皱紫菜色素突变体的分离与特性分析
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摘要
用不同剂量的紫外线(UV)对皱紫菜(Pyropia crispata)野生型品系(PC-WT)的壳孢子萌发体进行辐照,4周后,叶状体上均出现了黄绿、宝石红、黄橙等颜色的色素变异细胞块,经统计表明,剂量为80μW/cm~2的诱变效果最好。用酶解法将含有色素变异细胞块的叶状体细胞单个分离出来进行培养,从再生叶状体中分离到PC-JH(桔黄褐色)、PC-ZH(宝石红色)、PC-HL(黄绿色)、PC-QL(浅绿色)等纯色突变体,通过单性生殖分别获得纯合品系。各突变品系的F_1叶状体的颜色和形态均一,且与其母体叶状体相同,表明各突变品系的颜色等突变性状均能稳定遗传。与野生型品系相比,各色素突变品系的F_1叶状体的活体吸收光谱、色素蛋白的含量及相互间的比值均发生了明显变化,且相互之间也存在明显差异。综上所述,紫外线对皱紫菜叶状体具有良好的诱变效果,本实验所获得的色素突变品系,将成为皱紫菜遗传育种研究的重要材料。
The conchospore gametophytic blades of the wild-type strain of Pyropia crispata were irradiated with different doses of ultraviolet ray. After being cultured for about 4 weeks, many color-mutated cell clusters including yellow green, ruby red, yellow orange and so on,appeared on the irradiated blade. The statistical analysis showed most of the color-mutated cell types are yellow and the dose of 80 μW/cm~2 attained the best mutagenic effect. Single mutated cell was isolated enzymatically from the color-mutated blades and was regenerated into blades, in which some single-color mutants were selected: PC-JH(orange brown), PC-ZH(ruby red), PC-HL(yellow green) and PC-QL(light green). Their homozygous strain were gotten by parthenogenesis, respectively. The color and morphology of the F_1 gametophytic blades of each mutant strain showed uniform color and were same with its mother blade. The results indicated mutant characters such as color can be stably inherited in each mutant strain. Each mutant strain featured their own characteristics in in vivo absorption spectra of the blade, contents of chlorophyll a(Chl.a), phycocyanin(PC), phycoerythrin(PE) and their ratio, different from wild-type strain's. Compared with the wild-type strain(PC-WT), the content of PC and the value of PC/Chl.a of the two green-type strains(PC-HL, PC-QL) increased significantly, while the value of PE/PC decreased significantly and was close to one. The content of three pigment proteins in the orange-yellow mutant(PC-JH) was reduced, and the PC/Chl.a value was also lower, but the PE/Chl.a and PE/PC values were higher. The PE content of the date-red mutant(PC-ZH) increased significantly. Correspondingly, the values of PE/Chl.a and PE/PC also increased significantly. These results indicated that UV had a good mutagenic effect on Pyropia crispata blade, and the obtained four mutant strains were stable in color. They will become important materials for genetic breeding in Pyropia crispata.
The conchospore gametophytic blades of the wild-type strain of Pyropia crispata were irradiated with different doses of ultraviolet ray. After being cultured for about 4 weeks, many color-mutated cell clusters including yellow green, ruby red, yellow orange and so on,appeared on the irradiated blade. The statistical analysis showed most of the color-mutated cell types are yellow and the dose of 80 μW/cm~2 attained the best mutagenic effect. Single mutated cell was isolated enzymatically from the color-mutated blades and was regenerated into blades, in which some single-color mutants were selected: PC-JH(orange brown), PC-ZH(ruby red), PC-HL(yellow green) and PC-QL(light green). Their homozygous strain were gotten by parthenogenesis, respectively. The color and morphology of the F_1 gametophytic blades of each mutant strain showed uniform color and were same with its mother blade. The results indicated mutant characters such as color can be stably inherited in each mutant strain. Each mutant strain featured their own characteristics in in vivo absorption spectra of the blade, contents of chlorophyll a(Chl.a), phycocyanin(PC), phycoerythrin(PE) and their ratio, different from wild-type strain's. Compared with the wild-type strain(PC-WT), the content of PC and the value of PC/Chl.a of the two green-type strains(PC-HL, PC-QL) increased significantly, while the value of PE/PC decreased significantly and was close to one. The content of three pigment proteins in the orange-yellow mutant(PC-JH) was reduced, and the PC/Chl.a value was also lower, but the PE/Chl.a and PE/PC values were higher. The PE content of the date-red mutant(PC-ZH) increased significantly. Correspondingly, the values of PE/Chl.a and PE/PC also increased significantly. These results indicated that UV had a good mutagenic effect on Pyropia crispata blade, and the obtained four mutant strains were stable in color. They will become important materials for genetic breeding in Pyropia crispata.
引文
[1] SUTHERLAND J E,LINDSTROM S C,NELSON W A,et al.A new look at an ancient order:genericrevision of the Bangiales (Rhodophyta)[J].Journal of Phycology,2011,47(5):1131-1151.
[2] 曾呈奎,张德瑞,张峻甫,等.中国经济海藻志[M].北京:科学出版社,1962:103-104.ZENG C K,ZHANG D R,ZHANG J F,et al.China’s economic seaweed[M].Beijing:Science Press,1962:103-104.
[3] 王华芝,严兴洪,李琳.条斑紫菜(Porphyra yezoensis)耐高温品系的筛选及特性分析[J].海洋与湖沼,2012,43(2):363-369.WANG H Z,YAN X H,LI L.Selection and characterization of a high-temperature resistant strain of Porphyra yezoensis Ueda(Bangiales,Rhodophyta)[J].Oceanologia et Limnologia Sinica,2012,43(2):363-369.
[4] 陈伟洲,许俊宾,吴文婷,等.三种紫菜叶状体对高温胁迫的生理响应[J].热带海洋学报,2015,34(1):49-55.CHEN W Z,XU J B,WU W T,et al.Physiological responses of three species of Pyropia thallus to high temperature stress[J].Journal of Tropical Oceanography,2015,34(1):49-55.
[5] ARUGA Y.Color and the pigments of Porphyra yezoensis[J].Iden,1980,34(9):8-13.
[6] ARUGA Y,MIURA A.In vivo absorption spectra and pigment contents of the two types of color mutants of Porphyra[J].Japanese Journal of Phycology,1984,32(3):243-250.
[7] OHME M,KUNIFUJI Y,MIURA A.Cross experiments of the color mutants in Porphyra yezoensis Ueda[J].Japanese Journal of Phycology,1986,34:101-106.
[8] YAN X H,LI L,ARUGA Y.Genetic analysis of the position of meiosis in Porphyra haitanensis Chang et Zheng (Bangiales,Rhodophyta)[J].Journal of Applied Phycology,2005,17(6):467-473.
[9] 严兴洪,李琳,有贺佑胜.坛紫菜减数分裂位置的杂交试验分析[J].水产学报,2006,30(1):1-8.YAN X H,LI L,ARUGA Y.Cross experiments and analysis of the position of meiosis in Porphyra haitanensis (Rhodophyta)[J].Journal of Fisheries of China,2006,30(1):1-8.
[10] WANG J F,ZHU J Y,ZHOU W,et al.Early development patterns and morphogenesis of blades in four species of Porphyra (Bangiales,Rhodophyta)[J].Journal of Applied Phycology,2010,22(3):297-303.
[11] 吴宏肖,严兴洪,宋武林,等.坛紫菜与Pyropia radi种间杂交重组优良品系的选育与特性分析[J].水产学报,2014,38(8):1079-1088.WU H X,YAN X H,SONG W L,et al.Selection and characterization of an improved strain produced by genetic recombinant of interspecific hybridization between Pyropia haitanensis and Pyropia radi[J].Journal of Fisheries of China,2014,38(8):1079-1088.
[12] 刘美君,黄林彬,严兴洪.条斑紫菜种内杂交优良品系的筛选与特性分析[J].中国水产科学,2015,22(1):33-43.LIU M J,HUANG L B,YAN X H.Isolation and characterization of the improved strain HW-4 by in-traspecific hybridization in Pyropia yezoensis[J].Journal of Fishery Sciences of China,2015,22(1):33-43.
[13] DING H C,Lü F,WU H X,et al.Selection and characterization of an improved strain produced by inter-species hybridization between Pyropia sp.from India and Pyropia haitanensis from China[J].Aquaculture,2018,491:177-183.
[14] 李淑平,严兴洪.60Co-γ射线辐照对长紫菜的诱变效果及优良品系分离与特性分析[J].海洋学报,2015,37(10):69-79.LI S P,YAN X H.Isolation and characterization of an improved strain of Pyropia dentata (Bangiales,Rhodophyta) after being irradiated by 60Co-γ ray[J].Acta Oceanologica Sinica,2015,37(10):69-79.
[15] 戴继勋,张全启,包振民,等.紫菜原生质体的纯系培育、诱变处理和种间细胞融合的研究[J].海洋与湖沼,1990,21(3):293-296.DAI J X,ZHANG Q Q,BAO Z M,et al.Studies on the pure line culture,mutagenization and interspecific fusion of Porphyra protoplasts[J].Oceanologia et Limnologia Sinica,1990,21(3):293-296.
[16] 严兴洪.紫外线辐射与条斑紫菜原生质体后代发育和变异[J].上海水产大学学报,1992,1(1/2):71-78.YAN X H.Effects of UV irradiation on the development and variations of protoplast progenies in Porphyra yezoensis[J].Journal of Shanghai Fisheries University,1992,1(1/2):71-78.
[17] 刘宝.角蛋白降解菌的紫外线和硫酸二乙酯复合诱变[D].南京:南京农业大学,2012:14.LIU B.Complex mutagenesis of keratinose degrading strain with ultraviolet ray and diethyl sufate[D].Nanjing:Nanjing Agricultural University,2012:14.
[18] 何培民,秦松,严小军,等.海藻生物技术及其应用[M].北京:化学工业出版社,2007:59-61.HE P M,QIN S,YAN X J,et al.Seaweed biological technology and application[M].Beijing:Chemical Industry Press,2007:59-61.
[19] 严兴洪,李琳,陈俊华,等.坛紫菜的单性生殖与遗传纯系分离[J].高技术通讯,2007,17(2):205-210.YAN X H,LI L,CHEN J H,et al.Parthenogenesis and isolation of genetic pure strains in Porphyra haitanensis (Bangiales,Rhodophyta)[J].High Technology Letters,2007,17(2):205-210.
[20] 王素娟,张小平,徐志东,等.坛紫菜营养细胞和原生质体培养的研究I[J].海洋与湖沼,1986,17(3):217-221.WANG S J,ZHANG X P,XU Z D,et al.A study on the cultivation of the vegetative cells and protoplasts of P.yezoensis[J].Oceanologia et Limnologia Sinica,1986,17(3):217-221.
[21] KATO M,ARUGA Y.Comparative studies on the growth and photosynthesis of the pigmentation mutants of Porphyra yezoensis in laboratory culture[J].Japanese Journal of Phycology,1984,32:333-347.
[22] 赵东旭,杨新芳,刘亚丽,等.紫外线辐射对小麦幼苗生长的影响[J].河南科学,1996,14(s1):32-35.ZHAO D X,YANG X F,LIU Y L,et al.Effects of ultraviolet radiation on growth of wheat seedlings[J].Henan Sciences,1996,14(s1):32-35.
[23] 张敏.紫外线对霞多丽葡萄品质及相关生化物质的影响[D].晋中:山西农业大学,2016:16-30.ZHANG M.The affect of ultraviolet radiation to the quality and relevant substances of Chardonnay[D].Jinzhong:Shanxi Agricultural University,2016:16-30.
[24] 岳明,王勋陵.紫外线辐射对小麦和燕麦竞争性平衡的影响——小麦和燕麦生物量结构与冠层结构[J].环境科学学报,1999,19(5):526-531.YUE M,WANG Y L.Effect of enhanced ultraviolet radiation on competitive balance of wheat and oat-the structure of biomass and canopy[J].Acta Scientiae Circumstantiae,1999,19(5):526-531.
[25] 陈爱芹.紫外线诱变处理对马铃薯茎段再生植株生理生化变异的影响[D].长沙:湖南农业大学,2008:26-34.CHEN A Q.Physiological and biochemical variation of the potato stem section regeneration plants by ultraviolet radiation[D].Changsha:Hunan Agricultural University,2008:26-34.
[26] 李波,贾秀峰,焦德志,等.紫外线诱变苜蓿愈伤组织抗旱性的研究[J].干旱地区农业研究,2005,26(6):103-105.LI B,JIA X F,JIAO D Z,et al.Influence of ultraviolet ray to drought resistance of alfalfa calluses[J].Agricultural Research in the Arid Areas,2005,26(6):103-105.
[27] 黄世玉,王志勇.紫外线辐射坛紫菜离体细胞的初步试验[J].集美大学学报,1997,2(1):25-30.HUANG S Y,WANG Z Y.Preliminary experiment on isolated cells from Porphyra haitanensis (Rhodophyta) radiated by ultraviolet ray[J].Journal of Jimei University,1997,2(1):25-30.
[28] 严兴洪,张淑娟,黄林彬.60Co-γ射线对条斑紫菜(Porphyra yezoensis)的诱变效果与色素突变体分离[J].海洋与湖沼,2009,40(1):56-61.YAN X H,ZHANG S J,HUANG L B.Induction and isolation of pigmentation mutants of Porphyra yezoensis Ueda (Bangiales,Rhodophyta) by 60Co-γ ray irradiation[J].Oceanologia et Limnologia Sinica,2009,40(1):56-61.
[29] YAN X H,ARUGA Y.Induction of pigmentation mutants by treatment of monospore germlings with NNG in Porphyra yezoensis Ueda (Bangiales,Rhodophyta)[J].Algae,1997,12(1):39-52.
[30] YAN X H,FUJITA Y,ARUGA Y.Induction and characterization of pigmentation mutants in Porphyra yezoensis(Bangiales,Rhodophyta)[J].Journal of Applied Phycology,2000,12(1):69-81.
[31] 王素娟,马凌波,许璞,等.60Co-γ射线诱变条斑紫菜丝状体的研究[J].海洋科学,1999(4):43-47.WANG S J,MA L B,XU P,et al.Study on using gamma-ray to induce mutation in conchocelis of Porphyra yezoensis[J].Marine Sciences,1999(4):43-47.
[32] 李琳,严兴洪.坛紫菜绿色突变体的分离与特性分析[J].上海水产大学学报,2006,15(1):30-35.LI L,YAN X H.Isolation and characterization of a green type mutant in Porphyra haitanensis Chang et Zheng (Bangiales,Rhodophyta)[J].Journal of Shanghai Fisheries University,2006,15(1):30-35.
[33] 匡梅,许璞,王素娟.γ-射线对条斑紫菜和坛紫菜诱变作用的初步研究[J].上海水产大学学报,1997,6(4):241-245.KUANG M,XU P,WANG S J.A preliminary study of the mutagensis of γ-rays on Porphyra yezoensis and P.haitanensis[J].Journal of Shanghai Fisheries University,1997,6(4):241-245.
[34] 严兴洪,梁志强,宋武林,等.坛紫菜人工色素突变体的诱变与分离[J].水产学报,2005,29(2):166-172.YAN X H,LIANG Z Q,SONG W L,et al.Induction and isolation of artificial pigmentation mutants in Porphyra haitanensis Chang et Zheng (Bangiales,Rhodophyta)[J].Journal of Fisheries of China,2005,29(2):166-172.
[35] 严兴洪,田中次郎,有贺佑胜.条斑紫菜色彩突变体的诱导、分离和特性分析[J].水产学报,2000,24(3):221-228.YAN X H,TANAKA J,ARUGA Y.Isolation and characterization of pigmentation mutants in Porphyra yezoensis Ueda (Bangiales,Rhodophyta)[J].Journal of Fisheries of China,2000,24(3):221-228.
[2] 曾呈奎,张德瑞,张峻甫,等.中国经济海藻志[M].北京:科学出版社,1962:103-104.ZENG C K,ZHANG D R,ZHANG J F,et al.China’s economic seaweed[M].Beijing:Science Press,1962:103-104.
[3] 王华芝,严兴洪,李琳.条斑紫菜(Porphyra yezoensis)耐高温品系的筛选及特性分析[J].海洋与湖沼,2012,43(2):363-369.WANG H Z,YAN X H,LI L.Selection and characterization of a high-temperature resistant strain of Porphyra yezoensis Ueda(Bangiales,Rhodophyta)[J].Oceanologia et Limnologia Sinica,2012,43(2):363-369.
[4] 陈伟洲,许俊宾,吴文婷,等.三种紫菜叶状体对高温胁迫的生理响应[J].热带海洋学报,2015,34(1):49-55.CHEN W Z,XU J B,WU W T,et al.Physiological responses of three species of Pyropia thallus to high temperature stress[J].Journal of Tropical Oceanography,2015,34(1):49-55.
[5] ARUGA Y.Color and the pigments of Porphyra yezoensis[J].Iden,1980,34(9):8-13.
[6] ARUGA Y,MIURA A.In vivo absorption spectra and pigment contents of the two types of color mutants of Porphyra[J].Japanese Journal of Phycology,1984,32(3):243-250.
[7] OHME M,KUNIFUJI Y,MIURA A.Cross experiments of the color mutants in Porphyra yezoensis Ueda[J].Japanese Journal of Phycology,1986,34:101-106.
[8] YAN X H,LI L,ARUGA Y.Genetic analysis of the position of meiosis in Porphyra haitanensis Chang et Zheng (Bangiales,Rhodophyta)[J].Journal of Applied Phycology,2005,17(6):467-473.
[9] 严兴洪,李琳,有贺佑胜.坛紫菜减数分裂位置的杂交试验分析[J].水产学报,2006,30(1):1-8.YAN X H,LI L,ARUGA Y.Cross experiments and analysis of the position of meiosis in Porphyra haitanensis (Rhodophyta)[J].Journal of Fisheries of China,2006,30(1):1-8.
[10] WANG J F,ZHU J Y,ZHOU W,et al.Early development patterns and morphogenesis of blades in four species of Porphyra (Bangiales,Rhodophyta)[J].Journal of Applied Phycology,2010,22(3):297-303.
[11] 吴宏肖,严兴洪,宋武林,等.坛紫菜与Pyropia radi种间杂交重组优良品系的选育与特性分析[J].水产学报,2014,38(8):1079-1088.WU H X,YAN X H,SONG W L,et al.Selection and characterization of an improved strain produced by genetic recombinant of interspecific hybridization between Pyropia haitanensis and Pyropia radi[J].Journal of Fisheries of China,2014,38(8):1079-1088.
[12] 刘美君,黄林彬,严兴洪.条斑紫菜种内杂交优良品系的筛选与特性分析[J].中国水产科学,2015,22(1):33-43.LIU M J,HUANG L B,YAN X H.Isolation and characterization of the improved strain HW-4 by in-traspecific hybridization in Pyropia yezoensis[J].Journal of Fishery Sciences of China,2015,22(1):33-43.
[13] DING H C,Lü F,WU H X,et al.Selection and characterization of an improved strain produced by inter-species hybridization between Pyropia sp.from India and Pyropia haitanensis from China[J].Aquaculture,2018,491:177-183.
[14] 李淑平,严兴洪.60Co-γ射线辐照对长紫菜的诱变效果及优良品系分离与特性分析[J].海洋学报,2015,37(10):69-79.LI S P,YAN X H.Isolation and characterization of an improved strain of Pyropia dentata (Bangiales,Rhodophyta) after being irradiated by 60Co-γ ray[J].Acta Oceanologica Sinica,2015,37(10):69-79.
[15] 戴继勋,张全启,包振民,等.紫菜原生质体的纯系培育、诱变处理和种间细胞融合的研究[J].海洋与湖沼,1990,21(3):293-296.DAI J X,ZHANG Q Q,BAO Z M,et al.Studies on the pure line culture,mutagenization and interspecific fusion of Porphyra protoplasts[J].Oceanologia et Limnologia Sinica,1990,21(3):293-296.
[16] 严兴洪.紫外线辐射与条斑紫菜原生质体后代发育和变异[J].上海水产大学学报,1992,1(1/2):71-78.YAN X H.Effects of UV irradiation on the development and variations of protoplast progenies in Porphyra yezoensis[J].Journal of Shanghai Fisheries University,1992,1(1/2):71-78.
[17] 刘宝.角蛋白降解菌的紫外线和硫酸二乙酯复合诱变[D].南京:南京农业大学,2012:14.LIU B.Complex mutagenesis of keratinose degrading strain with ultraviolet ray and diethyl sufate[D].Nanjing:Nanjing Agricultural University,2012:14.
[18] 何培民,秦松,严小军,等.海藻生物技术及其应用[M].北京:化学工业出版社,2007:59-61.HE P M,QIN S,YAN X J,et al.Seaweed biological technology and application[M].Beijing:Chemical Industry Press,2007:59-61.
[19] 严兴洪,李琳,陈俊华,等.坛紫菜的单性生殖与遗传纯系分离[J].高技术通讯,2007,17(2):205-210.YAN X H,LI L,CHEN J H,et al.Parthenogenesis and isolation of genetic pure strains in Porphyra haitanensis (Bangiales,Rhodophyta)[J].High Technology Letters,2007,17(2):205-210.
[20] 王素娟,张小平,徐志东,等.坛紫菜营养细胞和原生质体培养的研究I[J].海洋与湖沼,1986,17(3):217-221.WANG S J,ZHANG X P,XU Z D,et al.A study on the cultivation of the vegetative cells and protoplasts of P.yezoensis[J].Oceanologia et Limnologia Sinica,1986,17(3):217-221.
[21] KATO M,ARUGA Y.Comparative studies on the growth and photosynthesis of the pigmentation mutants of Porphyra yezoensis in laboratory culture[J].Japanese Journal of Phycology,1984,32:333-347.
[22] 赵东旭,杨新芳,刘亚丽,等.紫外线辐射对小麦幼苗生长的影响[J].河南科学,1996,14(s1):32-35.ZHAO D X,YANG X F,LIU Y L,et al.Effects of ultraviolet radiation on growth of wheat seedlings[J].Henan Sciences,1996,14(s1):32-35.
[23] 张敏.紫外线对霞多丽葡萄品质及相关生化物质的影响[D].晋中:山西农业大学,2016:16-30.ZHANG M.The affect of ultraviolet radiation to the quality and relevant substances of Chardonnay[D].Jinzhong:Shanxi Agricultural University,2016:16-30.
[24] 岳明,王勋陵.紫外线辐射对小麦和燕麦竞争性平衡的影响——小麦和燕麦生物量结构与冠层结构[J].环境科学学报,1999,19(5):526-531.YUE M,WANG Y L.Effect of enhanced ultraviolet radiation on competitive balance of wheat and oat-the structure of biomass and canopy[J].Acta Scientiae Circumstantiae,1999,19(5):526-531.
[25] 陈爱芹.紫外线诱变处理对马铃薯茎段再生植株生理生化变异的影响[D].长沙:湖南农业大学,2008:26-34.CHEN A Q.Physiological and biochemical variation of the potato stem section regeneration plants by ultraviolet radiation[D].Changsha:Hunan Agricultural University,2008:26-34.
[26] 李波,贾秀峰,焦德志,等.紫外线诱变苜蓿愈伤组织抗旱性的研究[J].干旱地区农业研究,2005,26(6):103-105.LI B,JIA X F,JIAO D Z,et al.Influence of ultraviolet ray to drought resistance of alfalfa calluses[J].Agricultural Research in the Arid Areas,2005,26(6):103-105.
[27] 黄世玉,王志勇.紫外线辐射坛紫菜离体细胞的初步试验[J].集美大学学报,1997,2(1):25-30.HUANG S Y,WANG Z Y.Preliminary experiment on isolated cells from Porphyra haitanensis (Rhodophyta) radiated by ultraviolet ray[J].Journal of Jimei University,1997,2(1):25-30.
[28] 严兴洪,张淑娟,黄林彬.60Co-γ射线对条斑紫菜(Porphyra yezoensis)的诱变效果与色素突变体分离[J].海洋与湖沼,2009,40(1):56-61.YAN X H,ZHANG S J,HUANG L B.Induction and isolation of pigmentation mutants of Porphyra yezoensis Ueda (Bangiales,Rhodophyta) by 60Co-γ ray irradiation[J].Oceanologia et Limnologia Sinica,2009,40(1):56-61.
[29] YAN X H,ARUGA Y.Induction of pigmentation mutants by treatment of monospore germlings with NNG in Porphyra yezoensis Ueda (Bangiales,Rhodophyta)[J].Algae,1997,12(1):39-52.
[30] YAN X H,FUJITA Y,ARUGA Y.Induction and characterization of pigmentation mutants in Porphyra yezoensis(Bangiales,Rhodophyta)[J].Journal of Applied Phycology,2000,12(1):69-81.
[31] 王素娟,马凌波,许璞,等.60Co-γ射线诱变条斑紫菜丝状体的研究[J].海洋科学,1999(4):43-47.WANG S J,MA L B,XU P,et al.Study on using gamma-ray to induce mutation in conchocelis of Porphyra yezoensis[J].Marine Sciences,1999(4):43-47.
[32] 李琳,严兴洪.坛紫菜绿色突变体的分离与特性分析[J].上海水产大学学报,2006,15(1):30-35.LI L,YAN X H.Isolation and characterization of a green type mutant in Porphyra haitanensis Chang et Zheng (Bangiales,Rhodophyta)[J].Journal of Shanghai Fisheries University,2006,15(1):30-35.
[33] 匡梅,许璞,王素娟.γ-射线对条斑紫菜和坛紫菜诱变作用的初步研究[J].上海水产大学学报,1997,6(4):241-245.KUANG M,XU P,WANG S J.A preliminary study of the mutagensis of γ-rays on Porphyra yezoensis and P.haitanensis[J].Journal of Shanghai Fisheries University,1997,6(4):241-245.
[34] 严兴洪,梁志强,宋武林,等.坛紫菜人工色素突变体的诱变与分离[J].水产学报,2005,29(2):166-172.YAN X H,LIANG Z Q,SONG W L,et al.Induction and isolation of artificial pigmentation mutants in Porphyra haitanensis Chang et Zheng (Bangiales,Rhodophyta)[J].Journal of Fisheries of China,2005,29(2):166-172.
[35] 严兴洪,田中次郎,有贺佑胜.条斑紫菜色彩突变体的诱导、分离和特性分析[J].水产学报,2000,24(3):221-228.YAN X H,TANAKA J,ARUGA Y.Isolation and characterization of pigmentation mutants in Porphyra yezoensis Ueda (Bangiales,Rhodophyta)[J].Journal of Fisheries of China,2000,24(3):221-228.