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Application of Heavy Ion Beam Irradiation in Ornamental Flowers Breeding

重离子束辐射技术在花卉育种中的应用



全 文 :园 艺 学 报 2013,40(9):1837–1845 http: // www. ahs. ac. cn
Acta Horticulturae Sinica E-mail: yuanyixuebao@126.com
收稿日期:2013–04–27;修回日期:2013–08–28
基金项目:国家林业局林业公益性行业科研专项(201204609);‘十二五’国家科技支撑计划课题(2013BAD01B07);北京市科学技术
研究院青年骨干计划项目(2012-016)
* 通信作者 Author for correspondence(E-mail:bjr301@126.com)
重离子束辐射技术在花卉育种中的应用
孔 滢 1,2,白锦荣 1,*,尚宏忠 1,王乃彦 1,2
(1 北京市辐射中心,北京 100875;2 北京师范大学核科学与技术学院,北京 100875)
摘 要:重离子束辐射是一种新兴的辐射诱变技术。与传统的 γ 射线、X 射线等相比,重离子束具有
高传能线密度,能够产生更高的相对生物学效应,具有诱变率更高、诱变范围更大的特点。常用的离子
源主要有 C、N、Ne 等,不同植物材料的不同部位和发育阶段对重离子束的敏感度效果不一。通过结合
植物组织培养、辐射材料处理、多次辐射和基因工程等技术,目前已对菊花、香石竹、美女樱和夏堇等
20 多种植物开展了重离子束辐射育种研究,获得了新种质和新品种,部分新品种在市场推广后获得了良
好的经济效益。因此,重离子束辐射技术在观赏植物育种上有着广泛的应用前景。
关键词:重离子束;辐射;观赏植物;育种
中图分类号:S 68 文献标志码:A 文章编号:0513-353X(2013)09-1837-09

Application of Heavy Ion Beam Irradiation in Ornamental Flowers
Breeding
KONG Ying1,2,BAI Jin-rong1,*,SHANG Hong-zhong1,and WANG Nai-yan1,2
(1Beijing Radiation Center,Beijing 100875,China;2College of Nuclear Science and Technology,Beijing Normal
University,Beijing 100875,China)
Abstract:Heavy ion beam irradiation is an emerging radiation mutagenesis technology. Compared
with traditional radiation such as X-rays and gamma rays,heavy ion beam provides a high linear energy
transfer(LET)and relative biological effectiveness(RBE). This technique induces mutations at a higher
rate and wider mutation spectrum. When irradiated,the sensitivity of different parts and developmental
stages of different plant materials varies. Radiation ion source commonly used is C,N,Ne etc. Through a
combination of tissue culture,materials processing,re-irradiation and genetic engineering,this approach
has been applied in more than 20 ornamental plants,including chrysanthemum,carnation,verbena,torenia
etc. Many novel mutations and varieties have been obtained,some of them have been commercialized with
large economic benefits. Heavy ion beam technology is an excellent tool to improve mutation breeding of
horticulture plants with high efficiency.
Key words:heavy ion beam;irradiation;ornamental plant;breeding

近年来,重离子束辐射(heavy-ion beam irradiation)作为一种新兴的辐射诱变源越来越受到关
注。重离子是指荷能质量数大于 4 的带电粒子(周利斌 等,2008),具有高的传能线密度(带电粒

1838 园 艺 学 报 40 卷
子在单位长度径迹上传递的能量,linear energy transfer,LET)和相对生物学效应(relative biological
effectiveness,RBE),其单位剂量的诱变效应高于 X 射线、γ 射线、电子束等(Okamura et al.,2003;
Shikazono et al.,2003;周利斌 等,2008)。重离子通过加速器注入生物体内,同生物材料中的分子、
原子发生一系列的碰撞(王菊芳 等,2007),能在局部区域释放出高能量,显著诱导单链或双链 DNA
断裂、末端受损,从而不容易修复(Goodhead,1995)。
重离子束辐射是一种很有效的诱导技术手段,可以有效的改变目标性状,能在不影响其他特征
的基础上提供一个广泛的变异范围(Shikazono et al.,2005),通过 γ 射线辐射未能获得的特殊花色
可以通过离子辐射育种诱导出(Okamura et al.,2003),从而提高园艺和农作物的诱变频率。在对拟
南芥(Arabidopsis thaliana)的碳离子束辐照研究中发现,一半的变异为点变异,另一半有着大的
DNA 改变,例如插入、易位和大片段缺失(Shikazono et al.,2005)。一般认为重离子束能产生比较
大的 DNA 变换,插入、易位和大片段缺失,而不是点变异(Tanaka & Hase,2009)。目前开展重离
子束辐射育种的单位及其设备主要有日本理化学研究所的放射性同位素束流设施(Riken RI Beam
Factory);日本原子能机构的高崎量子应用研究所(Takasaki Ion Accelerators for Advanced Radiation
Application);日本若狭湾能源研究中心的多用途同步串联加速器(Wakasa Wan Energy Research
Center Multi-purpose Accelerator with Synchrotron and Tandem);中国科学院近代物理研究所(Institute
of Modern physics)和意大利国家核物理研究所的南方国家实验室(Laboratori Nazionali del Sud)等。
1 辐射材料与方法
不同的植物材料对辐射的敏感性有很大差异,表现在不同物种、品种以及同一植株的不同发育
阶段和器官上(Kazama et al.,2008;孙兰弟,2008)。目前用于重离子束辐射的材料有干种子、吸
水饱和种子、芽、叶片、花瓣、愈伤组织、组培小苗、体细胞胚等,它们对重离子束的敏感度为:
愈伤组织 > 发育中胚 > 茎节 > 吸水饱和种子 > 干种子(Abe et al.,2007,2012;Kazama et al.,
2008)。
一般说来,组培苗具有较强的分化和形成新组织、器官的能力,是辐射育种较理想的材料(彭
绿春 等,2007);且植物组织培养用的外植体材料含水量较干种子要高得多,辐射敏感性相应也高
得多(周利斌 等,2007)。但也有一些例外,例如在仙客来(Cyclamen persicum)中,比起愈伤组
织、体细胞胚等,用重离子束辐射块茎更能有效的改变花色性状(Sugiyama et al.,2008)。在菊花
(Chrysanthemum morifolium)中,和叶片相比,辐射花瓣后结合组织培养能获得诱变率更高的花色
突变体,推测花色相关基因在花瓣中表达更为活跃,从而更容易发生变异(Hase et al.,2009)。
进行辐射所用的离子源主要有 C、N、O、Ne、Ar、Fe 等。目前的研究结果显示,离子束辐射
诱变的影响并非取决于离子源的种类,而是取决于它的传能线密度 LET。目前应用较多的离子源是
碳离子,常用 LET 值的范围为 100 ~ 200 keV · μm-1(Atsushi & Yoshihiro,2010)。中国科学院近代
物理研究所用低能氮、碳离子束辐照万寿菊(Tagetes erecta)、一串红(Salvia splendens)、矮牵牛
(Petunia hybrida)、长春花(Catharanthus roseus)、鸡冠花(Celosia cristata)的种子和大丽花(Dahlia
pinnata)的花芽,获得了观赏性状改变的诱变植株(卫增泉 等,2003;周利斌 等,2008)。
2 对观赏植物的诱变
对观赏植物而言花色花型是重要的观赏性状,通过进行重离子束辐射,诱导出了很多新的表型。
例如获得了复杂花色和条纹花色的菊花以及腋芽减少的菊花突变体,并对该诱变植株进行了再次辐
9 期 孔 滢等:重离子束辐射技术在花卉育种中的应用 1839

射,获得了腋芽减少且在低温下能够开花的诱变植株(Atsushi & Yoshihiro,2010)。重离子束辐射
后的香石竹(Dianthus caryophyllus)植株表现出了比 X 射线辐照更多的花色变异种类(Okamura et
al.,2003)。通过重离子束诱导还获得了花朵粉色、樱桃红色,有条纹和斑点的矮牵牛植株(Okamura
et al.,2009)。在月季(Rosa)中,获得了花色更加艳丽、花瓣数量、花的大小和花形都发生改变的
诱变植株(Yamaguchi et al.,2003)。在夏堇(Torenia fournieri)中,通过离子束辐射获得了两类花
色变异体,一种是缺乏花色合成的基因,另一种是花色基因的表达发生了改变(Miyazaki et al.,2006)。
同时,对许多观赏植物而言,不育性状是很重要的。一方面能够增加花朵数量和延长花期,另
一方面雄性不育的植株不会通过授粉将其基因传递到自然环境中(Sugiyama et al.,2008)。美女樱
(Verbena hybrida)经过重离子束辐射后成功的分离出不育变异体‘Temari Bright Pink’,是世界上
第 1 个通过重离子束辐射技术获得的新品种,在 2002 年春季进入商业市场(Kazama et al.,2008)。
此外,由于辐射诱变的不确定性,还获得了一些株形、植株高度、叶色等性状发生变异的突变体。
目前开展重离子辐射的植物材料及结果见表 1。

表 1 重离子束辐射观赏植物获得的变异后代
Table 1 Mutant lines developed in various ornamentals using heavy ion beam breeding
序号
No.
植物材料
Plants species
辐射部位
Irradiated
materials
离子束种类和参数
Ion-beam and
physical parameters
获得突变体
Mutation induction
参考文献
References
1 矮牵牛 Petunia hybrida‘Kirin
Hana-Saka Rose’
离体芽
In vitro buds
12C6+(320 MeV) 花色粉和樱桃红、花瓣中细白条纹和
斑点的变异植株 Flower color
mutants with pink,cherry,minute
striped white,and spotted petals
Okamura et al.,
2009
2 矮牵牛 Pentunia hybrida
‘BBss11’
播种小苗
Seedlings
12C6+(320 MeV) 花色洋红、紫红、淡粉、白、蓝色白
边和深紫红等变异植株 Mutants of
magenta,purple,light pink,white,
blue picotee and burgundy flowers
Hase et al.,2009,
2010
3 巴西水竹叶 Tradescantia
fluminensis
嫩枝生长点
Growing points
of shoots
12C(95.8 MeV · u-1)彩叶、花葶紫、花萼白、花朵淡粉的
变异植株 Mutant with color leaves,
purple stalk,cholorophyll-deficient
calyces and light pink flower
He et al.,2011
4 大丽花 Dahlia‘Miharu’ 嫩枝
Shoots
14N(135 MeV · u-1) 花色深粉、浅粉、粉白等和花径变异
的诱变植株 Flower color mutants
(dark pink,pale pink and pink-white
etc.)and flower diameter mutants
Abe et al.,2005
5 大丽花 Dahlia spp. 组培苗生长点
Tissue-cultured
shoots
12C6+(320 MeV) 花色、花形变异的诱变植株 Mutants
on flower color and flower shape
Uyama et al.,2012
6 飞燕草 Delphinium‘Monoka’ 叶片
Leaf
12C6+(320 MeV) 花色淡粉、红粉、黄粉,花序小花增
多和矮生的变异植株 Flower color
mutants(light pink,red pink and yellow
pink),mutant with increased number of
spikes and dwarf mutant
Chinone et al.,
2009;Kitamura et
al.,2010
7 鸽石斛 Dendrobium crumenatum 原球茎
Protocorm-like
bodies
12C6+(320 MeV) 花朵变大、花葶变长的变异植株
Mutants with large flowers,a longer
flower stalk
Affrida et al.,2008
8 宫灯百合 Sandersonia aurantiaca 干种子
Dry seeds
14N(28.5 keV · μm-1)白化变异植株 Albino mutant plants Horita et al.,2002;
Mohan Jain,2006;
Abe et al.,2012
9 菊花 Chrysanthemum morifolium
‘Reagan Red’
舌状花
Ray floret
12C6+(320 MeV) 株高、花形、花色、花朵大小和数量,
叶色和叶形等突变 Mutants on plant
height,flower shape,flower color,
number of flower bud and flower size,
leaf color and shape
Affrida et al.,
2010;Shakinah et
al.,2011
1840 园 艺 学 报 40 卷
续表 1
序号
No.
植物材料
Plants species
辐射部位
Irradiated
materials
离子束种类和参数
Ion-beam and physical
parameters
获得突变体
Mutation induction
参考文献
References
10 菊花
Chrysanthemum‘Jimba’
叶片
Leaves
12C5+(220 MeV);
12C6+(320 MeV)
低温下开花和腋芽减少的变异植株
Mutants with few axillary buds,low
temperature flowering trait
Ueno et al.,2003
11 菊花 Chrysanthemum‘Jimba’腋
芽减少的诱变植株‘Aladdin’
few axillary buds mutant
‘Aladdin’induced from
Chrysanthemum‘Jimba’
叶片
Leaves
12C5+(220 MeV);
12C6+(320 MeV)
腋芽减少且在低温下能够开花的变
异植株 Mutants with few axillary
flower buds and low temperature
flowering
Ueno et al.,2004
12 菊花
Chrysanthemum‘Hakusui’
离体试管芽
In vitro buds
12C6+(320 MeV) 不定芽减少和低温下开花的变异植
株 Mutants with less adventitious
buds,no flowering delay under low
temperature condition
Asami et al.,
2010;Hase et al.,
2011
13 农杆菌介导的转 pac1 基因菊花
Dendranthema grandiflora
agrobacterium-mediated trangenic
chrysanthemum
叶片
Leaf segments
12C6+(320 MeV) 舌状花浅粉、深粉、浅橙、白和黄色
等变异植株 Pale pink,dark pink,
salmon,white and yellow etc. ray
florets mutants
Okamura et al.,
2008
14 菊花
Chrysanthemum morifolium
‘H13’
叶片
Leaf segments
12C5+(220 MeV) 舌状花深红、朱红、粉红、红白、红
白重瓣,和 T 型花簇的变异植株
Floret color mutants ( dark-red ,
vermilion,pink,white-red,white-red
double blooms),and mutant with T
type of flower cluster
Furutani et al.,
2008;Matsumura
et al.,2010
15 菊花 Chrysanthemum morifolium
‘Shiroyamate’
舌状花
Ray florets
12C5+(220 MeV) 舌 状 花 黄 和 淡 黄 的 变 异 植 株
Yellow and pale yellow ray florets
mutants
Matsumura et al.,
2010
16 菊花
Chrysanthemum morifolium
‘Taihei’
茎节的腋芽
Nodes with
axillary buds
12C(220 MeV) 花朵浅粉、深粉、白、黄、浅黄、橙、
深橙、红等的变异植株 Mutants
with pale pink,deep pink,white,
yellow,pale yellow,orange,deep
orange and red flowers
Yamaguchi et al.,
2009
17 蓝眼菊 Osteospermum‘Mother
Symphony’
叶片
Leaf segments
12C5+(220 MeV);
12C6+(320 MeV)
花色白、奶油、深黄和橙,斑叶和矮
生的变异植株 Flower color mutants
(white, cream, deep yellow and
orange),dwarf mutants and variegated
leaf mutants
Iizuka et al.,2006
18 蓝眼菊 Osteospermum‘Mother
Symphony’经离子辐射后的诱变
株 OM7 OM7 induced from
radiated mutants Osteospermum
‘Mother Symphony’
叶片
Leaf section
12C5+(220 MeV);
12C6+(320 MeV)
花色白紫、淡橙、和花瓣背面黄色的
变异植株 White purple and light
bitter orange flower mutants , and
mutant with light yellow in the back
side of the petal
Iizuka et al.,2008
19 蓝眼菊诱变株 Osteospermum
‘Mother Symphony’OM7 再次离
子辐射后诱变株 OM706
OM706 induced from re-irradiated
mutants Osteospermum ‘Mother
Symphony’OM7
叶片
Leaf sections
12C5+(220 MeV);
12C6+(320 MeV)
花朵黄白、浅橙、浅黄、淡黄色和花
背面亮黄色的变异植株 Flower
color mutants with yellowish white,
whitish light orange , whitish light
yellow,and lighter yellow in the back
side of the petal
Okada et al.,2009
20 毛油点草 Tricyrtis hirta 花被片诱导的
胚性愈伤组织
Embryogenic
calluses
induced from
tepals
12C6+(135 MeV · u-1)矮生、叶形、花朵大小和数量变异的
植株 Mutants on dwarf,leaf shape,
flower number and flower size
Nakano et al.,
2010a
21 美女樱 Verbena × hybrida
‘Temari Sakura’,‘Temari Coral
Pink’
含有侧生分生
组织的茎节
Nodes
containing
lateral
meristems
14N7+(1.89 GeV) 花数量增多、花期变长、不育的变异
植株 Mutants with a large number of
inflorescences,better longevity,and
sterile trait
Kanaya et al.,
2008

9 期 孔 滢等:重离子束辐射技术在花卉育种中的应用 1841

续表 1
序号
No.
植物材料
Plants species
辐射部位
Irradiated
materials
离子束种类和参数
Ion-beam and physical
parameters
获得突变体
Mutation induction
参考文献
References
22 天竺葵 Pelargonium‘Splendide’ 茎节上的腋

Stem node
containing a
single axillary
bud
12C(23 keV · μm-1) 雄蕊萎缩、无花粉的雄性不育诱
变 植 株 Male sterile mutants
lacked pollen-producing ability
Sugiyama et al.,
2005;Mohan
Jain,2006;Abe
et al.,2012
23 夏堇 Torenia‘Summer Wave
Blue’
叶组织和不
含侧生分生
组织的茎节
Leaf tissue
and stem
internodes
without lateral
meristems
14N(135 MeV · u-1);
20Ne(135 MeV · u-1)
花朵淡蓝、蓝、淡粉和粉色的变
异植株 Pale blue,blue,pale pink
and pink flower mutants
Miyazaki et al.,
2006
24 夏堇和转基因(CHS 和 DFR)株

Torenia fournieri‘Crown Violet’
and transgenic lines
组培苗叶片
Leaf disks
from in vitro
plants
12C(1.62 GeV);
20Ne(2.70 GeV)
花色更柔和、镶边、条纹等和花
瓣圆形、波状、流苏状等变异植
株 Flower color and coloration
pattern mutants ( tone-shifted ,
bordered , streaked etc. ), petal
shape and corolla divergence
mutants(rounded,wavy,fringed
etc.)
Sasaki et al.,
2008
25 仙客来 Cyclamen persicum
‘Fragrance Mini’
体胚小苗的
块茎 Tubers
of plantlets
induced from
somatic
embryos
12C6+(1.62 GeV) 花色淡蓝紫色加白色条纹、淡红
紫色等和花瓣形状变异的变异嵌
合体及雄性不育植株 Chimeric
mutants on flower color(reddish
purple,striped pattern of white-
purple)and petal forming,and male
sterile mutants
Sugiyama et al.,
2008
26 仙客来 Cyclamen persicum × C.
purpurascens‘Kaori-no-mai’
种子组培暗
培养后的黄
化叶柄
Etiolated
petioles
induced from
germinated
seedlings
(kept in
dark)
12C6+(320 MeV) 花红紫色的变异植株
Red-purple flowers mutant
Kondo et al.,
2009,2010
27 香石竹 Dianthus caryophyllus
‘Vital’
叶片
Leaf segments
12C5+(220 MeV) 花红、深粉、粉、浅粉、黄、双
色、条纹等和花型石竹型、月季
型等的变异植株 Flower color
mutants(red,dark pink,pink,light
pink,yellow,bi-colored,striped)
and flower shape mutants(Dianthus
type and rose type)
Okamura et al.,
2003
28 一串红 Salvia splendens 干种子
Dry seeds
12C6+(80.55 MeV · u-1) 深红色和鲜红色花朵变异植株
Dark red and fresh red flower color
mutants
Wu et al.,2009
29 月季 Rosa‘Orange Rosamini’ 组培苗的茎

Stems in vitro
culture
12C(220 MeV) 花色更加鲜艳、浅红色、花瓣数
量、花朵大小、和花型发生变异
的突变体 Mutants on flower
color ( intense flower color and
reddish flower),petals number,
flower size and flower shape
Yamaguchi et
al.,2003
30 非洲紫罗兰 Saintpaulia ionahta
‘Mauve’,‘Indikon’
幼嫩叶片
Young leaves
12C6+(960 MeV) 获得了叶绿素缺失突变体
Chlorophyll deficiency mutants
Zhou et al.,2006

1842 园 艺 学 报 40 卷
3 与其他技术的结合
为了提高目的性状的诱变频率和稳定性,常需要将重离子束辐射技术与其他技术相结合。
一是组织培养技术。辐射诱变育种过程中,突变嵌合体的分离和稳定变异体的获得是关键内容
之一(彭绿春 等,2007)。重离子束辐射结合组织培养能够更有效的对突变体进行诱导和筛选,将
其中的有益突变固定和增殖,有效的加速了新品种培育(Okamura et al.,2003;Zhou et al.,2006;
Sugiyama et al.,2008;Matsumura et al.,2010;Nakano et al.,2010b)。同时由于花卉主要是靠无性
繁殖方式进行扩繁,重离子辐照结合组织培养为花卉产业提供了新的育种途径(周利斌 等,2008),
在培育园艺新品种上有着广阔的应用前景。
二是辐射材料处理技术。对辐射材料采取恰当的预处理能够有效提高目标性状的诱变频率。例
如发现用高浓度蔗糖溶液对矮牵牛和萝卜小苗进行预处理,可以显著提高色素的积累,提高花色变
异植株的诱变率(Hara et al.,2003;Hase et al.,2010)。
三是重复重离子束辐射技术。对突变体进行离子再辐射以获得丰富的变异性状(Yamaguchi et
al.,2003;Iizuka et al.,2008)。例如为了获得纯白色舌状花的蓝眼菊,对诱变植株进行多次重离子
束辐射(Okada et al.,2009)。
四是基因工程技术。通过重离子束辐射技术获得新的诱变植株后,可以通过基因工程技术获得
新基因。例如通过重离子束辐照拟南芥后获得了多种突变体,从抗紫外线 B 突变体 uvi1 和 uvi4 分
离鉴定了相关基因(Tanaka et al.,2002;Hase et al.,2006);从花青素积累 ast 突变体或花青素缺乏
tt19 突变体中获得了部分相关基因(Tanaka et al.,1997;Kitamura et al.,2004);从锯齿状花瓣和萼
片的突变体 frill1 获得了与该表型相关的基因等(Hase et al.,2000,2005)。
4 讨论与展望
同常规辐射育种相比,重离子束辐射育种有着诱变效率更高,诱变谱更广的特点。结合植物无
性繁殖技术,对辐射后的材料进行栽培、选育和突变体鉴定等,获得具有优良性状的新品种,缩短
了育种年限。然而重离子束辐射育种中也存在着一些缺点。例如诱变率不高,仍需要在大量诱变植
株中进行选育;目的性较差,辐射突变方向不定(程金水,2000);以及重复性不高,在不同的试验
中获得相同的变异植株较为困难(Kondo et al.,2008;Sasaki et al.,2008)。此外,辐射剂量的多少
除了对植物的诱变效果有所影响外,也对植物生长发育有所影响,例如碳离子束辐射石斛
(Dendrobium mirbellianum)后生长十分缓慢(Affrida et al.,2008);以及一串红的花色突变体在所
有的生长阶段都受到抑制,但在开花和结实上没有任何影响(Wu et al.,2009)。经重离子束辐射后,
毛油点草的胚性愈伤组织的生长也受到了抑制,但在低剂量辐射时明显促进了愈伤组织中体细胞胚
的生长(Nakano et al.,2010b)。因此一般选择相对较低的辐射剂量,既有较高的诱变率,又不会对
植物生长有明显抑制。总体而言,重离子束辐射在花卉新品种培育方面有着很好的研究、开发与应
用前景。

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