γ射线对菊花（Dendranthema morifolium（Ramat.）Tzvel.）的辐射诱变效应研究
摘要
菊花别名鞠、寿客、帝女花、黄华等,是我国的传统名花,栽培历史已有1600多年,与兰花、梅花、竹并列合称为“四君子”。菊花色彩艳丽,姿态万千,不畏严寒,傲雪怒放,自古以来深受人们的喜爱,而且菊花的用途非常广泛,地被菊及岩菊可布置花坛、花境及岩石园:杭菊可入药,或作清凉饮料用。另外,菊花是世界上四大切花之一,近年来菊花的销售量在切花总量中一直位居榜首,约占总量的30%。菊花作为无性繁殖的观赏植物,一些品种花粉极少或没有花粉,且遗传上高度杂合且为多倍体,比较适合辐射诱变育种。本试验研究了不同剂量~(60)Co-γ射线辐照菊花‘神马'(‘Jinba')的组培苗和地被菊Zh99-1的种子,从形态指标、叶片相对含水量、叶绿素含量、种子发芽率和抗氧化酶系统等方面研究辐射对菊花M1代的生物效应。以期为菊花辐射有诱变育种提供理论支持。主要结果如下:
1.~(60)Co-γ射线辐照‘神马'组培苗的半致死剂量在20Gy左右,继代培养可提高辐射后代的成活率,增加变异群体的数目。射线辐射对M1代大多数性状产生了抑制作用。射线辐射对‘神马'M1代的叶片相对含水量无显著影响,M1代叶片相对含水量显著低于对照,说明辐射处理降低了M1代菊花的叶片保水能力。辐射对‘神马'M1代的叶绿素含量无显著影响,高温处理使辐射M1代的叶绿素含量显著上升。地被菊种子在受到辐射处理后,发芽势随辐射剂量的增加而逐渐降低,低剂量(10Gy和20Gy)对种子发芽率无明显影响,高剂量(30Gy、40Gy和50Gy)显著抑制了种子的发芽率,剂量越高抑制作用越明显。
2.~(60)Co-γ射线辐照后‘神马'组培苗M1代叶片H_2O_2含量显著低于对照,可能与过氧化氢酶(CAT)活性和谷胱甘肽还原酶(GR)活性的显著升高有关。没有经过辐射处理的菊花叶片,在受到高温处理后也出现了H_2O_2含量的显著下降,可能与CAT活性和过氧化物酶(POD)活性的显著升高有关。高温处理后辐射M1代H_2O_2含量的显著下降同时伴随着GR活性的显著升高。低剂量(10Gy和20Gy)辐射处理使‘神马'组培苗M1代超氧化物歧化酶(SOD)活性显著提高,较高剂量(30Gy)则无明显影响。表明菊花在受到不同胁迫时,发挥主要作用的抗氧化酶可能不同。
3.~(60)Co-γ射线辐照后M1代地被菊幼苗,随剂量增加H_2O_2含量先增加后减少,10Gy辐射处理除GR活性显著降低外,对其他抗氧化酶活性均无显著影响。从20Gy辐射处理开始随辐射剂量增加CAT活性也出现了先增加后减少的趋势,而SOD随辐射剂量增加活性逐渐提高。20Gy及其以上各剂量处理使地被菊辐射处理后代成活率急剧下降,POD活性却急剧升高,说明POD活性可以作为地被菊种子辐射后代成活率的代谢指标。
4.~(60)Co-γ射线辐照处理对‘神马'组培苗M1代丙二醛(MDA)含量无显著影响,高温处理使辐射M1代MDA含量显著高于对照,辐射处理可能降低了M1代菊花的耐热性。10Gy辐射处理对M1代地被菊幼苗MDA含量无显著影响,20Gy辐射处理显著提高了MDA含量并且随着剂量的升高MDA含量逐渐增加。
Chrysanthemum is Chinese traditional flower and it's cultivation history has been 1,600 years,with the orchid,plum blossom,bamboo tied together referred to as the“four gentlemen.”Colorful chrysanthemums was very popular all the time,and it has extensive use,ground-cover chrysanthemum and rock chrysanthemum can be arranged flower beds,flower and rock garden habitats;Florists Daisy can be medicine or soft drinks.In addition,the chrysanthemum is one of the world's four major cut flowers in recent years,cut chrysanthemum has been the largest sales share in the cut-flower trade, accounting for about 30%of the total.As vegetative propagation of ornamental plants, lots of chrysanthemum varieties have a little or no pollen,and a high degree of genetic heterozygosity and polyploid makes chrysanthemum suite for radiation-induced breeding.
In this experiment we researched different doses of ~(60)Co_(-γ),ray irradiated tissue culture seedlings of cut Chrysanthemum'Jinba'and seeds of Zh99-1,research the biological effects of Chrysanthemum M1 generation with the form of indicators,leaf relative water content,chlorophyll content,seed germination rate and antioxidant enzyme systems.It can provide theoretical support for mutation breeding of chrysanthemum.The main results are listed as follows:
1.The semi-lethal dose ofγray irradiation tissue culture seedling of 'Jinba' is about 20Gy.Subculture can increase the survival rate of offspring after radiation,increasing the number of variant groups.Radiation inhibits the majority of traits of M1 generation. Ray radiation have no significant effect on the leaf relative water content of 'Jinba' M1 generation,but after high temperature treatment the relative leaf water content of M1 with radiation was significantly lower than control.The results showed radiation reduced the capacity of leaves water-retention on M1 generation of chrysanthemum. Radiation have no significant effect on chlorophyll content of 'Jinba' M1 generation, high temperature treatment made a significant increase in chlorophyll content on M1 generation.Chrysanthemum seeds were treated by radiation,with the germination potential decreased in radiation dose increased.Low-dose(10 Gy and 20 Gy)on the seed germination rate have no significant impact,high-dose(30 Gy,40 Gy and 50 Gy) significantly inhibited the seed germination rate,the higher the dose the more obvious inhibition.
2.The H_2O_2 content of leaves was significantly lower than control after 'Jinba' tissue seedlings of M1 generation were irradiated byγ-ray,the results may be related to the activity of catalase(CAT)and glutathione reductase(GR)significantly increased. The chrysanthemum leaves have not been dealt by radiation with high-temperature treatment also appeared the H_2O_2 content in a significant drop.The results may be related to the activity of CAT and peroxidase(POD) increased significantly.After high temperature treatment of radiation-generation of M1,the H_2O_2 content decreased significantly at the same time along with the GR activity increased significantly. Low-dose(10Gy and 20Gy) radiation made tissue of 'Jinba' M1 generation's superoxide dismutase(SOD) activity was significantly increased;a higher dose(30Gy) had no significant effect.This results show that chrysanthemum by the different stress,play a major role in the antioxidant enzymes may be different.
3.The M1 generation plants that irradiated byγray,with the dose increased,the H_2O_2 content first increased and then reduced.10Gy radiations in addition to a significant reduction in GR activity,the activity of other antioxidant enzymes were not significantly affected.From the beginning of irradiation with 20Gy radiation dose increased CAT activity also increased after the first downward trend,while the SOD activity increased with the radiation dose a gradual increase.Dose of 20Gy and above all sharp decline the survival rate of radiation future generations,but there was a dramatic increase in POD activity that can be used as a metabolic marker of radiation future generations'survival rate on ground-cover chrysanthemum.
4.γray irradiation treatment on the tissue of 'Jinba' M1 generation have no significant impact to content of malondialdehyde(MDA).High temperature treatment made MDA content of radiation M1 significant higher than the control,radiation may reduce the heat resistance of chrysanthemum's M 1 generation.10Gy radiation treatment on the M 1 by the chrysanthemum seedlings have no significant effect to MDA content, 20Gy radiation treatment significantly increased MDA content and increased with the dose gradually increased MDA content.
1.~(60)Co-γ射线辐照‘神马'组培苗的半致死剂量在20Gy左右,继代培养可提高辐射后代的成活率,增加变异群体的数目。射线辐射对M1代大多数性状产生了抑制作用。射线辐射对‘神马'M1代的叶片相对含水量无显著影响,M1代叶片相对含水量显著低于对照,说明辐射处理降低了M1代菊花的叶片保水能力。辐射对‘神马'M1代的叶绿素含量无显著影响,高温处理使辐射M1代的叶绿素含量显著上升。地被菊种子在受到辐射处理后,发芽势随辐射剂量的增加而逐渐降低,低剂量(10Gy和20Gy)对种子发芽率无明显影响,高剂量(30Gy、40Gy和50Gy)显著抑制了种子的发芽率,剂量越高抑制作用越明显。
2.~(60)Co-γ射线辐照后‘神马'组培苗M1代叶片H_2O_2含量显著低于对照,可能与过氧化氢酶(CAT)活性和谷胱甘肽还原酶(GR)活性的显著升高有关。没有经过辐射处理的菊花叶片,在受到高温处理后也出现了H_2O_2含量的显著下降,可能与CAT活性和过氧化物酶(POD)活性的显著升高有关。高温处理后辐射M1代H_2O_2含量的显著下降同时伴随着GR活性的显著升高。低剂量(10Gy和20Gy)辐射处理使‘神马'组培苗M1代超氧化物歧化酶(SOD)活性显著提高,较高剂量(30Gy)则无明显影响。表明菊花在受到不同胁迫时,发挥主要作用的抗氧化酶可能不同。
3.~(60)Co-γ射线辐照后M1代地被菊幼苗,随剂量增加H_2O_2含量先增加后减少,10Gy辐射处理除GR活性显著降低外,对其他抗氧化酶活性均无显著影响。从20Gy辐射处理开始随辐射剂量增加CAT活性也出现了先增加后减少的趋势,而SOD随辐射剂量增加活性逐渐提高。20Gy及其以上各剂量处理使地被菊辐射处理后代成活率急剧下降,POD活性却急剧升高,说明POD活性可以作为地被菊种子辐射后代成活率的代谢指标。
4.~(60)Co-γ射线辐照处理对‘神马'组培苗M1代丙二醛(MDA)含量无显著影响,高温处理使辐射M1代MDA含量显著高于对照,辐射处理可能降低了M1代菊花的耐热性。10Gy辐射处理对M1代地被菊幼苗MDA含量无显著影响,20Gy辐射处理显著提高了MDA含量并且随着剂量的升高MDA含量逐渐增加。
Chrysanthemum is Chinese traditional flower and it's cultivation history has been 1,600 years,with the orchid,plum blossom,bamboo tied together referred to as the“four gentlemen.”Colorful chrysanthemums was very popular all the time,and it has extensive use,ground-cover chrysanthemum and rock chrysanthemum can be arranged flower beds,flower and rock garden habitats;Florists Daisy can be medicine or soft drinks.In addition,the chrysanthemum is one of the world's four major cut flowers in recent years,cut chrysanthemum has been the largest sales share in the cut-flower trade, accounting for about 30%of the total.As vegetative propagation of ornamental plants, lots of chrysanthemum varieties have a little or no pollen,and a high degree of genetic heterozygosity and polyploid makes chrysanthemum suite for radiation-induced breeding.
In this experiment we researched different doses of ~(60)Co_(-γ),ray irradiated tissue culture seedlings of cut Chrysanthemum'Jinba'and seeds of Zh99-1,research the biological effects of Chrysanthemum M1 generation with the form of indicators,leaf relative water content,chlorophyll content,seed germination rate and antioxidant enzyme systems.It can provide theoretical support for mutation breeding of chrysanthemum.The main results are listed as follows:
1.The semi-lethal dose ofγray irradiation tissue culture seedling of 'Jinba' is about 20Gy.Subculture can increase the survival rate of offspring after radiation,increasing the number of variant groups.Radiation inhibits the majority of traits of M1 generation. Ray radiation have no significant effect on the leaf relative water content of 'Jinba' M1 generation,but after high temperature treatment the relative leaf water content of M1 with radiation was significantly lower than control.The results showed radiation reduced the capacity of leaves water-retention on M1 generation of chrysanthemum. Radiation have no significant effect on chlorophyll content of 'Jinba' M1 generation, high temperature treatment made a significant increase in chlorophyll content on M1 generation.Chrysanthemum seeds were treated by radiation,with the germination potential decreased in radiation dose increased.Low-dose(10 Gy and 20 Gy)on the seed germination rate have no significant impact,high-dose(30 Gy,40 Gy and 50 Gy) significantly inhibited the seed germination rate,the higher the dose the more obvious inhibition.
2.The H_2O_2 content of leaves was significantly lower than control after 'Jinba' tissue seedlings of M1 generation were irradiated byγ-ray,the results may be related to the activity of catalase(CAT)and glutathione reductase(GR)significantly increased. The chrysanthemum leaves have not been dealt by radiation with high-temperature treatment also appeared the H_2O_2 content in a significant drop.The results may be related to the activity of CAT and peroxidase(POD) increased significantly.After high temperature treatment of radiation-generation of M1,the H_2O_2 content decreased significantly at the same time along with the GR activity increased significantly. Low-dose(10Gy and 20Gy) radiation made tissue of 'Jinba' M1 generation's superoxide dismutase(SOD) activity was significantly increased;a higher dose(30Gy) had no significant effect.This results show that chrysanthemum by the different stress,play a major role in the antioxidant enzymes may be different.
3.The M1 generation plants that irradiated byγray,with the dose increased,the H_2O_2 content first increased and then reduced.10Gy radiations in addition to a significant reduction in GR activity,the activity of other antioxidant enzymes were not significantly affected.From the beginning of irradiation with 20Gy radiation dose increased CAT activity also increased after the first downward trend,while the SOD activity increased with the radiation dose a gradual increase.Dose of 20Gy and above all sharp decline the survival rate of radiation future generations,but there was a dramatic increase in POD activity that can be used as a metabolic marker of radiation future generations'survival rate on ground-cover chrysanthemum.
4.γray irradiation treatment on the tissue of 'Jinba' M1 generation have no significant impact to content of malondialdehyde(MDA).High temperature treatment made MDA content of radiation M1 significant higher than the control,radiation may reduce the heat resistance of chrysanthemum's M 1 generation.10Gy radiation treatment on the M 1 by the chrysanthemum seedlings have no significant effect to MDA content, 20Gy radiation treatment significantly increased MDA content and increased with the dose gradually increased MDA content.
引文
[1]蒋甲福,陈发棣,管志勇,房伟民.小菊种子辐射生物学效应的研究[J].核农学报,2004,18(6):431-434
[2]高键,卢惠萍.花卉辐射诱变育种研究进展[J].安徽农业大学学报(自然科学版),2000,(3):22-24
[3]许耀奎.作物诱变育种[M].上海:上海科技出版社,1985,25
[4]Broertjes C,Roeet S,Bokelmann G S.Mutation breeding of Chrysanthemum morifolium Ram using in vivo and in vitro adventitious bud techniques[J].Euphytica,1976,25:11-19
[5]Broertjes C,Van Harten A M.Application of mutation breeding methods.In:A J Abbott and R R Attkin(Editors):Improving vegetatively propagated crops London:Academic Press.1987,335-338
[6]王海燕,何小弟,黄永高.菊花组织培养技术在育种上的应用研究[J].江苏林业科技,2003,30(4):25-26
[7]齐孟文,王化国.我国花卉辐射育种的进展与剖析[J].核农学通报,1997,18(6),288-290
[8]刘金勇.菊花品种遗传改良的研究进展[J].湖南林业科技,2004,31(1),49-52
[9]郭安熙,范家霖,杨保安,王柏楠,张建伟,陈树国,王世荣,王然.菊花花色辐射诱变研究[J].核农学报,1997,11(2):65-73
[10]傅玉兰,郑路.冬菊新品种选育[J].安徽农业大学学报,1994,(1)59-62
[11]黄建昌,肖艳.~(60)Co-γ射线处理对菊花生长及遗传诱变效应研究[J].园艺学进展,第五辑:635-639
[12]Banerji B K,Datta S K.Induction of somatic mutation in Chrysanthemum cultivar ‘Anupam'[J].J.Nucleus Agriculture Biology,1991,19:252-256
[13]林祖军,孙纪霞,迮福惠,崔广琴,辛国胜,刘学庆,王建玲.电子束在花卉诱变育种上的应用[J].核农学报,2002,16(6):351-354
[14]丁兵,王江,赵飞,李玉花.高压静电场在菊花天然杂交种选育中的应用[J].东北林业大学学报,2006,34(2):44-45
[15]卢钰,刘军,丰震,张美蓉,韩进,杨传强.菊花育种研究现状及今后的研究方向[J].山东农业大学学报(自然科学版),2004,35(1):145-149
[16]杨保安,范家霖,张建伟,王柏楠,郭淑红.辐射与组培复合育成‘霞光'等14个菊花新品种[J].河南科学,1996,14(1):57-60
[17]Nagatomi S,Miyahira E.Induction of flower mutation comparing with chronic and acute gamma irradiation using tissue culture techniques in Chrysanthemum morifolium ramat[J].Acta Hortculture,2000,508:69-74
[18]王彭伟,李鸿渐,张效平.切花菊单细胞突变育种研究[J].园艺学报,1996,23(3):285-288
[19]Mandal A K A,Chakrabarty D,Datta S K.Application of in vitro techniques in mutation breeding of Chrysanthemum[J].Plant Cell,Tissue and Organ Culture,2000,60(1):33-38
[20]Broertjes C,Lock C A M.Radiation-induced low-temperature tolerate solid mutants of Chrysanthemum morifolium[J].Euphytica,1985,34:97-103
[21]Huttema J B M,Preil W,Gussehoven G C,Schneiderr M.Methods for selection of low-temperature tolerance mutants of Chrysanthemum morifolium Ramat using irradiated cell suspension cultures[J].Plant Breeding,1991,120:140-147
[22]Broertjes C,Keen A.Adventitious shoots:do they develop from one cell?[J].Euphytica,1980,29:73-87
[23]Jerzy M,Zalewska M.Flower color recurrence in Chrysanthemum and gerbera mutants propagated in vitro from meristems and leaf explants[J].Acta Hort,1997,447:611-614
[24]陈发棣,腾年军,房伟民,管志勇.三个菊花品种器官愈伤组织辐射效应的研究[J].中南林学院学报,2003,23(5):49-52
[25]洪亚辉,朱兆海,黄璜,赵燕,徐锋.菊花组织培养与辐射诱变研究[J].湖南农业大学学报(自然科学版),2003,29(6):457-461
[26]Huettermann J,Koehnlein W,Teoule R.Effects of ionizing radiation of DNA:physical,chemical and biological aspects[M].New York,Springer-Verlag New York Inc.,1978,35-36
[27]许伟,李春涛,丁增成,徐宏青.唐菲园林彩叶地被植物品种与辐射诱变育种[J].安徽农业科学,2002,30(3):435-436
[28]孙大容.花生育种学[M].北京:中国农业出版社,1998,53-55
[29]佟道儒.烟草育种学[M].北京:中国农业出版社,1996,37-40
[30]章文成,杨占山,杨淑琴.辐射对血管内皮细胞生物学特性的影响[J].苏州大学学报,2005,25:193-196
[31]牟颖,刘树铮.X射线对小鼠胸腺细胞凋亡的影响[J].中华放射医学与防护杂志,1996,16:296-298
[32]肖辉海,陈良碧.植物体细胞无性系变异育种[J].湖南文理学院学报,2003,15:41-44
[33]FAO/IAEA.Selection in mutation breeding.Vienna IAEA,1984:157-169
[34]洪亚辉,朱兆海,黄璜,赵燕,徐锋.菊花组织培养与辐射诱变研究[J].湖南农业大学学报(自然科学版),2003,29(6):457-461
[35]J F Jackson.Radiation-induced chromosome aberrations[J].Mutation Research,1957,181:17-29
[36]毛炎麟.植物诱变育种学[M].北京:中国农业出版社,1996,97-100
[37]沈守江.诱变因素研究利用的进展[M].北京:原子能出版社,1995,187-189
[38]冯志杰.高等植物的辐射敏感性[M].北京:原子能出版社,1995,75-81
[39]A Anonymous.In vitro technology for mutation breeding.International Atomic Energy Agency,Vienna IAEA,1986,TECDoc-392.58
[40]刘军,许甫荣,郑春开.核科学百年讲座 第八讲 核科学技术在农业领域的应用[J].物理,2003,32(8):553-557
[41]龚守良.电离辐射旁效应[J].吉林大学学报,2003,29:864-866
[42]赵勇,龚守良,刘树铮.低剂量电离辐射对免疫器官局部微环境的免疫调节作用[J].中国病理生理杂志,1998,14:1241
[43]EI-Metainy A,Takagi M,Tano S,Yamaguchi H.Radiation-induced single-strand breaks in the DNA of dormant barley seeds[J].Mutation Research,1971,23:337-344
[44]G P Howland,R W Hart,M L Yette.Repair of DNA strand breaks after gamma-irradiation of protoplasts isolated from cultured wild carrot cells[J].Mutation Research,1975,27:81-87
[45]Tano S,Yamaguchi H.Repair of radiation-induced single-strand breaks in DNA of barley embryos[J].Mutation Research,1977,42:71-78
[46]Sarasin A,Bourre F,Benoit A,Daya-Grosjean L,Gentil A.Molecular analysis of mutagenesis in mammalian cells[J].International journal of radiation biology & related studies in physics,chemistry & medicine,1985,47:479-486
[46]山口彦之,育种学最新四进步,1977,18:17-36
[48]Bartkowiak A,Leyko W,Fried R.A comparative characterization of cytosolic superoxide dismutase/(SOD) from hog liver and erythrocytes[J].Comparative biochemistry and physiology,1979,62(1):61-66
[49]Farrington J A,Ebert M,Land E J,Fletcher K.Bipyridylium quaternary salts and related compounds.V.Pulse radiolysis studies of the reaction of paraquat radical with oxygen.Implications for the mode of action of bipyridyl herbicides[J].Biochimica et biophysica acta,1973,314(3):372-381
[50]H M Hassan,I Fridovich.Regulation of the synthesis of superoxide dismutase in Escherichia coli.Induction by methyl viologen[J].Journal of Biological Chemistry,1977,252(21):7667
[51]王楚桃,白先权,朱立权.~(60)Co-γ射线辐照对密环菌生长及酶活性的影响[J].西南农业大学学报,1998,20(1):19-23
[52]罗以贵,强继业,闫芳芳,刘芮.~(60)Co-γ射线辐照对云烟87烟苗某些生理抗性指标的影响[J].安徽农业科学,2008,36(12):4850-4851
[53]宋道军,徐登益.修复抑制剂对不同敏感性作物超氧化物歧化酶、过氧化氢酶和过氧化物酶活性的影响[J].核农学报,1997,11(2):93-96
[54]胥耀平,刘西平.不同pH、温度和水分胁迫对栾树SOD的影响[J].西北林学院学报,1996,11(4):91-93
[55]曹宗,吴相钰.植物生理学[M].北京:人民教育出版社,1990:60-61
[56]Endo T.Comparison of the effects of gamma-rays and maleic hydrazide on enzyme systems of maize seed[J].Radiation Botany,1967(7):35-40
[57]Giacomelli M,Cerviqni T.Changes in peroxidase activity,ascorbic and dehydroascorbic acid content in Vicia sativa grown under chronic gamma irradiation[J].Radiation Botany,1964(4):395-403
[58]刘芮,强继业,马佶.~(60)Co-γ射线辐射对长寿花某些酶指标的影响[J].青海农林科技,2006,2:31-33
[59]闫芳芳,强继业.~(60)Co-γ射线辐射对红掌几种酶活性及MDA含量的影响[J].云南农业大学学报,2006,21(5):690-692
[60]叶挺梅.~(60)Co-γ射线辐射对红星凤梨生理生化指标的影响[J].安徽农业科学,2006,34(10):2061-2062
[61]吴光升,强继业.γ,射线辐射对袖珍椰子生理指标影响的初步研究[J].种子,2006,25(3):22-24
[62]江文林.~(60)Co-γ射线辐射对一品红7项生理指标的影响[J].江西农业科学,2008,20(4):30-32
[63]叶挺梅,强继业,秦文弟,雷晓.~(60)Co-γ射线辐射对芦荟生理指标的影响[J].安徽农业科学,2005,33(22):234-235
[64]陈宏,靳阳,张喜红,滕佳霞,洪仁远.~(60)Co-γ射线不同剂量射线对水稻幼苗生长、根系蛋白及一些氧化酶活性的影响[J].南开大学学报(自然科学版),1999,32(2):116-119
[65]王泽港,冯敏,胡建伟,高岳,吕海燕,罗时石,郑浩先,马飞,葛才林.γ射线辐照对四种蔬菜叶片SOD活性和MDA含量影响的回归分析[J].核农学,2005,19(2):134-137
[66]Warfield D L,Nilan R A,Witters R E.Effect of ethylene and ionizing radiation on Saintpaulia peroxidase activity.Scientific paper NO.4333,1973,1:18
[67]R Balachandran,P C Kesavan.Effect of caffeine on peroxidase activity and gamma-ray-induced oxic and anoxic damage in Hordeum[J].Environmental and Experimental Botany.1978(6):375-379
[68]M Inoue,R B van Huystee.Effects of caffeine treatment alone and in combination with gamma exposure on cultured peanut cells[J].Canadian Journal of Botany.1984,62(9):1890-1895
[69]白玲玉,马玉珠,华珞,韦东甫.低剂量~(60)Co-γ射线辐照对白菜苗期酶活性及同工酶的影响[J].核农学报,1996,10(1):21-24
[70]强继业,陈宗瑜,李佛琳,高晓燕,许银莲.~(60)Co-γ射线辐照对球根海棠、蝴蝶兰生长及SOD和CAT活性的影响[J].种子,2004,23(4):8-11
[71]陈云飞,强继业.γ射线辐照对植物保护性酶活性和MDA含量的影响[J].安徽农业科学,2006,34(10):2034-2035
[72]方允中,郑荣梁.自由基生物学的理论与应用[M].北京:科学出版社(第一版),2002,213
[73]屈集平,高山林.盾叶薯蓣多倍体和辐射诱变株系部分生理活性的比较[J].药物生物技术,2007,14(3):196-200
[74]吴双,顾月华,赵凌,刘佳娟.软X射线同步辐射对小麦细胞膜脂质过氧化产物及相关酶活性的影响[J].核技术,2003,26(8):577-581
[75]张献清,曾学平,穆士杰,吴道澄,夏爱军,吴源茹,易静.低剂量γ射线辐照对人红细胞免疫功能和血清抗氧化酶活性的影响[J].中国医学物理学杂志,2008,25(3):667-669
[76]林植芳,李双顺,桂林珠,孙古畴,郭俊彦.水稻叶片的衰老与超氧化物歧化酶活性及脂质过氧化作用的关系[J].植物学报(英文版),1984,26(6):605-615
[77]Weatherly P E.Studies on water relation of cotton plant.I.The field measurement of water deficits in leaves[J].New Phytologist,1950,49:91-97
[78]张宪政.植物叶绿素含量测定——丙酮乙醇混合液法[J].辽宁农业科学,1986(3):26-28
[79]苏正淑,张宪政.几种测定叶绿素含量的方法比较[J].植物生理学通讯1989(5):77-78
[80]Beyer W F,Fridovich I.Assaying for superoxide dismutase activity:Some large consequences of minor changes in condition[J].Anal Biochem.,1987,161:559-566
[81]Wang W X,Vinocur B,Altman A.Plant responses to drought,salinity and extreme temperatures:towards genetic engineering for stress tolerance[J].Planta,2003,218:1-14
[82]Stanislawa Pukacka,Ewelina Ratajczak.Production and scavenging of reactive oxygen species in Fagus sylvatica seeds during storage at varied temperature and humidity[J].Journal of Plant Physiology,2005,162(8):873-885
[83]Heath R L,Packer L.Photoperoxidation in isolated chloroplasts.I.Kinetics and stoichiometry of fatty acid peroxidation[J].Arch.Biochem.Biophys.,1968,125:189-198
[84]Alexieva V,Sergiev I,Mapelli S,et al.The effect of drought and ultraviolet radiation on growth and stress markers in pea and wheat[J].Plant Cell Environ.,2001,24:1337-1344
[85]Nakano Y,Asada K.Hydrogen peroxide is scavenged by ascorbate-specific peroxidase in spinach chloroplasts[J].Plant Cell Physiol,1981,22:867-880
[86]Halliwell B,Foyer C H.Properties and physiological function of a glutathione reductase purified from spinach leaves by affinity chromatography[J].Planta,1978,139:9-17
[87]Bradford M M.A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding[J].Anal Biochem.,1976,72:248-254
[88]Levitt A D.Responses of plant to environmental stresses Volume 1[M].New York:Academic Press,1990,55
[89]王晶,刘录祥,赵世荣,郭会君,赵林姝,陈文华.~(60)Co-γ射线对菊花组培苗的诱变效应[J].农业生物技术学报.2006,14:241-244
[90]高健.钴-60Gammma 射线辐照中国水仙的诱变效应和机理研究[博士学位论文].北京:中国农业科学院,2000.61-64
[91]师生波,贲桂英,赵新全,韩发.增强UV-B辐射对高山植物麻花艽净光和速率的影响[J].植物生态学报,2001,25(5):520-524
[92]Costa H,Gallego S M,Tomaro M L.Effect of UV-B radiation on antioxidant defense system in sunflower cotyledons[J].Plant Sci.,2002,162:939-945
[93]徐冠仁.植物诱变育种学[M].北京:中国农业出版社,1996(第一版),1996,91-93
[94]Bowler C,van Montagu M,Inz(?) D.Superoxide dismutase and stress tolerance[J].Annu.Rev.Plant Physiol.Plant Mol.Biol.,1992,43:83-116.
[95]Allen R D.Dissection of oxidative stress tolerance using transgenic plants[J].Plant Physiol.,1995,107:1049-1054.
[2]高键,卢惠萍.花卉辐射诱变育种研究进展[J].安徽农业大学学报(自然科学版),2000,(3):22-24
[3]许耀奎.作物诱变育种[M].上海:上海科技出版社,1985,25
[4]Broertjes C,Roeet S,Bokelmann G S.Mutation breeding of Chrysanthemum morifolium Ram using in vivo and in vitro adventitious bud techniques[J].Euphytica,1976,25:11-19
[5]Broertjes C,Van Harten A M.Application of mutation breeding methods.In:A J Abbott and R R Attkin(Editors):Improving vegetatively propagated crops London:Academic Press.1987,335-338
[6]王海燕,何小弟,黄永高.菊花组织培养技术在育种上的应用研究[J].江苏林业科技,2003,30(4):25-26
[7]齐孟文,王化国.我国花卉辐射育种的进展与剖析[J].核农学通报,1997,18(6),288-290
[8]刘金勇.菊花品种遗传改良的研究进展[J].湖南林业科技,2004,31(1),49-52
[9]郭安熙,范家霖,杨保安,王柏楠,张建伟,陈树国,王世荣,王然.菊花花色辐射诱变研究[J].核农学报,1997,11(2):65-73
[10]傅玉兰,郑路.冬菊新品种选育[J].安徽农业大学学报,1994,(1)59-62
[11]黄建昌,肖艳.~(60)Co-γ射线处理对菊花生长及遗传诱变效应研究[J].园艺学进展,第五辑:635-639
[12]Banerji B K,Datta S K.Induction of somatic mutation in Chrysanthemum cultivar ‘Anupam'[J].J.Nucleus Agriculture Biology,1991,19:252-256
[13]林祖军,孙纪霞,迮福惠,崔广琴,辛国胜,刘学庆,王建玲.电子束在花卉诱变育种上的应用[J].核农学报,2002,16(6):351-354
[14]丁兵,王江,赵飞,李玉花.高压静电场在菊花天然杂交种选育中的应用[J].东北林业大学学报,2006,34(2):44-45
[15]卢钰,刘军,丰震,张美蓉,韩进,杨传强.菊花育种研究现状及今后的研究方向[J].山东农业大学学报(自然科学版),2004,35(1):145-149
[16]杨保安,范家霖,张建伟,王柏楠,郭淑红.辐射与组培复合育成‘霞光'等14个菊花新品种[J].河南科学,1996,14(1):57-60
[17]Nagatomi S,Miyahira E.Induction of flower mutation comparing with chronic and acute gamma irradiation using tissue culture techniques in Chrysanthemum morifolium ramat[J].Acta Hortculture,2000,508:69-74
[18]王彭伟,李鸿渐,张效平.切花菊单细胞突变育种研究[J].园艺学报,1996,23(3):285-288
[19]Mandal A K A,Chakrabarty D,Datta S K.Application of in vitro techniques in mutation breeding of Chrysanthemum[J].Plant Cell,Tissue and Organ Culture,2000,60(1):33-38
[20]Broertjes C,Lock C A M.Radiation-induced low-temperature tolerate solid mutants of Chrysanthemum morifolium[J].Euphytica,1985,34:97-103
[21]Huttema J B M,Preil W,Gussehoven G C,Schneiderr M.Methods for selection of low-temperature tolerance mutants of Chrysanthemum morifolium Ramat using irradiated cell suspension cultures[J].Plant Breeding,1991,120:140-147
[22]Broertjes C,Keen A.Adventitious shoots:do they develop from one cell?[J].Euphytica,1980,29:73-87
[23]Jerzy M,Zalewska M.Flower color recurrence in Chrysanthemum and gerbera mutants propagated in vitro from meristems and leaf explants[J].Acta Hort,1997,447:611-614
[24]陈发棣,腾年军,房伟民,管志勇.三个菊花品种器官愈伤组织辐射效应的研究[J].中南林学院学报,2003,23(5):49-52
[25]洪亚辉,朱兆海,黄璜,赵燕,徐锋.菊花组织培养与辐射诱变研究[J].湖南农业大学学报(自然科学版),2003,29(6):457-461
[26]Huettermann J,Koehnlein W,Teoule R.Effects of ionizing radiation of DNA:physical,chemical and biological aspects[M].New York,Springer-Verlag New York Inc.,1978,35-36
[27]许伟,李春涛,丁增成,徐宏青.唐菲园林彩叶地被植物品种与辐射诱变育种[J].安徽农业科学,2002,30(3):435-436
[28]孙大容.花生育种学[M].北京:中国农业出版社,1998,53-55
[29]佟道儒.烟草育种学[M].北京:中国农业出版社,1996,37-40
[30]章文成,杨占山,杨淑琴.辐射对血管内皮细胞生物学特性的影响[J].苏州大学学报,2005,25:193-196
[31]牟颖,刘树铮.X射线对小鼠胸腺细胞凋亡的影响[J].中华放射医学与防护杂志,1996,16:296-298
[32]肖辉海,陈良碧.植物体细胞无性系变异育种[J].湖南文理学院学报,2003,15:41-44
[33]FAO/IAEA.Selection in mutation breeding.Vienna IAEA,1984:157-169
[34]洪亚辉,朱兆海,黄璜,赵燕,徐锋.菊花组织培养与辐射诱变研究[J].湖南农业大学学报(自然科学版),2003,29(6):457-461
[35]J F Jackson.Radiation-induced chromosome aberrations[J].Mutation Research,1957,181:17-29
[36]毛炎麟.植物诱变育种学[M].北京:中国农业出版社,1996,97-100
[37]沈守江.诱变因素研究利用的进展[M].北京:原子能出版社,1995,187-189
[38]冯志杰.高等植物的辐射敏感性[M].北京:原子能出版社,1995,75-81
[39]A Anonymous.In vitro technology for mutation breeding.International Atomic Energy Agency,Vienna IAEA,1986,TECDoc-392.58
[40]刘军,许甫荣,郑春开.核科学百年讲座 第八讲 核科学技术在农业领域的应用[J].物理,2003,32(8):553-557
[41]龚守良.电离辐射旁效应[J].吉林大学学报,2003,29:864-866
[42]赵勇,龚守良,刘树铮.低剂量电离辐射对免疫器官局部微环境的免疫调节作用[J].中国病理生理杂志,1998,14:1241
[43]EI-Metainy A,Takagi M,Tano S,Yamaguchi H.Radiation-induced single-strand breaks in the DNA of dormant barley seeds[J].Mutation Research,1971,23:337-344
[44]G P Howland,R W Hart,M L Yette.Repair of DNA strand breaks after gamma-irradiation of protoplasts isolated from cultured wild carrot cells[J].Mutation Research,1975,27:81-87
[45]Tano S,Yamaguchi H.Repair of radiation-induced single-strand breaks in DNA of barley embryos[J].Mutation Research,1977,42:71-78
[46]Sarasin A,Bourre F,Benoit A,Daya-Grosjean L,Gentil A.Molecular analysis of mutagenesis in mammalian cells[J].International journal of radiation biology & related studies in physics,chemistry & medicine,1985,47:479-486
[46]山口彦之,育种学最新四进步,1977,18:17-36
[48]Bartkowiak A,Leyko W,Fried R.A comparative characterization of cytosolic superoxide dismutase/(SOD) from hog liver and erythrocytes[J].Comparative biochemistry and physiology,1979,62(1):61-66
[49]Farrington J A,Ebert M,Land E J,Fletcher K.Bipyridylium quaternary salts and related compounds.V.Pulse radiolysis studies of the reaction of paraquat radical with oxygen.Implications for the mode of action of bipyridyl herbicides[J].Biochimica et biophysica acta,1973,314(3):372-381
[50]H M Hassan,I Fridovich.Regulation of the synthesis of superoxide dismutase in Escherichia coli.Induction by methyl viologen[J].Journal of Biological Chemistry,1977,252(21):7667
[51]王楚桃,白先权,朱立权.~(60)Co-γ射线辐照对密环菌生长及酶活性的影响[J].西南农业大学学报,1998,20(1):19-23
[52]罗以贵,强继业,闫芳芳,刘芮.~(60)Co-γ射线辐照对云烟87烟苗某些生理抗性指标的影响[J].安徽农业科学,2008,36(12):4850-4851
[53]宋道军,徐登益.修复抑制剂对不同敏感性作物超氧化物歧化酶、过氧化氢酶和过氧化物酶活性的影响[J].核农学报,1997,11(2):93-96
[54]胥耀平,刘西平.不同pH、温度和水分胁迫对栾树SOD的影响[J].西北林学院学报,1996,11(4):91-93
[55]曹宗,吴相钰.植物生理学[M].北京:人民教育出版社,1990:60-61
[56]Endo T.Comparison of the effects of gamma-rays and maleic hydrazide on enzyme systems of maize seed[J].Radiation Botany,1967(7):35-40
[57]Giacomelli M,Cerviqni T.Changes in peroxidase activity,ascorbic and dehydroascorbic acid content in Vicia sativa grown under chronic gamma irradiation[J].Radiation Botany,1964(4):395-403
[58]刘芮,强继业,马佶.~(60)Co-γ射线辐射对长寿花某些酶指标的影响[J].青海农林科技,2006,2:31-33
[59]闫芳芳,强继业.~(60)Co-γ射线辐射对红掌几种酶活性及MDA含量的影响[J].云南农业大学学报,2006,21(5):690-692
[60]叶挺梅.~(60)Co-γ射线辐射对红星凤梨生理生化指标的影响[J].安徽农业科学,2006,34(10):2061-2062
[61]吴光升,强继业.γ,射线辐射对袖珍椰子生理指标影响的初步研究[J].种子,2006,25(3):22-24
[62]江文林.~(60)Co-γ射线辐射对一品红7项生理指标的影响[J].江西农业科学,2008,20(4):30-32
[63]叶挺梅,强继业,秦文弟,雷晓.~(60)Co-γ射线辐射对芦荟生理指标的影响[J].安徽农业科学,2005,33(22):234-235
[64]陈宏,靳阳,张喜红,滕佳霞,洪仁远.~(60)Co-γ射线不同剂量射线对水稻幼苗生长、根系蛋白及一些氧化酶活性的影响[J].南开大学学报(自然科学版),1999,32(2):116-119
[65]王泽港,冯敏,胡建伟,高岳,吕海燕,罗时石,郑浩先,马飞,葛才林.γ射线辐照对四种蔬菜叶片SOD活性和MDA含量影响的回归分析[J].核农学,2005,19(2):134-137
[66]Warfield D L,Nilan R A,Witters R E.Effect of ethylene and ionizing radiation on Saintpaulia peroxidase activity.Scientific paper NO.4333,1973,1:18
[67]R Balachandran,P C Kesavan.Effect of caffeine on peroxidase activity and gamma-ray-induced oxic and anoxic damage in Hordeum[J].Environmental and Experimental Botany.1978(6):375-379
[68]M Inoue,R B van Huystee.Effects of caffeine treatment alone and in combination with gamma exposure on cultured peanut cells[J].Canadian Journal of Botany.1984,62(9):1890-1895
[69]白玲玉,马玉珠,华珞,韦东甫.低剂量~(60)Co-γ射线辐照对白菜苗期酶活性及同工酶的影响[J].核农学报,1996,10(1):21-24
[70]强继业,陈宗瑜,李佛琳,高晓燕,许银莲.~(60)Co-γ射线辐照对球根海棠、蝴蝶兰生长及SOD和CAT活性的影响[J].种子,2004,23(4):8-11
[71]陈云飞,强继业.γ射线辐照对植物保护性酶活性和MDA含量的影响[J].安徽农业科学,2006,34(10):2034-2035
[72]方允中,郑荣梁.自由基生物学的理论与应用[M].北京:科学出版社(第一版),2002,213
[73]屈集平,高山林.盾叶薯蓣多倍体和辐射诱变株系部分生理活性的比较[J].药物生物技术,2007,14(3):196-200
[74]吴双,顾月华,赵凌,刘佳娟.软X射线同步辐射对小麦细胞膜脂质过氧化产物及相关酶活性的影响[J].核技术,2003,26(8):577-581
[75]张献清,曾学平,穆士杰,吴道澄,夏爱军,吴源茹,易静.低剂量γ射线辐照对人红细胞免疫功能和血清抗氧化酶活性的影响[J].中国医学物理学杂志,2008,25(3):667-669
[76]林植芳,李双顺,桂林珠,孙古畴,郭俊彦.水稻叶片的衰老与超氧化物歧化酶活性及脂质过氧化作用的关系[J].植物学报(英文版),1984,26(6):605-615
[77]Weatherly P E.Studies on water relation of cotton plant.I.The field measurement of water deficits in leaves[J].New Phytologist,1950,49:91-97
[78]张宪政.植物叶绿素含量测定——丙酮乙醇混合液法[J].辽宁农业科学,1986(3):26-28
[79]苏正淑,张宪政.几种测定叶绿素含量的方法比较[J].植物生理学通讯1989(5):77-78
[80]Beyer W F,Fridovich I.Assaying for superoxide dismutase activity:Some large consequences of minor changes in condition[J].Anal Biochem.,1987,161:559-566
[81]Wang W X,Vinocur B,Altman A.Plant responses to drought,salinity and extreme temperatures:towards genetic engineering for stress tolerance[J].Planta,2003,218:1-14
[82]Stanislawa Pukacka,Ewelina Ratajczak.Production and scavenging of reactive oxygen species in Fagus sylvatica seeds during storage at varied temperature and humidity[J].Journal of Plant Physiology,2005,162(8):873-885
[83]Heath R L,Packer L.Photoperoxidation in isolated chloroplasts.I.Kinetics and stoichiometry of fatty acid peroxidation[J].Arch.Biochem.Biophys.,1968,125:189-198
[84]Alexieva V,Sergiev I,Mapelli S,et al.The effect of drought and ultraviolet radiation on growth and stress markers in pea and wheat[J].Plant Cell Environ.,2001,24:1337-1344
[85]Nakano Y,Asada K.Hydrogen peroxide is scavenged by ascorbate-specific peroxidase in spinach chloroplasts[J].Plant Cell Physiol,1981,22:867-880
[86]Halliwell B,Foyer C H.Properties and physiological function of a glutathione reductase purified from spinach leaves by affinity chromatography[J].Planta,1978,139:9-17
[87]Bradford M M.A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding[J].Anal Biochem.,1976,72:248-254
[88]Levitt A D.Responses of plant to environmental stresses Volume 1[M].New York:Academic Press,1990,55
[89]王晶,刘录祥,赵世荣,郭会君,赵林姝,陈文华.~(60)Co-γ射线对菊花组培苗的诱变效应[J].农业生物技术学报.2006,14:241-244
[90]高健.钴-60Gammma 射线辐照中国水仙的诱变效应和机理研究[博士学位论文].北京:中国农业科学院,2000.61-64
[91]师生波,贲桂英,赵新全,韩发.增强UV-B辐射对高山植物麻花艽净光和速率的影响[J].植物生态学报,2001,25(5):520-524
[92]Costa H,Gallego S M,Tomaro M L.Effect of UV-B radiation on antioxidant defense system in sunflower cotyledons[J].Plant Sci.,2002,162:939-945
[93]徐冠仁.植物诱变育种学[M].北京:中国农业出版社,1996(第一版),1996,91-93
[94]Bowler C,van Montagu M,Inz(?) D.Superoxide dismutase and stress tolerance[J].Annu.Rev.Plant Physiol.Plant Mol.Biol.,1992,43:83-116.
[95]Allen R D.Dissection of oxidative stress tolerance using transgenic plants[J].Plant Physiol.,1995,107:1049-1054.