变异株,建立了无性系变异群体,获得了2株花色变异株、7株出苗变异株及6株无子球变异株,实现了种质资源的创新。
Gladiolus hybridus Hort is a perennial corm plant which belongs to Iridaceae Gladiolus family. It is one of four famous cutting-flowers all over the world. It plays an important role in the field of flower production and consumption. At present, the bulbs are mainly introduced from abroad. Few are bred in our country. There are only a few summer-flower-type varieties in current cultivars which resulted in narrow genetic resources among the cultivars. Little has been reported relating to Gladiolus genetic basic research. To establish mutant library by using somaclonal variation, chemical mutagenesis and molecular marker-assisted breeding is important to breed some new varieties or lines with independent intellectual property rights or lay the foundation for expanding the available germplasm resources.
In this study Gladiolus cultivar'Rose Supreme'was used as material and differen organs were used as explants to establish high frequency regeneration system. Callus derived from different organs were treated by chemical mutagen to expand variation spectrum at the same time. Mutants were screened from somatic clones by morphological, cytological observation and ISSR molecular marker assisted method. It laid the foundation for genetic map construction and breeding in Gladiolus. It also provided a new way for landscape plants which are difficult to hybrid and need asexual ways to propagate, long time to obtain pure lines.
1. GRose Supreme was used to establish regeneration systems from different organs such as cormels, leaves, petals, pedicels and filaments. The results indicated that regeneration systems can be established by using cormels, petals and pedicels as explants, while using leaves and filaments didn't succeed.The best medium for induction of callus is MS+2,4-D4.0mg-L-1+6-BA0.5mg-L-1. The best medium for somatic embryos induction is MS+2,4-D1.0mg·L-1+TDZ0.3mg-L-1.TDZ is the most important factors for somatic embryo induction. This is the first report to reveal the process of induction of somatic embryogenesis and their development.
2. When petal was used as explants to establish regeneration system, pigment of explants can be passed down to new organs with cell division. And the petal callus also can develop into petals directly. Plantlets induced by the petals carried the original pigment of peals in the process of plant architecture apparently. The phenomenon of the pigment expression in plantlets and chimerism also happens when pedicels were used as explants.The pigment expression does not appear when cormels were used as explants. The frequency of the petals somaclonal variation is higher than the pedicel under natural condition.The research revealed characteristics of plant architechture of regenerated plantlets derived from different organs.
3. Petal callus was treated by different concentration of colchicines for3w. Half fatal dose of colchicines was0.5mg·L-1.56.95%plantlets derived from callus treated by colchicines appeared sturdy, leaf hypertrophy. The stomata of mutants are larger than the control's. The chromosome number of control is2n=4x=30, while the chromosome number of mutants is4n=4x=60.
4. Petal callus was treated by different concentration of EMS for3w. Half lethal dose of EMS was9.0mg·L-1. Plantlets in Mo generation appeared8types of malformation. Contents of total soluble protein, soluble sugar and SOD, POD activity varied according to concentration and duration of EMS treatment.
5.136mutants were detected from1134somatic lines derived from different organs and callus treated by fatal dose of chemical mutagenesis. The mutation rate of different somatic lines was analyzed. Two flower-color mutants derived from petals clones were observed in third growth period,7geminating mutants derived from petal callus treated by EMS were observed. These mutants have been detected in the first growth period by1SSR molecular markers assisted method. The coincidence rate of1SSR detection and traits observation is100%; ISSR detected a total of124mutants were detected by ISSR molecular marker in the first growth period from somatic lines treated by colchicines and97mutants were actually observed in the second and third growth period, The coincidence rate of ISSR detection and traits observation is78.2%.
Innovation:
(1)The traditional methods of getting mutants were radiating bulbs with electronic beam or Co60. And callus of different organs of Gladiolus were first used as inducing materials for colchicine and EMS in vitro induction, and established different somaclonal, Expanding of the variation spectrum, and innovating the mutagenesis methods.
(1)The methods of combinating ISSR molecular markers and biological observation were used in the reserch.Somatic clones were established.Two mutants in flower,7mutants in germinating and6mutants in forming cormels were obtained in terms of innovation of germplasm resources.
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