doi:

DOI: 10.3724/SP.J.1118.2019.19024

Journal of Fishery Sciences of China (中国水产科学) 2019/26:3 PP.592-603

Advances in Pyropia(formerly Porphyra) genetics and breeding


Abstract:
Pyropia (formerly known as Porphyra) is an important large red algae (laver) that not only tastes delicious but also possesses several health-beneficial effects, such as antilipidemic and anti-ageing effects. In addition, Pyropia blades have a strong ability to absorb nitrogen and phosphorus and fix carbon, and its cultivation also plays an important role in the restoration and improvement of eutrophication in shallow seas. Herein, we introduce the germplasm resources of Pyropia in China and review the advances in its breeding technique, objectives, and achievements. At present, there are 24 species and varieties of laver in China, and they are mainly distributed from the south of the Huanghai and Bohai Seas to the southeast coast of Guangdong, and a few species are also common in Taiwan and Hainan Island. In the Huanghai and Bohai Seas, P. yezoensis, P. tenera, and P. katadai are the main species, and in the southeast coast, P. haitanensis, P. crispata, and P. dentata are the main species. Based on the natural groups and habitat environment, there are two cultivation areas, namely, the north and south of the Yangtze River cultivating P. yezoensis and P. haitanensis, respectively. Although the problem of seed quantity has been solved by traditional seedling methods, the problem of seed quality has not been resolved because inbreeding could easily decrease the quality of a cultivated germplasm. Therefore, genetic breeding of Pyropia is of great significance to promote its yield and quality. The breeding techniques of Pyropia have been improved with the progress in science and technology and with the accumulation of cultivation experience, through the initial field selection (selective breeding), mutation breeding and cross breeding, which are based on cell biology, and genetic engineering breeding. The purpose of genetic breeding of Pyropia is to provide varieties that meet the needs of production and development; therefore, the breeding objectives should also change with changes in production and market demand. In the early stage of breeding, yield and quality were used as key indicators to identify whether a variety is good. With changes in climate and marine environment, it is of great significance to cultivate varieties of Pyropia with high-temperature and low-salt resistance. At present, Chinese has cultivated 6 new varities of Pyropia, and 4 of them are P. haitanensis, 2 of them are P. yezoensis. Finally, we put forward some problems in the development of laver industry in China, assuming that superior seed selection and industrial development are interrelated. Overall, the paper provide useful information for further studies in Pyropia genetics and breeding.

Key words:Pyropia (Porphyra) breeding; germplasm resource; breeding technique; breeding objective

ReleaseDate:2019-07-04 08:55:18



[1] Yoshida T, Notoya M, Kikuchi N, et al. Catalogue of species of Porphyra in the world, with special reference to the type locality and bibliography[J]. Natural History Research Special Issue, 1997, 3:5-18.

[2] Guo L, Wang S J, Hao Q, et al. Adbances in the biological functions of polysaccharide and phycoerythrin in Porphyra[J]. Food Research and Development, 2010, 31(6):182-185.[郭雷, 王淑军, 郝倩, 等. 紫菜多糖和藻红蛋白生物活性的研究进展[J]. 食品研究与开发, 2010, 31(6):182-185.]

[3] Zhou H P, Chen Q H. Antiageing effect of the polysaccharide from Porphyra yezoensis Ueda[J]. Journal of China Pharmaceutical University, 1989, 20(4):231-234.[周慧萍, 陈琼华. 紫菜多糖抗衰老作用的实验研究[J]. 中国药科大学学报, 1989, 20(4):231-234.]

[4] Zhou H P, Chen Q H. Anticoagulative and antilipemic effect of polysaccharide from Porphyra yezoensis Ueda[J]. Journal of China Pharmaceutical University, 1990, 21(6):358-360.[周慧萍, 陈琼华. 紫菜多糖的抗凝血和降血脂作用[J]. 中国药科大学学报, 1990, 21(6):358-360.]

[5] Zhao S F. Marine Algae and Algae Culture Science[M]. Beijing:National Defense Industry Press, 2012:282.[赵素芬. 海藻与海藻栽培学[M]. 北京:国防工业出版社, 2012:282.]

[6] Zhu J Y, Yan X H, Ding L P, et al. Color Atlas of Chinese Laver[M]. Beijing:China Agriculture Press, 2016:22.[朱建一, 严兴洪, 丁兰平, 等. 中国紫菜原色图集[M]. 北京:中国农业出版社, 2016:22.]

[7] Yang Y F, Fei X G. Prospects for bioremediation of cultivation of large-sized seaweed in Eutrophic mariculture areas[J]. Journal of Ocean University of Qingdao, 2003, 33(1):53-57.[杨宇峰, 费修绠. 大型海藻对富营养化海水养殖区生物修复的研究与展望[J]. 青岛海洋大学学报, 2003, 33(1):53-57.]

[8] Drew K M. Conchocelis phase in the life history of Porphyra umbilicalis (L.) Kütz[J]. Nature, 1949, 164:748-749.

[9] Kurogi M. Life history study of Porphyra[J]. Research Report of Northeast Sea Fisheries Research Institute, 1953(2):67-103.[黑木宗尚. アマノリ類の生活史の研究[J]. 東北海區水産研究所研究報告, 1953(2):67-103.]

[10] Tseng C K, Chang T J. Study of Porphyra I. The life history of Porphyra tenera[J]. Journal of Integrative Plant Biology, 1954, 3(3):287-302.[曾呈奎, 张德瑞. 紫菜的研究Ⅰ. 甘紫菜的生活史[J]. 植物学报, 1954, 3(3):287-302.]

[11] Zheng B F, Li J. Chinese Seaweed Flora (Volume Ⅱ, Book I):Porphyridiales, Erythropeltidales Garbary Hansen et Scagel, Goniotrichales Skuja, Bangiales[M]. Beijing:Science Press, 2009.[郑宝福, 李钧. 中国海藻志(第二卷, 第一册):紫球藻目, 红盾藻目, 角毛藻目, 红毛菜目[M]. 北京:科学出版社, 2009.]

[12] Kurogi M. Systematics of Porphyra in Japan[M]//Abbott I A, Kurogi M. Contributions to Systematics of Benthic Marine Algae of the North Pacific. Kobe:Japanses Society of Phycology, 1972.

[13] Tseng C K, Chang T J. On Porphyra margintat sp. nov. and its systematic position[J]. Journal of Integrative Plant Biology, 1958, 7(1):15-25.[曾呈奎, 张德瑞. 边紫菜及其系统学地位[J]. 植物学报, 1958, 7(1):15-25.]

[14] Fujian Fisheries Bureau. Artificial Cultivation of Porphyra haitanensis[M]. Fuzhou:Fujian People's Publishing House, 1979:1-101.[福建水产局. 坛紫菜人工养殖[M]. 福州:福建人民出版社, 1979:1-101.]

[15] Ma J H, Cai S Q. Cultivation and Process of Porphyra yezoensis[M]. Beijing:Science Press, 1996:1-58.[马家海, 蔡守清. 条斑紫菜的栽培与加工[M]. 北京:科学出版社, 1996:1-58.]

[16] Li S P, Yan X H. Isolation and characterization of Pyropia dentate (Bangiales, Rhodophyta) after being irradiated by 60Co-γ ray[J]. Haiyang Xuebao, 2015, 37(10):69-79.[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.]

[17] Zhang Q. The artificial mutation breeding of Pyropia crispata and its interspecific hybridization with Pyropia haitanensis[D]. Shanghai:Shanghai Ocean University, 2018.[张倩. 皱紫菜(Pyropia crispata)的人工诱变育种及其与坛紫菜的种间杂交试验[D]. 上海:上海海洋大学, 2018.]

[18] Zhao S. Mutagenic effects of short-wavelength ultraviolet rays (UV-C) on Pyropia suborbiculata Kjellman and isolation of mutants[D]. Shanghai:Shanghai Ocean University, 2018.[赵爽. 紫外线对圆紫菜的诱变效果与突变体的分离[D]. 上海:上海海洋大学, 2018.]

[19] 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.[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.]

[20] Chen S S, Ding H C, Yan X H. Isolation and characterization of an improved strain of Porphyra chauhanii (Bangiales, Rhodophyta) with high-temperature resistance[J]. Journal of Applied Phycology, 2016, 28(5):3031-3041.

[21] Zhang Y Y, Yan X H. India-endemic Pyropia radi (Bangiales, Rhodophyta):biology and life history[J]. Oceanologia et Limnologia Sinica, 2014, 45(1):52-59.[张岩岩, 严兴洪. 一种印度产紫菜(Pyropia radi)的生物学特性与部分生活史[J]. 海洋与湖沼, 2014, 45(1):52-59.]

[22] You S C. The work and operation of national laver germplasm bank[J]. China Fisheries, 2004(9):66-67.[尤抒忱. 国家级紫菜种质库工作与运行情况[J]. 中国水产, 2004(9):66-67.]

[23] Tseng C K. Biology of Economic Algae Germplasm Seedlings[M]. Jinan:Shandong Science and Technology Press, 1999:50-59.[曾呈奎. 经济海藻种质种苗生物学[M]. 济南:山东科学技术出版社, 1999:50-59.]

[24] Zhang Y J, Yang Y X, Wang Q Y, et al. Studies on the breeding and genetics of Porphyra yezoensis Ueda[J]. Transactions of Oceanology and Limnology, 1985(4):44-51.[张佑基, 杨以勋, 王清印, 等. 条斑紫菜(Porphyra yezoensis Ueda)遗传和育种的研究[J]. 海洋湖沼通报, 1985(4):44-51.]

[25] Luo Q J, Yang R, Lin S Z, et al. "Zhedong No. 1" of Pyropia haitanensis[J]. China Fisheries, 2015(11):57-59.[骆其君, 杨锐, 林少珍, 等. 坛紫菜"浙东1号"[J]. 中国水产, 2015(11):57-59.]

[26] He P M, Qin S, Yan X J, et al. Seaweed Biotechnology and Its Application[M]. Beijing:Chemical Industry Press, 2007:86-98.[何培民, 秦松, 严晓军, 等. 海藻生物技术及其应用[M]. 北京:化学工业出版社, 2007:86-98.]

[27] Lee H J, Choi J I, Kim G H. Characterization of Porphyra lucasii pigment mutant induced by gamma irradiation[J]. Phycological Research, 2018, 66(3):199-208.

[28] Niwa K, Hayashi Y, Abe T, et al. Induction and isolation of pigmentation mutants of Porphyra yezoensis (Bangiales, Rhodophyta) by heavy-ion beam irradiation[J]. Phycological Research, 2009, 57(3):194-202.

[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):39-54.

[30] 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, 2016, 15(1):30-35.[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.]

[31] Yan X, 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.

[32] Lee H J, Choi J I. Isolation and characterization of a high-growth-rate strain in Pyropia yezoensis induced by ethyl methane sulfonate[J]. Journal of Applied Phycology, 2018, 30(4):2513-2522.

[33] Wang J F, Xu P, Zhu J Y, et al. Studies on the pigment mutation in Porphyra (Bangiales, Rhodophyta)[J]. Marine Fisheries Research, 2007, 28(2):28-35.[王金锋, 许璞, 朱建一, 等. 紫菜属海藻色素突变的研究[J]. 海洋水产研究, 2007, 28(2):28-35.]

[34] Liang H. Plant Genetics and Breeding[M]. Guangzhou:Guangdong Higher Education Press, 2002:197-206.[梁红. 植物遗传与育种[M]. 广州:广东高等教育出版社, 2002:197-206.]

[35] Yan X H, Liang Z Q, Song W L, et al. Induction and isolation of artificial pigmentation mutants of Porphyra haitanensis Chang et Zheng (Bangiales, Rhodophyta)[J]. Journal of Fisheries of China, 2005, 29(2):166-172.[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.]

[36] 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.[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.]

[37] National Fisheries Technology Extension Center. Promotion of New Varieties of Aquatic Guide in 2009[M]. Beijing:China Agriculture Press, 2010:42-57.[全国水产技术推广总站. 2009水产新品种推广指南[M]. 北京:中国农业出版社, 2010:42-57.]

[38] National Fisheries Technology Extension Center. Promotion of New Varieties of Aquatic Guide in 2014[M]. Beijing:China Agriculture Press, 2014:165-181.[全国水产技术推广总站. 2014水产新品种推广指南[M]. 北京:中国农业出版社, 2014:165-181.]

[39] Lü F, Yan X H, Liu C J, et al. Selection of a high-temperature tolerant strain of Porphyra haitanensis and its cultivation in sea area[J]. Journal of Shanghai Ocean University, 2010, 19(4):457-462.[吕峰, 严兴洪, 刘长军, 等. 坛紫菜耐高温品系的选育与海区中试[J]. 上海海洋大学学报, 2010, 19(4):457-462.]

[40] 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.[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.]

[41] 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.[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.]

[42] 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.[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.]

[43] Ding H C, Zhang B L, Yan X H. Isolation and characterization of a heat-resistant strain with high yield of Pyropia yezoensis Ueda (Bangiales, Rhodophyta)[J]. Aquaculture and Fisheries, 2016, 1:24-33.

[44] 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.[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.]

[45] National Fisheries Technology Extension Center. Promotion of New Varieties of Aquatic Guide in 2014[M]. Beijing:China Agriculture Press, 2014:145-164.[全国水产技术推广总站. 2014水产新品种推广指南[M]. 北京:中国农业出版社, 2014:145-164.]

[46] National Fisheries Technology Extension Center. Promotion of New Varieties of Aquatic Guide in 2015[M]. Beijing:China Agriculture Press, 2015:1-150.[全国水产技术推广总站. 2015水产新品种推广指南[M]. 北京:中国农业出版社, 2015:1-150.]

[47] Suto S. Intergeneric and interspecific crossing of lavers (Porphyra)[J]. Nippon Suisan Gakkaishi, 1963, 29(8):739-748.

[48] Miura A, Shin J A. Crossbreeding in cultivars of Porphyra yezoensis (Bangiales, Rhodophyta)-Preliminary report[J]. Korean Journal of Phycology, 1989, 4(2):207-211.

[49] Chen C S, Xu Y, Ji D H, et al. A preliminary study on cultivation of the crossbred Porphyra haitanensis thallus and their economic characteristics[J]. Journal of Fisheries of China, 2007, 31(1):97-104.[陈昌生, 徐燕, 纪德华, 等. 坛紫菜品系间杂交藻体选育及经济性状的初步研究[J]. 水产学报, 2007, 31(1):97-104.]

[50] National Fisheries Technology Extension Center. Promotion of new Varieties of Aquatic Guide in 2012[M]. Beijing:China Agriculture Press, 2012:134-153.[全国水产技术推广总站. 2012水产新品种推广指南[M]. 北京:中国农业出版社, 2012:134-153.]

[51] Liu M J, Huang L B, Yan X H. Isolation and characterization of the improved strain HW-4 by intraspecific hybridization in Pyropia yezoensis[J]. Journal of Fishery Sciences of China, 2015, 22(1):33-43.[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.]

[52] Tang L C, Ding H C, Yan X H. Selection and characterization of the improved strain (LC-14) in Pyropia yezoensis (Bangiales, Rhodophyta)[J]. Journal of Fishery Sciences of China, 2017, 24(4):738-745.[唐隆晨, 丁洪昌, 严兴洪. 条斑紫菜优良品系(LC-14)的筛选与特性分析[J]. 中国水产科学, 2017, 24(4):738-745.]

[53] Jiang H, Ding H C, Yan X H. Selection and characterization of an improved strain (A-18) by hybridization recombinant in Pyropia yezoensis (Bangiales, Rhodophyta)[J]. Haiyang Xuebao, 2018, 40(2):95-103.[江灏, 丁洪昌, 严兴洪. 条斑紫菜杂交重组品系(A-18)的筛选与特性分析[J]. 海洋学报, 2018, 40(2):95-103.]

[54] 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.[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.]

[55] Kim N G. Culture study on the hybrid by interspecific crossing between Porphyra pseudolinearis and P. dentata (Bangiales, Rhodophyta), two dioecious species in culture[J]. Alage, 2011, 16(1):79-86.

[56] Shin J A. Crossing between Porphyra yezoensis and P. tenera[J]. Algae, 1999, 14(1):73-77.

[57] 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.

[58] Gu L Z, Ding H C, Yan X H. Phenomenon of cell breakdown and phenotypic observation on surviving blades in interspecific hybridization progeny between Pyropia haitanensis and Pyropia dentata[J]. Journal of Fisheries of China, 2018, 42(1):60-71.[谷灵芝, 丁洪昌, 严兴洪. 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[J]. 水产学报, 2018, 42(1):60-71.]

[59] Tang Y L. Isolation and cultivation of the vegetative cells and protoplasts of Porphyra suborbiculata Kjellm[J]. Journal of Shandong College of Oceanology, 1982, 12(4):37-50.[唐延林. 紫菜营养细胞和原生质体的分离和培养[J]. 山东海洋学院学报, 1982, 12(4):37-50.]

[60] Fujita Y, Migita S. Fusion of protoplasts from thalli of two different color types in Porphyra yezoensis Ueda and development of fusion products[J]. Japanese Journal of Phycology, 1987, 35(3):201-208.

[61] Araki T, Morishita T. Fusion of protoplasts from wild type Porphyra yezoensis and green type P. tenera thalli (Rhodophyta)[J]. Nippon Suisan Gakkaishi, 1990, 56(7):1161.

[62] 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.[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.]

[63] Chen C S. Electrofusion of protoplasts from Porphyra haitanensis and Porphyra yezoensis thalli (Rhodophyta)[J]. Chinese Journal of Biotechnology, 1992, 8(1):65-69.[陈昌生. 坛紫菜和条斑紫菜的原生质体电融合[J]. 生物工程学报, 1992, 8(1):65-69.]

[64] Chen C S. Protoplast fusion of Porphyra haitanensis and P. psudolinearis and its culture[J]. Journal of Oceanography in Taiwan Strait, 1993, 12(2):180-184.[陈昌生. 坛紫菜和拟线形紫菜的原生质体融合及培养[J]. 台湾海峡, 1993, 12(2):180-184.]

[65] Mizukami Y, Okauchi M, Kito H, et al. Culture and development of electrically fused protoplasts from red marine algae, Porphyra yezoensis and P. suborbiculata[J]. Aquaculture, 1995, 132(3-4):361-367.

[66] Yan X H. Intrageneric fusion of protoplasts from gametophytic blades and development of fusion products in Porphyra spp. (Rhodophyta)[J]. Journal of Shanghai Fisheries University, 2001, 10(2):109-117.[严兴洪. 紫菜种内原生质体的融合和融合体再生[J]. 上海水产大学学报, 2001, 10(2):109-117.]

[67] Moon D A, Goff L J. Molecular characterization of two large DNA plasmids in the red alga Porphyra pulchra[J]. Current Genetics, 1997, 32(2):132-138.

[68] Goff L J, Ashen J, Moon D. The evolution of parasites from their hosts:A case study in the parasitic red algae[J]. Evolution, 1997, 51(4):1068-1078.

[69] Kuang M, Wang S J, Li Y, et al. Transient expression of exogenous GUS gene in Porphyra yezoensis (Rhodophyta)[J]. Chinese Journal of Oceanology Limnology, 1998, 16(S1):56-61.

[70] Fukuda S, Mikami K, Uji T, et al. Factors influencing efficiency of transient gene expression in the red macrophyte Porphyra yezoensis[J]. Plant Science, 2008, 174(3):329-339.

[71] Hirata R, Uji T, Fukuda S, et al. Development of a nuclear transformation system with a codon-optimized selection marker and reporter genes in Pyropia yezoensis (Rhodophyta)[J]. Journal of Applied Phycology, 2014, 26(4):1863-1868.

[72] Uji T, Hirata R, Fukuda S, et al. A codon-optimized bacterial antibiotic gene used as selection marker for stable nuclear transformation in the marine red alga Pyropia yezoensis[J]. Marine Biotechnology, 2014, 16(3):251-255.

[73] Kong F N, Zhao H L, Liu W X, et al. Construction of plastid expression vector and development of genetic transformation system for the seaweed Pyropia yezoensis[J]. Marine Biotechnology, 2017, 19(2):147-156.

[74] Feng C, Ding H C, Yan X H. Transcriptomic profiling of Pyropia haitanensis blade in responding to low-salinity stress[J]. Journal of Fisheries of China, 2016, 40(12):1842-1849.[冯畅, 丁洪昌, 严兴洪. 低盐胁迫下坛紫菜叶状体的转录组分析[J]. 水产学报, 2016, 40(12):1842-1849.]

[75] Xu Y, Huang L, Ji D H, et al. Construction of a dense genetic linkage map and mapping quantitative trait loci for economic traits of a doubled haploid population of Pyropia haitanensis (Bangiales, Rhodophyta)[J]. BMC Plant Biology, 2015, 15:228.

[76] Yang H, Mao Y X, Kong F N, et al. Profiling of the transcriptome of Porphyra yezoensis with Solexa sequencing technology[J]. Chinese Science Bulletin, 2011, 56(20):2119-2130.

[77] Im S, Choi S, Hwang M S, et al. De novo assembly of transcriptome from the gametophyte of the marine red algae Pyropia seriata and identification of abiotic stress response genes[J]. Journal of Applied Phycology, 2015, 27(3):1343-1353.

[78] Sun P P, Mao Y X, Li G Y, et al. Comparative transcriptome profiling of Pyropia yezoensis (Ueda) M.S. Hwang & H.G. Choi in response to temperature stresses[J]. BMC Genomics, 2015, 16:463.

[79] Wang W L, Teng F, Lin Y H, et al. Transcriptomic study to understand thermal adaptation in a high temperature-tolerant strain of Pyropia haitanensis[J]. PLoS ONE, 2018, 13(4):e0195842.

[80] Wang W L, Lin Y H, Teng F, et al. Comparative transcriptome analysis between heat-tolerant and sensitive Pyropia haitanensis strains in response to high temperature stress[J]. Algal Research, 2018, 29:104-112.

[81] Chang J, Chen L D, Xu Y, et al. Selection of the internal control gene for expression analyses of Pyropia haitanensis under high light stress by quantitative real-time PCR[J]. Journal of Fisheries of China, 2017, 41(7):1064-1072.[昌晶, 陈陆丹, 徐燕, 等. 高光胁迫下坛紫菜定量PCR内参基因的筛选[J]. 水产学报, 2017, 41(7):1064-1072.]

[82] Mei G S, Ji D H, Li B, et al. Molecular cloning and expression analysis under high temperature stress of ribosomal protein S15a gene from Porphyra haitanensis[J]. Journal of Fisheries of China, 2012, 36(12):1826-1833.[梅高尚, 纪德华, 李兵, 等. 坛紫菜核糖体蛋白S15a基因的克隆及高温胁迫表达分析[J]. 水产学报, 2012, 36(12):1826-1833.]

[83] Dai Z Z, Li B, Xu Y, et al. Cloning and expression pattern analysis of two heat shock protein (Hsp90) genes from Pyropia haitanensis[J]. Journal of Fisheries of China, 2014, 38(3):340-349.[代真真, 李兵, 徐燕, 等. 坛紫菜两种Hsp90基因的克隆及表达特征分析[J]. 水产学报, 2014, 38(3):340-349.]

[84] Lin Y H, Wang W L, Xu Y, et al. Cloning and expression analysis of serine hydroxyl methyltransferase (SHMT) genes from Pyropia haitanensis[J]. Progress in Fishery Sciences, 2018, 39(5):122-129.[林颖辉, 王文磊, 徐燕, 等. 坛紫菜丝氨酸羟甲基转移酶基因的克隆及表达特征[J]. 渔业科学进展, 2018, 39(5):122-129.]

[85] Kong F N, Mao Y X, Yang H, et al. Cloning and characterization of the HLIP gene encoding high light-inducible protein from Porphyra yezoensis[J]. Journal of Applied Phycology, 2012, 24(4):685-692.

[86] Xiao H D, Chen C S, Xu Y, et al. Cloning and expression analysis of the chloroplast fructose-1,6-bisphosphatase gene from Pyropia haitanensis[J]. Acta Oceanological Sinica, 2014, 33(4):92-100.

[87] Ji D H, Li B, Xu Y, et al. Cloning and quantitative analysis of five heat shock protein 70 genes from Pyropia haitanensis[J]. Journal of Applied Phycology, 2015, 27(1):499-509.

[88] Chen C S, Dai Z Z, Xu Y, et al. Cloning, expression, and characterization of carbonic anhydrase genes from Pyropia haitanensis (Bangiales, Rhodophyta)[J]. Journal of Applied Phycology, 2016, 28(2):1403-1417.

[89] Im S, Lee H N, Jung H S, et al. Transcriptome-based identification of the desiccation response genes in marine red algae Pyropia tenera (Rhodophyta) and enhancement of abiotic stress tolerance by PtDRG2 in Chlamydomonas[J]. Marine Biotechnology, 2017, 19(3):232-245.

[90] Na Y, Lee H N, Wi J, et al. PtDRG1, a desiccation response gene from Pyropia tenera (Rhodophyta), exhibits chaperone function and enhances abiotic stress tolerance[J]. Marine Biotechnology, 2018, 20(5):584-593.

[91] Aruga Y. Color and the pigments of Porphyra yezoensis[J]. Iden, 1980, 34(9):8-13.

[92] Tan Y H, Huang L B, Yan X H. Selection and characterization of a low-salinity tolerant strain in Pyropia haitanensis (Bangiales, Rhodophyta)[J]. Oceanologia et Limnologia Sinica, 2014, 45(3):504-512.[檀应华, 黄林彬, 严兴洪. 坛紫菜耐低盐品系的选育与特性分析[J]. 海洋与湖沼, 2014, 45(3):504-512.]

[93] Chen C S, Ji D H, Xie C T, et al. Breeding selection and comparison of the economic traits on the low salinity resistant strains of Porphyra haitanensis[J]. Journal of Jimei University:Natural Science, 2009, 14(1):1-7.[陈昌生, 纪德华, 谢潮添, 等. 坛紫菜耐低盐品系的选育及经济性状的比较[J]. 集美大学学报:自然科学版, 2009, 14(1):1-7.]

[94] Monotilla W D, Notoya M. Morphological and physiological responses of Porphyra suborbiculata Kjellman (Bangiales, Rhodophyta) blades from five localities[J]. Botanica Marina, 2004, 47(4):323-334.

[95] 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.[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.]