西沙珊瑚疾病调查与扁枝滨珊瑚微生物群落结构分析
摘要
本文记录了2010年5月至10月及2011年4月至8月在西沙群岛海域开展珊瑚疾病调查的结果,发现普哥滨珊瑚(Porites pukoensis)、扁枝滨珊瑚(Porites andrewsi)、蔷薇珊瑚(Montipora spp.)、杯形珊瑚(Pocillopora spp.)、鹿角珊瑚(Acropora spp.)、菊花珊瑚(Goniastrea spp.)、澄黄滨珊瑚(Porites lutea)和滨珊瑚(Porites spp.)等共14种珊瑚主要出现了白化病、白斑病、黑化病、黄色炎症样病症、粉红颗粒状综合症等9种不同症状的疾病,目前该海区为珊瑚疾病的频发区。其中,普哥滨珊瑚的黄色炎症样病症和扁枝滨珊瑚的白化病最为常见。普哥滨珊瑚黄色炎症样病症主要出现在永兴岛附近海区,患病部位存在大量黄色脓样分泌物,患病部位水螅体生长正常、萎缩或缺失,有时骨骼部分缺失,患病部位面积一般为0.4～2.4cm2；该疾病由机械损伤或其它原因引起,因机械损伤引起的伤口一般在2-3个月可以恢复,其中小面积伤口的黄色物质可在10-20d内消失、伤口基本恢复正常,而其它原因产生的黄色物质需1-3个月才能消失,有的甚至在观察期内无明显变化。扁枝滨珊瑚的白化病出现在七连屿一带的扁枝滨珊瑚分布区,该病存在整枝完全白化、局部大面积白化、散布白色斑点三种情况,白化部位水螅体缺失。
从扁枝滨珊瑚提取微生物宏基因组DNA,通过采用PCR-DGGE技术及序列分析,结果表明,在64条序列中有34条为可培养细菌,30条为不可培养细菌,它们分别与以下种亲缘关系最近：Vibrio communis、Vibrio harveyi、Vibrio penaeicida、Vibrio parahaemolyticus、Vibrio sp.、Mucus bacterium、Vibrio fortisstrain、PseudoAlteromonas sp.、Uncultured bacterium、Uncultured Alteromonas sp.、Uncultured alpha proteobacterium,其中白化扁枝滨珊瑚的亲缘关系最近的优势种群为Vibrio harveyi、Vibrio penaeicida、Vibrio parahaemolyticus、Vibrio sp.、 Vibrio cambellii,经2216E培养基培养的珊瑚微生物亲缘关系最近的优势菌为Alteromonas sp.,经TCBS培养基培养的珊瑚微生物亲缘关系最近的优势菌为Pseudo Alteromonas sp.。本文为我国西沙海域珊瑚疾病的首次报道,可为今后开展西沙珊瑚疾病防治研究提供参考。
The main stony coral diseases in Xisha Archipelago were surveyed from May to October in2010and from April to August in2011. Nine diseases, including coral bleaching, white spot disease, coral black disease, yellow inflammatory like syndrome, and pink syndrome, were found in14species of stony coral, such as Porites pukoensis, Porites andrewsi, Montipora spp., Pocillopora spp., Acropora spp., Goniastrea spp., Porites lutea, Porites spp.. The yellow inflammatory-like disease of Porites pukoensis and bleaching in Porites andrewsi were the most popular diseases in this area. The yellow inflammatory-like disease of Porites pukoensis was mainly distributed in the sea near Yongxing Island. Some yellow pus-like secreta presented in its lesion. The areas of lesion were from0.4to2.4cm2and the polyps in the lesion were normally growing, curling or missing. This disease might caused by mechanical damage or other unknown reasons. The recure of the disease caused by mechanical damage would be happened in two to three months. Although it would be disappeared in10to20days while the wound was small, the yellow secreta would be disappeared after one to three months while the disease was caused by other reasons. Bleaching of Porties andrewsi appeared in the sea of Qilian Island. The symptoms of this disease include complete bleaching, part bleaching or white spot disease. The polyps in its lesion missed and the skeleton was very white.
Microbial DNA genome was abstracted from Porties andrewsi, and it was analyzed by PCR-DGGE and sequence analysis technology. The results showed that most of the isolates fell within the Vibrio core group, with the highest gene sequence similarity being towards bacteria such as:Vibrio communis, Vibrio harveyi, Vibrio penaeicida, Vibrio parahaemolyticus, Vibrio sp., Mucus bacterium, Vibrio fortisstrain, PseudoAlteromonas sp., Uncultured bacterium, Uncultured Alteromonas sp., Uncultured alpha proteobacterium. The predominating population mainly belong Vibrio harveyi, Vibrio penaeicida, Vibrio parahaemoly ticus, Vibrio sp., Vibrio cambellii, in bleaching of Porties andrewsi. The predominating population mainly belong to Alteromonas sp. with2216E agar. The predominating population mainly belong to PseudoAlteromonas sp. with TCBS agar. This paper reported coral diseases in Xisha Archipelago for the first time and would provide information and data for further study of coral disease in Xisha in future.
从扁枝滨珊瑚提取微生物宏基因组DNA,通过采用PCR-DGGE技术及序列分析,结果表明,在64条序列中有34条为可培养细菌,30条为不可培养细菌,它们分别与以下种亲缘关系最近：Vibrio communis、Vibrio harveyi、Vibrio penaeicida、Vibrio parahaemolyticus、Vibrio sp.、Mucus bacterium、Vibrio fortisstrain、PseudoAlteromonas sp.、Uncultured bacterium、Uncultured Alteromonas sp.、Uncultured alpha proteobacterium,其中白化扁枝滨珊瑚的亲缘关系最近的优势种群为Vibrio harveyi、Vibrio penaeicida、Vibrio parahaemolyticus、Vibrio sp.、 Vibrio cambellii,经2216E培养基培养的珊瑚微生物亲缘关系最近的优势菌为Alteromonas sp.,经TCBS培养基培养的珊瑚微生物亲缘关系最近的优势菌为Pseudo Alteromonas sp.。本文为我国西沙海域珊瑚疾病的首次报道,可为今后开展西沙珊瑚疾病防治研究提供参考。
The main stony coral diseases in Xisha Archipelago were surveyed from May to October in2010and from April to August in2011. Nine diseases, including coral bleaching, white spot disease, coral black disease, yellow inflammatory like syndrome, and pink syndrome, were found in14species of stony coral, such as Porites pukoensis, Porites andrewsi, Montipora spp., Pocillopora spp., Acropora spp., Goniastrea spp., Porites lutea, Porites spp.. The yellow inflammatory-like disease of Porites pukoensis and bleaching in Porites andrewsi were the most popular diseases in this area. The yellow inflammatory-like disease of Porites pukoensis was mainly distributed in the sea near Yongxing Island. Some yellow pus-like secreta presented in its lesion. The areas of lesion were from0.4to2.4cm2and the polyps in the lesion were normally growing, curling or missing. This disease might caused by mechanical damage or other unknown reasons. The recure of the disease caused by mechanical damage would be happened in two to three months. Although it would be disappeared in10to20days while the wound was small, the yellow secreta would be disappeared after one to three months while the disease was caused by other reasons. Bleaching of Porties andrewsi appeared in the sea of Qilian Island. The symptoms of this disease include complete bleaching, part bleaching or white spot disease. The polyps in its lesion missed and the skeleton was very white.
Microbial DNA genome was abstracted from Porties andrewsi, and it was analyzed by PCR-DGGE and sequence analysis technology. The results showed that most of the isolates fell within the Vibrio core group, with the highest gene sequence similarity being towards bacteria such as:Vibrio communis, Vibrio harveyi, Vibrio penaeicida, Vibrio parahaemolyticus, Vibrio sp., Mucus bacterium, Vibrio fortisstrain, PseudoAlteromonas sp., Uncultured bacterium, Uncultured Alteromonas sp., Uncultured alpha proteobacterium. The predominating population mainly belong Vibrio harveyi, Vibrio penaeicida, Vibrio parahaemoly ticus, Vibrio sp., Vibrio cambellii, in bleaching of Porties andrewsi. The predominating population mainly belong to Alteromonas sp. with2216E agar. The predominating population mainly belong to PseudoAlteromonas sp. with TCBS agar. This paper reported coral diseases in Xisha Archipelago for the first time and would provide information and data for further study of coral disease in Xisha in future.
引文
[1]于登攀,邹仁林.鹿回头造礁珊瑚群落多样性的现状与动态.生态学报,1996,16(6)：259-264.
[2]王丽荣,赵焕庭.珊瑚礁生态系的一般特点.生态学杂志,2001,20(6)：41-45.
[3]牛文涛,刘玉新,林荣澄.珊瑚礁生态系统健康评价方法的研究进展.海洋学研究,2009,27(4)：77-85.
[4]朱葆华,王广策,黄勃等.温度、缺氧、氨氮和硝氮对3种珊瑚白化的影响.科学通报,2004,49(17)：1743-1748.
[5]关飞.世界珊瑚礁生存报告.沿海环境,2001,(11)：12-13.
[6]李淑,余克服,陈天然等.珊瑚共生虫黄藻密度结合卫星遥感分析2007年南沙群岛珊瑚热白化.科学通报,2011,56(10)：756-764.
[7]李淑,余克服,施祺等.造礁石珊瑚对低温的耐受能力及响应模式.应用生态学报,2009,20：2289-2295.
[8]李淑,余克服,施祺等.海南岛鹿回头石珊瑚对高温响应行为的实验研究.热带地理,2008,28：534-539.
[9]李淑,余克服.珊瑚礁白化研究进展.生态学报,2007,27(5)：2059-2069.
[10]李颖虹,黄小平,岳维忠.西沙永兴岛环境质量状况及管理对策.海洋环境科学,2004,23(1)：51-53.
[11]李颖虹,黄小平,岳维忠等.西沙群岛永兴岛珊瑚礁和礁坪生物生态学研究.海洋与湖沼,2004,35(2)：176-182.
[12]时小军,刘元兵,陈特固,余克服.全球气候变暖对西沙、南沙海域珊瑚生长的潜在威胁.热带地理,2008,28,342-368.
[13]吴钟解,王道儒,涂志刚等.西沙生态监控区造礁石珊瑚退化原因分析.海洋学报,2011,33(4)：140-146.
[14]邹仁林,陈友璋.我国浅水造礁石珊瑚地理分布的初步研究.南海海洋科学集刊,1983,4：89-95.
[15]邹仁林等.海南岛珊瑚垂直分带的初步研究.海洋与湖沼,1966,2(8)：153-160.
[16]邹仁林.中国动物志：造礁石珊瑚.北京：的科学出版社,2001：1-284.
[17]国家海洋局.中华人民共和国海洋行业标准——珊瑚礁生态监测技术规程.北京：中国标准出版社,2005,1-20.
[18]罗鹏,胡超群,谢珍玉等.凡纳滨对虾咸淡水养殖系统内细菌群落组成的PCR-DGGE分析.热带海洋学报,2006,25(2)：49-53.
[19]赵焕庭,袁家义.雷州半岛灯楼角珊瑚岸礁的特征.海洋地质与第四纪地质, 2002,22(2)：35-40.
[20]赵焕庭.中国现代珊瑚礁研究.世界科技研究与发展,1998,20(4)：98-105.
[21]聂宝符,梁美桃,朱袁智等.南海礁区现代造礁珊瑚类骨骼细结构的研究.北京：中国科学技术出版社,1991,1-78.
[22]莎姆布鲁克J.[美]著,黄培堂译《分子克隆实验指南》(第三版).科学出版社,2005,1,26-32.
[23]Aeby G, WORK T, FENNER D et al,2008. Coral and crutose coralline algae disease on the reefs of American Samoa. Proceeding of the 11th International Coral Reef Symposium,3:197-201.
[24]Aeby G S, ROSS M, WILLIAMS G J et al,2010. Disease dynamics of Montipora white syndrome within Kaneohe Bay, Oahu, Hawaii:distribution, seasonality, virulence, and transmissibility. Diseases of aquatic organisms DisAquat Org 91:1-8.
[25]Antonius A. New observations on coral destruction in reefs. Tenth Meeting of the Association of Island Marine Laboratories of the Caribbean (abstract). University of Puerto Rico (Mayaguez),1973,3.
[26]Antonius A. The "band" disease in coral reefs. Proceedings of the 4th International Coral Reef Symposium, Manila,1981,2,pp 7-14.
[27]Arboleda M D, Reichardt W T. Vibrio sp. causing Porites ulcerative white spot disease. Diseases of Aquatic Organisms Dia Aquat Org, (2010) 90,93-104.
[28]Baker A C, Starger C J, McClanahan T R, et al. Corals'adaptive response to climate change. Nature,2004,430:741.
[29]Banin E, Ben-Haim Y, Fine M et al,2000. Virulence mechanisms of the coral bleaching pathogen Vibrio shiloi. Proceedings 9th International Coral Reef Symposium,2.
[30]Ben-Haim Y, Rosenberg E. A novel Vibrio sp.pathogen of the coral Pocillipora damicornis. Marine Biology,2002,141(1):47-55.
[31]Brown B E,1997. Coral bleaching:causes and consequences. Coral Reefs,16, Suppl.:S129-S138.
[32]Buddemeier R W, Baker A C, Fautin D G, et al. The adaptive bleaching hypothesis.2004, p427-444.In:Rosenberg E & Loya Y(eds), Coral Health and Disease. Berlin:Springer.
[33]Bythell J, Pantos O, Richardson L. White Plague, white band, and other"white"diseases.In:Rosenberg E & Loya Y.(eds.) Coral Health and Disease. New York:Springer,2004:351-365.
[34]Chimettoa L A, Brocchia M, Thompsond F L et al. Vibrios dominate as culturable nitrogen-fixing bacteria of the Brazilian coral Mussismilia hispida. Systematic and Microbiology, (2008) 31,312-319.
[35]Connell J H. Diversity in tropical rain forests and coral reefs:High diversity of trees and corals ismaintained only in a nonequilibrium state. Science,1978,199: 1320-1370.
[36]Dustan P. Vitality of reef coral populations off Key Largo, Florida:Recruitment and mortality. Env. Geol.1977,2:51-58.
[37]Efrony R, Loya Y, Bacharach E Rosenberg. Phage therapy of coral disease. Coral Reef,2007,26:7-13.
[38]Efrony R, Atad I, Rosenberg E. Phage Therapy of Coral White Plague Disease: Properties of Phage BA 3.Curr Microbiol,2009,58:139-145.
[39]Gladfelter W B. White-band disease in Acropora palmata:implications for the structure and growth of shallow reefs. Bull Mar Sci,1982,32:639-643.
[40]Green E P, Bruckner A W. The significance of coral disease epizootiology for coral reef conservation. Biological Conservation,2000,96:347-361.
[41]Handelsman J, Rondon M R, Brady S F, et al. Molecular biological access to the chemistry of unknown soil microbes:a new frontier for natural products.Chem Biol,1998,5:R245-R249.
[42]Harvell, Drew, Jordan-Dahlgren et al. Coral disease, environmental drivers, and the balance between coral and microbial associates. Oceanography,2007,20(1): 172-195.
[43]Huang H, Zou R L, Liu S, et al. Impacts of warm effluent from the Daya Bay nuclear power plant on hermatypic coral community (I). The North Pacific Marine Science Organization 11th Annual Meeting,2002.
[44]Hughes T P, Baird A H, Bellwood D R et al. Climate Change,Human Impacts, and the Resilience of Coral Reefs. Science,2003,301:929-933.
[45]Jameson S C, Erdmann M V, Berger W H, et al. Development of biological criteria for coral reef ecosystem assessment. USEPA.1998. Office of Science and Technology, Health and Ecological Criteria Division, Washington, DC.
[46]Kimura T, Dai C F, Huang H, Status of coral reefs in east and north Asia(China, Korea and Japan). In:Proceedings of the 10th International Coral Reef Symposium. Japan, Japan Coral Reef Soc,2004,62.
[47]Kushmaro A, Loya Y, Fine M et al. Bacterial infection and coral bleaching. Science,1996,380:396.
[48]Kushmaro A, Rosenberg E, Fine M et al. Bleaching in the coral Oculina patagononica by Vibrio AK-1[J].Mar Ecol Prog Ser,1997,147:159-165.
[49]Kushmaro A, Banin E, Loya Y et al. Vibrio shiloi sp. nov., the causative agent of bleaching of the coral Oculina patagonica. International Journal of Systematic and Evolutionary Microbiology,2001,51,1383-1388.
[50]Lewis J B. Coral Reef Ecosystem. In:Longhurst A R(ed.). Analysis of Marine,1981, p127-158. Ecosystems. London. Now York. Torktoy, Sydey San Francisco. Academic Press.
[51]Littman R A, Willis B L, Bourne D G. Bacterial communities of juvenile corals infected with different Symbiodinium (dinoflagellate) clades. Mar Ecol Prog Ser 2009,389:45-59.
[52]Lindahl U. Coral reef rehabilitation through transplantation of staghorn corals: effects of artificialstabilization and mechanical damages. Coral Reefs,2003,22 (3):217-223.
[53]Loya Y. The coral reefs of Eilat-past, present and future:three decades of coral community structure studies.2004, p1-34. In:Rosenberg E & Loya Y(eds), Coral Health and Disease. Berlin:Springer.
[54]Loya Y. Community structure and species diversity of hermatypic corals at Eilat, Red sea. Mar Bio,1972,13:100-123.
[55]Loya Y. Possible effects of water pollution on the community structure of Red sea corals. Mar Bio,1975,29:177-185.
[56]Luna G M, Bongiorni L, Claudia Gili, Biavasco F, Danovaro R. Vibiro harveyi as a causative agent of the White Syndrome in tropical stony corals. Environment Microbiology Reports (2010) 2(1),120-127.
[57]Mumby P J, Dahlgren C P, Harborne A R et al. Fishing, Trophic Cascades, and the Process of Grazing on Coral Reefs. Science,2006,311,98-101.
[58]Omry Koren, Eugene Rosenberg, Bacteria Associated with the Bleached and Cave Coral Oculina patagonica. Microb Ecol,2008,55:523-539.
[59]Pantos O, Bythell J C. Bacterial community structure associated with white band disease in the elkhorn coral Acropora palmate determined using culture-independent 16S rRNA techniques. Diseases of Aquatic Organisms Dis Aquat Org.2006,69:79-88.
[60]Patterson K L, Porter J W, Ritchie K B,et al. The etiology of white pox, a lethal disease of the Caribbean elkhorn coral, Acropora palmata. Proceedings of the National Academy of Science, USA,2002,99(13):8725-8730.
[61]Ravindran J, Raghukumar C, Raghukumar S. Fungi in Porites lutea:association with healthy and diseased corals. Dis Aquat Organ,2001,47:219-228.
[62]Ravindran J, Raghukumar C. Pink line syndrome (PLS) in the scleractinian coral Porites lutea. Coral Reefs,2002,21:252.
[63]Ravindran J, Raghukumar C. Pink-line syndrome, a physiological crisis in the scleractinian coral Porites lutea. Marine Biology,2006,149:347-356
[64]Raymundo L H, Harvell C D, Reynolds T L. Porites ulcerative white spot disease: description, prevalence, and host range of a new coral disease affecting Indo-Pacific reefs. Diseases of Aquatic Organisms Dis Aquat Org,2003, 56:95-104.
[65]Raymundo L J, Rosell K B, Reboton C T, Kaczmarsky L. Coral disease onPhilippine reefs:genus Porites is a dominant host. Diseases of Aquatic Organisms Dis Aquat Org,2005,64:181-191.
[66]Reshef L, Koren O, Loya Y, Zilber-Rosenberg I, Rosenberg E. The coral probiotic hypothesis. Environ Microbiol,2006,8(12):2068-2073.
[67]Richard B. Aronson, William F. Precht. White-band disease and the changing face of Caribbean coral reefs. Hydrobiologia,2001,460:25-38.
[68]Richardson L L, Goldberg W M, Carlton R G et al. Coral disease outbreak in the Florida Keys:Plague Typer Ⅱ.Rev. Biol. Trop.,1998,46(5):187-198.
[69]Richardson L L, Smith G W, Richie K B, Carlton R G. Integrating microbiologica, microbiologica, microsensor, molecular, and physiologic techniques in the study of coral disease pathogenesis. Hydrobiologia,2001,460: 71-89.
[70]Richardson L L. Red band disease:a new cyanobacterial infestation of corals. Proceedings of the American Academy of Underwater Sciences Scientific Diving Symposium,1992,pp.153-160.
[71]Richardson L L. Coral diseases:what is really known? Trends in Ecology and Consveration.1998,13,438-443.
[72]Richardson L L. Coral diseases:What is really known? Trends Ecol Evol, 1998,13:438-443.
[73]Ritchie K B, Smith G W. Type II white-band disease. Revista de Biological Tropical,1998,46:199-203.
[74]Rogers C. Coral bleaching and disease should not be underestimated as causes of Caribbean coral reef decline. Proc Biol Sci,2009,276(1655):197-200.
[75]Rohwer F, Breitbart M, Jara J, et al. Diversity of bacteria associated with the Caribbean coral Montastrea franksi. Coral Reefs,2001,20:85-91.
[76]Rosenberg E and Ben-Haim Y. Microbial diseases of corals and global warming. Environ. Microbiol.2002,4:318-326.
[77]Rosenberg E, Barash Y. Microbial Diseases of Corals. In:Belkin and Colwell. Oceans and Health:Pathogens in the Marine Environment[M]. New York: Springer,2005,415-430.
[78]Rosenberg E, Falkovitz L. The Vibrio shiloi/Oculina patagonica model system of coral bleaching. Annu Rev Microbiol,2004,58:143-159.
[79]Smith G W, Well E. Coral reef diseases revised. In:Proceedings of the 10th International Coral Reef Symposium. Japan:Japanese Coral Reef Society,2004, 165.
[80]Stimson J, Sakai K, Sembali H. Interspecific comparison of the symbiotic relationship in corals with high and low rates of bleaching-induced mortality. Coral Reefs,2002,21:409-421.
[81]Sutherland K P, Porter J W, Torres C. Disease and immunity in Caribbean and Indo-Pacific zooxanthellate corals. Marine Ecology Progress Series Mar Ecol Prog Ser,2004,266:273-302.
[82]Vallaeys T, Topp E, Muyzer G, et al. Evaluation of denaturing gradient gel electrophoresis in the detection of 16S rDNA sequence variation in rhizobia and methanotrophs. FEMS Microbiology Ecology,1997,24:279-285.
[83]Vargas-Angel B. Coral health and disease assessment in the U.S. pacific remote island areas. Bulletin of Marine Science,2009,84(2):211-227.
[84]Venter J C, Remington K, Heidelberg J F,et al. Environmental genome shotgun sequencing of the Sargasso Sea.Science,2004,304:66-74.
[85]Veron J E N, Stafford-Smith M. Corals of the world. Australia:Austrlian Institute of Marine Science,2000,1(62-167),2(24-45),3(156-175,276-345).
[86]Wilkinson C. Status of Coral Reefs of the World:2004. Town sville, Queens land: Australian Institute of Marine Science Press,2004. (1):1-316.
[87]Wilkinson C. Status of coral reefs of the world:2000. Global Coral Reef Monitoring Network,Australian Institute of Marine Science, Cape Ferguson, Queensland,2000.
[88]Williams G J, Aeby G S, Cowie R O M et al. Predictive Modeling of Coral Disease Distribution within a Reef System. PLoS ONE,2010,5(2):e9264.
[89]Williams G J, Work T M, Aeby G S et al. Gross and microscopic morphology of lesions in Cnidaria from Palmyra Atoll, Central Pacific. Journal of Invertebrate Pathology,2011,106:165-173.
[90]Work T M, Richardson L L, Reynolds T L, Willis B L. Biomedical and veterinary science can increase our understanding of coral disease. Joural of Experimental Marine Biology and Ecology.2008,362:63-70.
[2]王丽荣,赵焕庭.珊瑚礁生态系的一般特点.生态学杂志,2001,20(6)：41-45.
[3]牛文涛,刘玉新,林荣澄.珊瑚礁生态系统健康评价方法的研究进展.海洋学研究,2009,27(4)：77-85.
[4]朱葆华,王广策,黄勃等.温度、缺氧、氨氮和硝氮对3种珊瑚白化的影响.科学通报,2004,49(17)：1743-1748.
[5]关飞.世界珊瑚礁生存报告.沿海环境,2001,(11)：12-13.
[6]李淑,余克服,陈天然等.珊瑚共生虫黄藻密度结合卫星遥感分析2007年南沙群岛珊瑚热白化.科学通报,2011,56(10)：756-764.
[7]李淑,余克服,施祺等.造礁石珊瑚对低温的耐受能力及响应模式.应用生态学报,2009,20：2289-2295.
[8]李淑,余克服,施祺等.海南岛鹿回头石珊瑚对高温响应行为的实验研究.热带地理,2008,28：534-539.
[9]李淑,余克服.珊瑚礁白化研究进展.生态学报,2007,27(5)：2059-2069.
[10]李颖虹,黄小平,岳维忠.西沙永兴岛环境质量状况及管理对策.海洋环境科学,2004,23(1)：51-53.
[11]李颖虹,黄小平,岳维忠等.西沙群岛永兴岛珊瑚礁和礁坪生物生态学研究.海洋与湖沼,2004,35(2)：176-182.
[12]时小军,刘元兵,陈特固,余克服.全球气候变暖对西沙、南沙海域珊瑚生长的潜在威胁.热带地理,2008,28,342-368.
[13]吴钟解,王道儒,涂志刚等.西沙生态监控区造礁石珊瑚退化原因分析.海洋学报,2011,33(4)：140-146.
[14]邹仁林,陈友璋.我国浅水造礁石珊瑚地理分布的初步研究.南海海洋科学集刊,1983,4：89-95.
[15]邹仁林等.海南岛珊瑚垂直分带的初步研究.海洋与湖沼,1966,2(8)：153-160.
[16]邹仁林.中国动物志：造礁石珊瑚.北京：的科学出版社,2001：1-284.
[17]国家海洋局.中华人民共和国海洋行业标准——珊瑚礁生态监测技术规程.北京：中国标准出版社,2005,1-20.
[18]罗鹏,胡超群,谢珍玉等.凡纳滨对虾咸淡水养殖系统内细菌群落组成的PCR-DGGE分析.热带海洋学报,2006,25(2)：49-53.
[19]赵焕庭,袁家义.雷州半岛灯楼角珊瑚岸礁的特征.海洋地质与第四纪地质, 2002,22(2)：35-40.
[20]赵焕庭.中国现代珊瑚礁研究.世界科技研究与发展,1998,20(4)：98-105.
[21]聂宝符,梁美桃,朱袁智等.南海礁区现代造礁珊瑚类骨骼细结构的研究.北京：中国科学技术出版社,1991,1-78.
[22]莎姆布鲁克J.[美]著,黄培堂译《分子克隆实验指南》(第三版).科学出版社,2005,1,26-32.
[23]Aeby G, WORK T, FENNER D et al,2008. Coral and crutose coralline algae disease on the reefs of American Samoa. Proceeding of the 11th International Coral Reef Symposium,3:197-201.
[24]Aeby G S, ROSS M, WILLIAMS G J et al,2010. Disease dynamics of Montipora white syndrome within Kaneohe Bay, Oahu, Hawaii:distribution, seasonality, virulence, and transmissibility. Diseases of aquatic organisms DisAquat Org 91:1-8.
[25]Antonius A. New observations on coral destruction in reefs. Tenth Meeting of the Association of Island Marine Laboratories of the Caribbean (abstract). University of Puerto Rico (Mayaguez),1973,3.
[26]Antonius A. The "band" disease in coral reefs. Proceedings of the 4th International Coral Reef Symposium, Manila,1981,2,pp 7-14.
[27]Arboleda M D, Reichardt W T. Vibrio sp. causing Porites ulcerative white spot disease. Diseases of Aquatic Organisms Dia Aquat Org, (2010) 90,93-104.
[28]Baker A C, Starger C J, McClanahan T R, et al. Corals'adaptive response to climate change. Nature,2004,430:741.
[29]Banin E, Ben-Haim Y, Fine M et al,2000. Virulence mechanisms of the coral bleaching pathogen Vibrio shiloi. Proceedings 9th International Coral Reef Symposium,2.
[30]Ben-Haim Y, Rosenberg E. A novel Vibrio sp.pathogen of the coral Pocillipora damicornis. Marine Biology,2002,141(1):47-55.
[31]Brown B E,1997. Coral bleaching:causes and consequences. Coral Reefs,16, Suppl.:S129-S138.
[32]Buddemeier R W, Baker A C, Fautin D G, et al. The adaptive bleaching hypothesis.2004, p427-444.In:Rosenberg E & Loya Y(eds), Coral Health and Disease. Berlin:Springer.
[33]Bythell J, Pantos O, Richardson L. White Plague, white band, and other"white"diseases.In:Rosenberg E & Loya Y.(eds.) Coral Health and Disease. New York:Springer,2004:351-365.
[34]Chimettoa L A, Brocchia M, Thompsond F L et al. Vibrios dominate as culturable nitrogen-fixing bacteria of the Brazilian coral Mussismilia hispida. Systematic and Microbiology, (2008) 31,312-319.
[35]Connell J H. Diversity in tropical rain forests and coral reefs:High diversity of trees and corals ismaintained only in a nonequilibrium state. Science,1978,199: 1320-1370.
[36]Dustan P. Vitality of reef coral populations off Key Largo, Florida:Recruitment and mortality. Env. Geol.1977,2:51-58.
[37]Efrony R, Loya Y, Bacharach E Rosenberg. Phage therapy of coral disease. Coral Reef,2007,26:7-13.
[38]Efrony R, Atad I, Rosenberg E. Phage Therapy of Coral White Plague Disease: Properties of Phage BA 3.Curr Microbiol,2009,58:139-145.
[39]Gladfelter W B. White-band disease in Acropora palmata:implications for the structure and growth of shallow reefs. Bull Mar Sci,1982,32:639-643.
[40]Green E P, Bruckner A W. The significance of coral disease epizootiology for coral reef conservation. Biological Conservation,2000,96:347-361.
[41]Handelsman J, Rondon M R, Brady S F, et al. Molecular biological access to the chemistry of unknown soil microbes:a new frontier for natural products.Chem Biol,1998,5:R245-R249.
[42]Harvell, Drew, Jordan-Dahlgren et al. Coral disease, environmental drivers, and the balance between coral and microbial associates. Oceanography,2007,20(1): 172-195.
[43]Huang H, Zou R L, Liu S, et al. Impacts of warm effluent from the Daya Bay nuclear power plant on hermatypic coral community (I). The North Pacific Marine Science Organization 11th Annual Meeting,2002.
[44]Hughes T P, Baird A H, Bellwood D R et al. Climate Change,Human Impacts, and the Resilience of Coral Reefs. Science,2003,301:929-933.
[45]Jameson S C, Erdmann M V, Berger W H, et al. Development of biological criteria for coral reef ecosystem assessment. USEPA.1998. Office of Science and Technology, Health and Ecological Criteria Division, Washington, DC.
[46]Kimura T, Dai C F, Huang H, Status of coral reefs in east and north Asia(China, Korea and Japan). In:Proceedings of the 10th International Coral Reef Symposium. Japan, Japan Coral Reef Soc,2004,62.
[47]Kushmaro A, Loya Y, Fine M et al. Bacterial infection and coral bleaching. Science,1996,380:396.
[48]Kushmaro A, Rosenberg E, Fine M et al. Bleaching in the coral Oculina patagononica by Vibrio AK-1[J].Mar Ecol Prog Ser,1997,147:159-165.
[49]Kushmaro A, Banin E, Loya Y et al. Vibrio shiloi sp. nov., the causative agent of bleaching of the coral Oculina patagonica. International Journal of Systematic and Evolutionary Microbiology,2001,51,1383-1388.
[50]Lewis J B. Coral Reef Ecosystem. In:Longhurst A R(ed.). Analysis of Marine,1981, p127-158. Ecosystems. London. Now York. Torktoy, Sydey San Francisco. Academic Press.
[51]Littman R A, Willis B L, Bourne D G. Bacterial communities of juvenile corals infected with different Symbiodinium (dinoflagellate) clades. Mar Ecol Prog Ser 2009,389:45-59.
[52]Lindahl U. Coral reef rehabilitation through transplantation of staghorn corals: effects of artificialstabilization and mechanical damages. Coral Reefs,2003,22 (3):217-223.
[53]Loya Y. The coral reefs of Eilat-past, present and future:three decades of coral community structure studies.2004, p1-34. In:Rosenberg E & Loya Y(eds), Coral Health and Disease. Berlin:Springer.
[54]Loya Y. Community structure and species diversity of hermatypic corals at Eilat, Red sea. Mar Bio,1972,13:100-123.
[55]Loya Y. Possible effects of water pollution on the community structure of Red sea corals. Mar Bio,1975,29:177-185.
[56]Luna G M, Bongiorni L, Claudia Gili, Biavasco F, Danovaro R. Vibiro harveyi as a causative agent of the White Syndrome in tropical stony corals. Environment Microbiology Reports (2010) 2(1),120-127.
[57]Mumby P J, Dahlgren C P, Harborne A R et al. Fishing, Trophic Cascades, and the Process of Grazing on Coral Reefs. Science,2006,311,98-101.
[58]Omry Koren, Eugene Rosenberg, Bacteria Associated with the Bleached and Cave Coral Oculina patagonica. Microb Ecol,2008,55:523-539.
[59]Pantos O, Bythell J C. Bacterial community structure associated with white band disease in the elkhorn coral Acropora palmate determined using culture-independent 16S rRNA techniques. Diseases of Aquatic Organisms Dis Aquat Org.2006,69:79-88.
[60]Patterson K L, Porter J W, Ritchie K B,et al. The etiology of white pox, a lethal disease of the Caribbean elkhorn coral, Acropora palmata. Proceedings of the National Academy of Science, USA,2002,99(13):8725-8730.
[61]Ravindran J, Raghukumar C, Raghukumar S. Fungi in Porites lutea:association with healthy and diseased corals. Dis Aquat Organ,2001,47:219-228.
[62]Ravindran J, Raghukumar C. Pink line syndrome (PLS) in the scleractinian coral Porites lutea. Coral Reefs,2002,21:252.
[63]Ravindran J, Raghukumar C. Pink-line syndrome, a physiological crisis in the scleractinian coral Porites lutea. Marine Biology,2006,149:347-356
[64]Raymundo L H, Harvell C D, Reynolds T L. Porites ulcerative white spot disease: description, prevalence, and host range of a new coral disease affecting Indo-Pacific reefs. Diseases of Aquatic Organisms Dis Aquat Org,2003, 56:95-104.
[65]Raymundo L J, Rosell K B, Reboton C T, Kaczmarsky L. Coral disease onPhilippine reefs:genus Porites is a dominant host. Diseases of Aquatic Organisms Dis Aquat Org,2005,64:181-191.
[66]Reshef L, Koren O, Loya Y, Zilber-Rosenberg I, Rosenberg E. The coral probiotic hypothesis. Environ Microbiol,2006,8(12):2068-2073.
[67]Richard B. Aronson, William F. Precht. White-band disease and the changing face of Caribbean coral reefs. Hydrobiologia,2001,460:25-38.
[68]Richardson L L, Goldberg W M, Carlton R G et al. Coral disease outbreak in the Florida Keys:Plague Typer Ⅱ.Rev. Biol. Trop.,1998,46(5):187-198.
[69]Richardson L L, Smith G W, Richie K B, Carlton R G. Integrating microbiologica, microbiologica, microsensor, molecular, and physiologic techniques in the study of coral disease pathogenesis. Hydrobiologia,2001,460: 71-89.
[70]Richardson L L. Red band disease:a new cyanobacterial infestation of corals. Proceedings of the American Academy of Underwater Sciences Scientific Diving Symposium,1992,pp.153-160.
[71]Richardson L L. Coral diseases:what is really known? Trends in Ecology and Consveration.1998,13,438-443.
[72]Richardson L L. Coral diseases:What is really known? Trends Ecol Evol, 1998,13:438-443.
[73]Ritchie K B, Smith G W. Type II white-band disease. Revista de Biological Tropical,1998,46:199-203.
[74]Rogers C. Coral bleaching and disease should not be underestimated as causes of Caribbean coral reef decline. Proc Biol Sci,2009,276(1655):197-200.
[75]Rohwer F, Breitbart M, Jara J, et al. Diversity of bacteria associated with the Caribbean coral Montastrea franksi. Coral Reefs,2001,20:85-91.
[76]Rosenberg E and Ben-Haim Y. Microbial diseases of corals and global warming. Environ. Microbiol.2002,4:318-326.
[77]Rosenberg E, Barash Y. Microbial Diseases of Corals. In:Belkin and Colwell. Oceans and Health:Pathogens in the Marine Environment[M]. New York: Springer,2005,415-430.
[78]Rosenberg E, Falkovitz L. The Vibrio shiloi/Oculina patagonica model system of coral bleaching. Annu Rev Microbiol,2004,58:143-159.
[79]Smith G W, Well E. Coral reef diseases revised. In:Proceedings of the 10th International Coral Reef Symposium. Japan:Japanese Coral Reef Society,2004, 165.
[80]Stimson J, Sakai K, Sembali H. Interspecific comparison of the symbiotic relationship in corals with high and low rates of bleaching-induced mortality. Coral Reefs,2002,21:409-421.
[81]Sutherland K P, Porter J W, Torres C. Disease and immunity in Caribbean and Indo-Pacific zooxanthellate corals. Marine Ecology Progress Series Mar Ecol Prog Ser,2004,266:273-302.
[82]Vallaeys T, Topp E, Muyzer G, et al. Evaluation of denaturing gradient gel electrophoresis in the detection of 16S rDNA sequence variation in rhizobia and methanotrophs. FEMS Microbiology Ecology,1997,24:279-285.
[83]Vargas-Angel B. Coral health and disease assessment in the U.S. pacific remote island areas. Bulletin of Marine Science,2009,84(2):211-227.
[84]Venter J C, Remington K, Heidelberg J F,et al. Environmental genome shotgun sequencing of the Sargasso Sea.Science,2004,304:66-74.
[85]Veron J E N, Stafford-Smith M. Corals of the world. Australia:Austrlian Institute of Marine Science,2000,1(62-167),2(24-45),3(156-175,276-345).
[86]Wilkinson C. Status of Coral Reefs of the World:2004. Town sville, Queens land: Australian Institute of Marine Science Press,2004. (1):1-316.
[87]Wilkinson C. Status of coral reefs of the world:2000. Global Coral Reef Monitoring Network,Australian Institute of Marine Science, Cape Ferguson, Queensland,2000.
[88]Williams G J, Aeby G S, Cowie R O M et al. Predictive Modeling of Coral Disease Distribution within a Reef System. PLoS ONE,2010,5(2):e9264.
[89]Williams G J, Work T M, Aeby G S et al. Gross and microscopic morphology of lesions in Cnidaria from Palmyra Atoll, Central Pacific. Journal of Invertebrate Pathology,2011,106:165-173.
[90]Work T M, Richardson L L, Reynolds T L, Willis B L. Biomedical and veterinary science can increase our understanding of coral disease. Joural of Experimental Marine Biology and Ecology.2008,362:63-70.