坛紫菜与长紫菜种间杂交后代的细胞崩溃现象与成活后代的表现型观察比较
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摘要
坛紫菜壳孢子萌发的第一次和第二次分裂为减数分裂,结束后形成了一个基因型不同的四分体,随后由这个四分体发育成一个基因型嵌合的叶状体。由坛紫菜与长紫菜种间杂交所产生的杂合丝状体成熟后释放出壳孢子,后者在分裂初期出现了大量的细胞崩溃死亡,通过对单个的壳孢子萌发体进行定点跟踪观察后发现,细胞崩溃死亡不仅会发生在减数分裂的直接产物即处于二细胞期、三细胞期、四细胞期的萌发体,当细胞数大于4个的萌发体在进行有丝分裂时也会出现细胞崩溃死亡。绝大部分的壳孢子能进行第一次减数分裂,形成处于二细胞期的萌发体,随后进行第二次减数分裂,形成处于三细胞期或四细胞期的萌发体。培养4 d,78.6%的壳孢子已发育成含2~4个细胞的萌发体,其余为未分裂的壳孢子萌发体;在含2~4个细胞的萌发体中未出现细胞崩溃死亡的萌发体的百分率为99.7%,0.3%的个体出现了细胞崩溃死亡。随着培养时间的延长,大多数处于二细胞期的萌发体发育成处于三细胞期和四细胞期的萌发体,而大规模的细胞崩溃死亡也出现在处于三细胞期和四细胞期的萌发体中。培养14 d,处于三细胞期和四细胞期的萌发体的百分率分别为38.6%和37.2%,它们当中含有崩溃死亡细胞的个体百分率分别为99.5%和99.2%。此外,8.9%的萌发体能够发育成含4个以上细胞的个体,但在随后的有丝分裂中也出现了细胞崩溃死亡现象。培养35 d,大部分的萌发体其体细胞已全部崩溃死亡,但仍有约1%的萌发体能成活下来,可它们并非是由一个完整的四分体发育而来,而是由四分体中的1~2个成活细胞发育而来。培养45 d,可把成活体的形态和颜色分成类亲本型和重组型两大类。研究表明,利用紫菜种间杂交的后代在壳孢子萌发初期会出现细胞崩溃死亡的现象,不仅可以用于辅助更精确地鉴别物种,而且可以从其成活后代中筛选出具有重组优势的新品系。
In Pyropia haitanensis, meiosis occurs during the first two divisions of germinating conchospores, and then the latter formed a tetrad with diffierent genotype, finally the tetrad developed into a genotype chimeric blade.The heterozygous conchocelies was formed in the interspecific hybridization between Pyropia haitanensis and Pyropia dentata, which could produce conchospores after maturation, but the conchospores had serious cell breakdown at the early development stage of germination. Through the tracking observation of single conchospore germling, we found that cell breakdown occurred not only in the direct products of meiosis(dyad, triad and tetrad),but also in the mitosis when the number of germling cell was more than 4. In culture, most of the conchospores could form dyad after the first meiotic division, and then form triad or tetrad through the second meiotic division.When cultured for 4 days, 78.6% of the conchospores had developed into germlings containing 2–4 cells, and the rest were undivided conchospores; The percentage of germlings containing 2–4 cells that did not have dead cells was 99.7%, and 0.3% of individuals had dead cells. With the prolongation of culture time, most of the dyads developed into the triads and tetrads, while large-scale cells breakdown appeared in the triad and tetrad. After culture for 14 days, the percentages of triad and tetrad were 38.6% and 37.2%, respectively, in which 99.5% and 99.2%contained dead cells, respectively. In addition, 8.9% of the germlings could develop into individuals with more than 4 cells, but cell death also appeared in the subsequent mitosis. When cultured for 35 days, most of the cells of germlings had all died, still, about 1% of the germlings survived, but they did not develop from the complete tetrads, just developed from 1-2 surviving tetrad cells. After 45 days of culture, the surviving individuals could be divided into parental and recombinational type based on their morphology and color. The results confirmed that the hybrid breakdown occured in the early development stage of conchospores in interspecific hybridization of Pyropia, which could not only be used to assist in the more accurate identification of species, but also we could select new strains with recombination advantage from the surviving offspring.
In Pyropia haitanensis, meiosis occurs during the first two divisions of germinating conchospores, and then the latter formed a tetrad with diffierent genotype, finally the tetrad developed into a genotype chimeric blade.The heterozygous conchocelies was formed in the interspecific hybridization between Pyropia haitanensis and Pyropia dentata, which could produce conchospores after maturation, but the conchospores had serious cell breakdown at the early development stage of germination. Through the tracking observation of single conchospore germling, we found that cell breakdown occurred not only in the direct products of meiosis(dyad, triad and tetrad),but also in the mitosis when the number of germling cell was more than 4. In culture, most of the conchospores could form dyad after the first meiotic division, and then form triad or tetrad through the second meiotic division.When cultured for 4 days, 78.6% of the conchospores had developed into germlings containing 2–4 cells, and the rest were undivided conchospores; The percentage of germlings containing 2–4 cells that did not have dead cells was 99.7%, and 0.3% of individuals had dead cells. With the prolongation of culture time, most of the dyads developed into the triads and tetrads, while large-scale cells breakdown appeared in the triad and tetrad. After culture for 14 days, the percentages of triad and tetrad were 38.6% and 37.2%, respectively, in which 99.5% and 99.2%contained dead cells, respectively. In addition, 8.9% of the germlings could develop into individuals with more than 4 cells, but cell death also appeared in the subsequent mitosis. When cultured for 35 days, most of the cells of germlings had all died, still, about 1% of the germlings survived, but they did not develop from the complete tetrads, just developed from 1-2 surviving tetrad cells. After 45 days of culture, the surviving individuals could be divided into parental and recombinational type based on their morphology and color. The results confirmed that the hybrid breakdown occured in the early development stage of conchospores in interspecific hybridization of Pyropia, which could not only be used to assist in the more accurate identification of species, but also we could select new strains with recombination advantage from the surviving offspring.
引文
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[2]吴航航,丁洪昌,严兴洪.条斑紫菜耐高温杂交重组品系的筛选与特性分析[J].水产学报,2017,41(5):711-722.Wu H H,Ding H C,Yan X H.Selection and characterization of a high-temperature resistant strain by hybridization recombination in Pyropia yezoensis[J].Journal of Fisheries of China,2017,41(5):711-722(in Chinese).
[3]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,2007:86-98.He P M,Qin S,Yan X J,et al.Seaweed biotechnology and its application[M].Beijing:Chemical Industry Press,2007:86-98(in Chinese).
[4]严兴洪,马少玉.坛紫菜抗高温品系的筛选[J].水产学报,2007,31(1):112-119.Yan X H,Ma S Y.Selection of a high-temperature resistant strain of Porphyra haitanensis(Rhodophyta)[J].Journal of Fisheries of China,2007,31(1):112-119(in Chinese).
[5]Yan X H,Lv F,Liu C J,et al.Selection and characterization of a high-temperature tolerant strain of Porphyra haitanensis Chang et Zheng(Bangiales,Rhodophyta)[J].Journal of Applied Phycology,2010,22(4):511-516.
[6]严兴洪,陈敏.坛紫菜耐低盐优良品系的筛选[J].上海水产大学学报,2008,17(3):316-320.Yan X H,Chen M.Selection of low-salinity resistant improved varieties in Porphyra haitanensis(Bangiales,Rhodophyta)[J].Journal of Shanghai Fisheries University,2008,17(3):316-320(in Chinese).
[7]徐燕,陈昌生,谢潮添,等.坛紫菜杂交品系优势的初步评价[J].海洋水产研究,2008,29(1):62-69.Xu Y,Chen C S,Xie C T,et al.Preliminary evaluation of hybridized strains of Porphyra haitanensis[J].Marine Fisheries Research,2008,29(1):62-69(in Chinese).
[8]纪德华,谢潮添,徐燕,等.坛紫菜品系间杂交子代杂种优势的ISSR分析[J].海洋学报,2008,30(6):147-153.Ji D H,Xie C T,Xu Y,et al.ISSR analysis on the heterosis in hybrids of Porphyra haitanensis[J].Acta Oceanologica Sinica,2008,30(6):147-153(in Chinese).
[9]刘海洋,李琳,严兴洪.坛紫菜与Pyropia radi的种间杂交实验[J].上海海洋大学学报,2013,22(6):882-887.Liu H Y,Li L,Yan X H.Study on the interspecies cross between Pyropia haitanensis and Pyropia radi[J].Journal of Shanghai Ocean University,2013,22(6):882-887(in Chinese).
[10]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(in Chinese).
[11]李淑平.长紫菜的人工诱变育种及与坛紫菜的种间杂交试验[D].上海:上海海洋大学,2015:33-37.Li S P.The artificial mutation breeding and the interspe-cific hybridization with Pyropia haitanensis of Pyropia dentata[D].Shanghai:Shanghai Ocean University,2015:33 -37(in Chinese).
[12]Zhang Y,Yan X H,Aruga Y.The sex and sex determination in Pyropia haitanensis(Bangiales,Rhodophyta)[J].PLo S One,2013,8(8):e73414.
[13]严兴洪,李琳,有贺祐胜.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(in Chinese).
[14]Shin J A,Miura A,Fu P F.Hybrid breakdown and breakthrough in interspecific crosses between Porphyra yezoensis and P.tenera[J].National History Research,1997,3:65-70.
[15]Niwa K,Kobiyama A,Sakamoto T.Interspecific hybridization in the haploid blade-forming marine crop Porphyra(Bangiales,Rhodophyta):Occurrence of allodiploidy in surviving F1 gametophytic blades[J].Journal of Phycology,2010,46(4):693-702.
[16]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(in Chinese).
[17]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.haitanensis I.[J].Oceanologia et Limnologia Sinica,1986,17(3):217-221(in Chinese).
[18]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,2006,29(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,2006,29(2):166-172(in Chinese).
[19]Kim N G.Culture study on the hybrid by interspecific crossing between Porphyra pseudolinearis and P.dentata(Bangiales,Rhodophyta),two dioecious speciesin culture[J].ALAGE,2011,16(1):79-86.
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