臺灣博碩士論文加值系統

珊瑚礁生態是一個貧營養鹽的海洋生態系，其中海葵與蟲黃藻(zooxanthellae)的共生是一個重要的生物現象，然而海葵和共生藻(Symbiodinium spp.)的共生關係以及牠們的交互作用機制仍未完全了解。本研究即探討當共生藻無法進行光合作用提供海葵營養物質時，海葵的反應。在光照及完全黑暗狀態下餵食豐年蝦與不餵食的三種處理方式來做比較。結果顯示經過24天，黑暗完全不餵食海葵的重量、蛋白質含量、共生藻密度、光合作用能力 (Fv / Fm) 從第6天開始下降到24天後，分別減少了5 %、80 %、60 %、20 % ( p<0.05)，一樣在黑暗中但有餵食宿主的蛋白量僅減少30 % ( p<0.05)，並也其共生藻密度與光合作用能力 (Fv / Fm) 與控制組比起來沒有顯著差異（p >0.05）。再者比較處理24天後海葵的生化指標，結果顯示穀氨酸去氫酶(GDH)、總胺基酸以及必需胺基酸有顯著差異(p<0.05)；於黑暗中無餵食與有餵食的海葵體內GDH 活性在第12天時開始上升至第24 天，上升的量已超過100 % ；然而在光照並且有餵食的海葵GDH 活性僅有增加17 % ；所以海葵宿主若處於完全黑暗環境中無論是沒有餵食或有餵食，其GDH活性都會比光照有餵食來的高（p < 0.05）；海葵在黑暗下有餵食平均會減少59.0 % 總FAA 以及26.4 % EFAA；在完全黑暗不餵食則平均會降低73.8 % 總FAA 以及76.8 % EFAA。另外以人工方式給予碳源來取代光照，結果在完全黑暗狀態下餵食豐年蝦或甘油，經過24天後比較對照組(照光+餵食豐年蝦)與兩處理組之間的重量、蛋白量、共生藻數量、光合作用能力II (Fv /Fm) 、GDH，皆有顯著差異( p < 0.05)；兩處理組(餵食豐年蝦及甘油)由第6天開始下降至第24天，其重量、蛋白量、共生藻密度、光合作用能力II (Fv / Fm)與照光時比起來一樣是較低的，則分別減少了重量為30 %、42 %；蛋白量為52 %、65 % ；光合作用能力II (Fv / Fm)為6 %、8 % 以及共生藻密度也減少了53 %、 44%，並且兩處理組的GDH 活性一樣增加，其增加的活性分別都超過100 %（p < 0.05）；僅有 MDH無明顯差異( p > 0.05)。由以上結果顯示，在黑暗的環境下共生藻無法行光合作用，不論是餵食豐年蝦或供給碳源，海葵的狀態與照光狀態比起來皆有明顯的差異，所以就算以人工方式額外供給氮源或碳源，只要共生藻無法行光合作用提供營養給宿主，宿主依然無法維持與照光時相似的生理狀態。

Coral reef ecosystem is a nutrient poor marine ecosystem which anemone and zooxanthellae symbiosis is an important biological phenomena. However, mechanisms of symbiosis of anemones and zooxanthellae and their interaction are not yet fully understood. The aim of this research was to explore the anemones response, which when symbiotic algae unable to carry out photosynthesis and provided nutrients to anemones. Three approaches for comparison in the state: (1) light and feeding the Artemia sp.; (2) complete darkness and feeding the Artemia sp.; and (3) complete darkness and not fed. The results showed the anemone’s weight, protein count, alga density, photosynthetic capacity (Fv / Fm) began to decline from the first 6 days to 24 days in darkness and feeding, its weight, protein count, alga density, photosynthetic capacity (Fv / Fm) decreased by 5%, 8%, 60%, 20% ( p < 0.05 ), but the same in the darkness and feeding of host which protein reduction of only 30% ( p < 0.05 ), despite a decrease in the amount, but did not like the no more than feeding anemone to many, and there alga density and photosynthesis capacity (Fv / Fm) compared to the control, there was no significant difference ( p > 0.05 ). Futher comparison anemones biochemical through 24 days indicators showed that glutamate dehydrogenase, total free amino acids and essential amino acids are significantly different ( p < 0.05 ), in the dark with or without feeding anemone’s vivo GDH activity began to rise in the first 12 dasy rising to 24 days volume has exceeded 100%, however, the GDH activity increased by 17% in the general state (light and feeding ), if host is in a completely dark environment, no matter there is no feeding or feeding, the activity will be high than the control ( p < 0.05 ), anemones average decrease of 59% total FAA and 26.4% EFAA in the darkness and feeding; in darkness without feeding will decrease on average 73.8% total FAA and76.8% EFAA. Another way we use of artificial to give carbon to replace lighting. The results are displayed in completely dark state feeding Artemia sp. or glycerol , comparing weight, protein count, alga density, photosynthetic capacity (Fv / Fm) and glutamate dehydrogenase of control (lighting and feeding) and between the two treatment after 24 days which all significant difference ( p < 0.05 ), compare two treatment (fed Artemia sp. and glycerol) and the control (lighting and feeding), two treatment were relatively low began to decrease from the fist 6 days to 24 days, its weight, protein count, alga density, photosynthetic capacity (Fv / Fm), respectively, reducing the weight of 30%, 42%; protein count of 52%, 65%; photosynthetic capacity (Fv / Fm) of 6%, 8% and alga density is reduced by 53%, 44%. Both treatment’s GDH activity is increased, which the activity of both increased by more than 100% ( p < 0.05 ), only malate dehydrogenase was no significant differences ( p > 0.05 ), from the above result, in the case of complete darkness caused by symbiotic algae can not photosynthesize, either not fed Artemia sp. or supply carbon, however the state of anemones compared with light status were have significant differences, so even to use of artificial means supplying additional nitrogen or carbon source, and as long as the symbiotic algae can not to carry out photosynthesis provide nutrition to the host, the host is still unable to maintain the light similar of physiological state.

目錄
中文摘要............................................................................................I
英文摘要......................................................................................... III
謝辭................................................................................................V
目錄...............................................................................................VI
圖次............................................................................................... V
表次...............................................................................................XI
第壹章 前言......................................................................................... 1
第貳章 文獻回顧...................................................................................... 5
2.1 海葵............................................................................................ 5
2.2 共生藻（Symbiodinium）........................................................................... 5
2.3 刺細胞動物與共生藻之間的共生關係................................................................... 6
2.4 宿主和共生體各自扮演的角色......................................................................... 8
2.5 珊瑚礁的重要性................................................................................... 10
2.6 白化（Bleaching）................................................................................ 11
2.7 白化的原因及影響.................................................................................. 11
2.8 白化的生化指標.................................................................................... 13
2.8.1 榖氨酸去氫酶 (glutamate dehydrogenase；GDH）..................................................... 13
2.8.2. 蘋果酸去氫酶 (malate dehydrogenase；MDH） ...................................................... 14
2.8.3 游離胺基酸（free amino acids；FAA）.............................................................. 15
第参章 材料與方法....................................................................................... 16
3.1 動物採集與飼養...................................................................................... 16
3.2 動物餵食........................................................................................... 16
3.2.1 豐年蝦孵化........................................................................................ 16
3.2.2 營養混合液配製..................................................................................... 17
3.3 海葵處理的裝置與條件................................................................................. 17
3.3.1 處理裝置.......................................................................................... 17
3.3.2 海葵中共生藻光合作用能力測定....................................................................... 17
3.3.3 海葵共生藻的分離與組織液的製備.......................................................................18
3.4 蘋果酸去氫酶 (MDH)與穀氨酸去氫酶 (GDH)的活性測定....................................................... 18
3.4.1 海葵組織之游離胺基酸 (FAA) 分析..................................................................... 19
3.5 宿主組織均質液中蛋白量的測定.......................................................................... 20
3.6 統計分析............................................................................................ 20
第肆章 結果.............................................................................................. 21
第伍章 討論.............................................................................................. 24
參考文獻................................................................................................. 58

圖目錄
圖1、(A)為共生狀態的海葵(Aiptasia pulchella)、(B)培養於黑暗狀態下， 24 天後的海葵............................. 29
圖2、孵化豐年蝦的裝置...................................................................................... 30
圖3、豐年蝦孵化24 小時後的情形............................................................................. 31
圖4、豐年蝦為向光性的生物，當孵化24 小時後則以遮光的方式使得豐
年蝦往下並朝著有光線的出水口集中............................................................................ 32
圖5、於海水中以豐年蝦餵食海葵(Aiptasia pulchella)的情形..................................................... 33
圖6、海葵麻醉裝置......................................................................................... 34
圖7、標準胺基酸以o-phthaldialdehyde 作管柱前衍生化後之HPLC 分析圖譜.......................................... 35
圖8、於不同處理下海葵(Aiptasia pulchella)宿主重量經過 24 天後的變化情形.......................................36
圖9、不同處理下海葵(Aiptasia pulchella)宿主組織中的蛋白含量經過24天後的變化情形................................ 37
圖10、不同處理下海葵(Aiptasia pulchella)宿主組織中的共生藻光合作用能力經過24 天後的變化情形..................... 38
圖11、不同處理下海葵(Aiptasia pulchella)宿主組織中的共生藻密度經過24 天後的變化情形............................. 39
圖12、不同處理下海葵(Aiptasia pulchella)宿主組織中之GDH 活性經過24 天後的變化情形.............................. 40
圖13、不同處理下海葵(Aiptasia pulchella)宿主組織中之MDH 活性經過24 天後的變化情形.............................. 41
圖14、不同處理下海葵(Aiptasia pulchella)經過 24 天後，宿主組織中總胺基酸含量的變化情形.......................... 42
圖15、不同條件下海葵(Aiptasia pulchella)經過 24 天後，宿主組織中胺基酸組成..................................... 43
圖16、不同條件下海葵(Aiptasia pulchella)經過 24 天後，宿主組織中胺基酸組成之變化................................ 44
圖17、不同環境處理下海葵(Aiptasia pulchella)宿主組織中之Total protein amino acid 經過 18 天後的變化情形......... 45
圖18、不同處理下海葵(Aiptasia pulchella)宿主組織中之Total essential amino acid 經過 24 天後的變化情形........... 46
圖19、兩種不同處理下海葵(Aiptasia pulchella)宿主重量經過24 天後的變化情形....................................... 47
圖20、兩種不同處理下海葵(Aiptasia pulchella)宿主組織中的蛋白含量經過24 天後的變化情形............................. 48
圖21、兩種不同處理下海葵(Aiptasia pulchella)宿主組織中的共生藻密度經過 24 天後的變化情形.......................... 49
圖22、於兩種不同處理下海葵(Aiptasia pulchella)宿主組織中的共生藻光合作用能力經過 24 天後的變化情形................. 50
圖23、兩種不同處理下海葵(Aiptasia pulchella)宿主組織中之GDH 活性經過 24 天後的變化情形............................ 51
圖24、兩種不同處理下海葵(Aiptasia pulchella)宿主組織中之MDH 活性經過 24 天後的變化情形............................ 52

表目錄
表1、Seawater buffer ( SW buffer) (pH 8.2) 的組成............................................................. 53
表2、海葵 (Aiptasia pulchella) 在共生、在完全黑暗有餵食豐年蝦與在完全黑暗沒有餵食的條件下，經過24 天後測試動物組織中之蛋白量、
FAA、MDH、GDH 等生化指標的含量................................................................................. 54
表3、海葵 (Aiptasia pulchella) 在共生、在完全黑暗有餵食豐年蝦與在完全黑暗沒有餵食的條件下，經過24 天後測試宿主之重量、共生藻密
度、光合作用能力等生理指標的含量................................................................................ 55
表4、海葵 (Aiptasia pulchella) 在共生、在完全黑暗有餵食豐年蝦與在完全黑暗有餵食營養液的條件下，經過 24 天後測試宿主之重量、共
生藻密度、光合作用能力等生理指標的含量........................................................................... 56
表5、海葵 (Aiptasia pulchella) 在共生、在共生、在完全黑暗有餵食豐年蝦與在完全黑暗有餵食營養液的條件下，經過24 天後測試動物組織
中之蛋白量、MDH、GDH 等生化指標的含量........................................................................... 57

陳昭倫 (2005) 珊瑚白化：滅絕的鑼聲？樂觀的演化適應？一個生物多樣性觀點。中央研究院週報1037：5-8。
陳昱竹 (2012) 利用薄荷腦白化珊瑚之研究：白化珊瑚之生理生化分析及重建異源共生藻之嘗試。大仁科技大學生物科技所碩士論文。台灣。
黃柏翔 (2011) 美麗海葵 HRS 蛋白在胞內共生所扮演的角色之研究。國立高雄大學生物科技研究所碩士論文。台灣。
Baird, A. H., Bhagooli, R., Ralph, P. J. &Takahashi, S. (2009). Coral bleaching: the role of the host. Trends in Ecology & Evolution 24(1):16-20.
Bellantuono, A. J., Hoegh-Guldberg, O. & Rodriguez-Lanetty, M. (2012).Resistance to thermal stress in corals without changes in symbiont composition. Proceedings of the Royal Society B: Biological Sciences 279(1731): 1100-1107.
Brown, B. E. (1997). Coral bleaching: causes and consequences. Coral Reefs 16: 129-138.
Buddemeier, R. W., Kleypas, J. A. & Aronson, R. B. (2004). Coral Reefs and Global Climate Change. Prepared for the Pew Center on Global Climate Change pp 1- 4.
Baker, A. C. (2003). Flexibility and specificity in coral-algal symbiosis: diversity, ecology, and biogeography of Symbiodinium. Annual Review of Ecology Evolution and Systematics pp 661-689.
Brown, B. E. (2000). The significance of pollution in eliciting the bleaching response in symbiotic cnidarians. International Journal of Environment and Pollution 13: 392–415 .
Ben-Haim,Y. & Rosenberg, E. (2002). A novel Vibrio sp. pathogen of the coral Pocillopora damicornis. Marine Biology 141: 47–55.
Brundrett, M. C. (2002). Coevolution of roots and mycorrhizas of land plants.
New Phytologist 154: 275–304.
Banin, E., Khare, S. K., Naider, F. & Rosenberg, E. (2001). Proline-rich peptide from the coral pathogen Vibrio shiloi that inhibits photosynthesis of zooxanthellae. Applied and environmental microbiology 67:1536.
Chen, C. A. & Dai, C. F. (2004). Local phase shift from Acropora-dominant to Condylactis-dominant community in the Tiao-Shi Reef, Kenting National Park, southern Taiwan. Coral Reefs 23(4): 508-508.
Clarke, K. R. &Warwick, R. M. (1994). Change in Marine Communities. Plymouth Marine Laboratory: 144.
Coffroth, M. A. & Santos, S. R. (2005). Genetic Diversity of Symbiotic Dinoflagellates in the Genus Symbiodinium. Protist 156(1): 19-34.
Chen, M.C., Cheng, Y. M., Sung, P. J., Kuo, C. E. & Fang, L. S. (2003). Molecular identification of Rab7 (ApRab7) in Aiptasia pulchella and its exclusion from phagosomes harboring zooxanthellae Biochemical and biophysical research communications 308: 586-595.
Chen, M.C., Cheng, Y.M., Hong, M.C. & Fang, L.S. (2004) . Molecular cloning of Rab5 (ApRab5) in Aiptasia pulchella and its retention in phagosomes harboring live zooxanthellae. Biochemical and biophysical research communications 324: 1024-1033.
Chen, M. C., Hong, M. C., Huang, Y. S., Liu, M. C., Cheng, Y. M. & Fang, L. S. (2005). ApRab11, a cnidarian homologue of the recycling regulatory protein Rab11, is involved in the establishment and maintenance of the Aiptasia-Symbiodinium endosymbiosis.
Biochemical and biophysical research communications 338: 1607-1616.
Downs, C. A., Kramarsky, W. E., Martinez, J., Kushmaro, A., Woodley, C. M., Loya, Y. & Ostrander, G. K. (2009). Symbiophagy as a cellular mechanism for coral bleaching. Autophagy 5: 211-216.
Dubinsky, Z. (1990). Coral Reefs: Ecosystems of the World. Elsevier Science 25: 550 .
Douglas, A. E. & Smith, DC. (1983). The cost of symbionts to their host in green hydra. Endocytobiology 2: 631-648.
Douglas, A. E. (2005). Bugs within bugs: symbiotic bacteria in garden insects.
Microbiology Today 32: 76-79.
Dubinsky, Z. & Jokiel, P. (1994). Ratio of energy and nutrient fluxes 60
regulates symbiosis between zooxanthellae and corals. Pacific Science 48(3): 313-324.
Douglas, A. E. (1994). Symbiotic Interactions. Oxford University Press.
Douglas, A. E. (2003). Coral bleaching--how and why? Marine Pollution
Bulletin 46:385-392.
Dunn, S.R., Bythell, J.C., Le Tissier, M.D.A., Burnett, W.J. & Thomason,
J.C. (2002). Programmed cell death and cell necrosis activity during hyperthermic stress-induced bleaching of the symbiotic sea anemone Aiptasia sp. Experimental Marine Biology and Ecology 272: 29-53.
Dunne, R. P. & Brown, B. E. (2001). The influence of solar radiation on bleaching of shallow water reef corals in the Andaman Sea, 1993-1998. Coral Reefs 20(3): 201-210
Fitt, W., Brown, B., Warner, M. & Dunne, R. (2001). Coral bleaching: interpretation of thermal tolerance limits and thermal thresholds in tropical corals. Coral Reefs 20(1): 51-65.
Falkowski, P.G., Dubinsky, Z., Muscatine L. & Porter, J.W. (1984). Light and the bioenergetics of a symbiotic coral. Bioscience 34: 705—709.
Falkowski, P.G., Dubinsky, Z., Muscatine, L. & McCloskey, L. (1993). Population control insymbiotic corals. Ammonium ions and organic materials maintain the density of zooxanthellae. Bioscience 43:606–611.
Fransolet, D., Roberty S. & Plumier J. C. (2012). Establishment of endosymbiosis: The case of cnidarians and Symbiodinium. Experimental Marine Biology and Ecology 420: 1-7.
Frost, T. M. & Wiilliamson, C. E. (1980). In situ determination of the effect of symbiotic algae on the growth of the freshwater sponges Spongilla lacustris. Ecology. 6: 1367-1370.
Grottoli, A. G., Rodrigues, L. J. & Juarez, C. (2004). Lipids and stable carbon
isotopes in two species of Hawaiian corals, Porites compressa and Montipora verrucosa, following a bleaching event. Marine Biology 145(3).
Grabowska, A., Nowicki, M. & Kwinta, J. (2011). Glutamate dehydrogenase 61
of the germinating triticale seeds: gene expression, activity distribution and kinetic characteristics. Acta Physiologiae Plantarum 33(5): 1981-1990.
Goulet, T. L. (2006). Most corals may not change their symbionts. Marine ecology progress series 321: 1-7.
Gates, R.D., Baghdasarian, G. & Muscatine, L. (1992). Temperature stress causes host cell detachment in symbiotic cnidarians: implications for coral bleaching. The Biological Bulletin 182: 324.
Hallock P. (2001). Coral reefs, carbonate sediments, nutrients and global change. In Stanley Jr. GD (ed) The History and Sedimentology of Ancient Reef Systems. Springer-Verlag pp387—427.
Hoegh-Guldberg, O. (1999). Climate change, coral bleaching and the future of the world's coral reefs. Marine and Freshwater Research 50(8): 839-866.
Hoegh-Guldberg, O. & Jones, R. (1999). Photoinhibition and photoprotection in symbiotic dinoflagellates from reef-building corals Marine Ecology-progress Series 183: 73-86.
Hoegh-Guldberg, O., Mumby, P. J., J., H. A., Steneck, R. S., Greenfield, P., Gomez, E., Harvell, C. D., Sale, P. F., Edwards, A. J., Caldeira, K., Knowlton, N., Eakin, C. M., Iglesias-Prieto, R., Muthiga, N., Bradbury, R. H., Dubi, A. & Hatziolos, M. E. (2007). Coral reefs under rapid climate change and ocean acidification. Science 318(5857): 1737-1742.
Hoegh-Guldberg, O. & Smith, G. J. (1989). Influence of the population density of zooxanthellae and supply of ammonium on the biomass and metabolic characteristics of the reef corals Seriatopora hystrix and Stylophora pistillata. Marine Ecology Progress Series 57: 173-186.
Hughes, T. P. (1994). Catastrophes, phase-shifts and large-scale degradation of Caribbean reef. Science 265: 1547-1551.
Hughes, T. P., Baird, A. H., Bellwood, D. R., Card, M., Connolly, S. R., Folke, C., Grosberg, R., Hoegh-Guldberg, O., Jackson, J. B. C., Kleypas, J., Lough, J. M., Marshall, P., Nyström, M., Palumbi, S. R., 62 Pandolfi, J. M., Rosen, B. & Roughgarden, J. (2003). Climate Change, Human Impacts, and the Resilience of Coral Reefs. Science 301(5635): 929-933.
Hughes, T. P., Rodrigues, M. J., Bellwood, D. R., Ceccarelli, D., Hoegh-Guldberg, O., McCook, L., Moltschaniwskyj, N., Pratchett, M. S., Steneck, R. S. & Willis, B. (2007). Phase shifts, herbivory, and the resilience of coral reefs to climate change. Current Biology 17(4): 360-365.
Harvell, C. D., Kim, K., Burkholder, J. M., Colwell, R. R., Epstein, P. R., Grimes, D. J., Hofmann, E. E., Lipp, E. K., Osterha us , A. D. M. E., Overstreet, R. M., Porter, J. W., Smith, G. W. & Vasta, G. R. (1999). Emerging marine diseases––climate links and anthropogenic factors. Science 285:1505–1510.
Jackson, J. B. C., Kirby, M. X., Berger, W. H., Bjorndal, K. A., Botsford, L. W., Bourque, B. J., Bradbury, R. H., Cooke, R., Erlandson, J., Estes, J. A., Hughes, T. P., Kidwell, S., Lange, C. B., Lenihan, H. S., Pandolfi, J. M., Peterson, C. H., Steneck, R. S., Tegner, M. J. & Warner, R. R. (2001). Historical Overfishing and the Recent Collapse of Coastal Ecosystems. Science 293(5530): 629-637.
Jones, R. J. (1997). Zooxanthellae loss as a bioassay for assessing stress in
corals. Marine Ecology Progress Series 149: 163-171.
Jones, R. J. (2004). Testing the ‘photoinhibition’ model of coral bleaching using chemical inhibitors. Marine Ecology Progress Series 284: 133-145.
Jones, R. (2005). The ecotoxicological effects of Photosystem II herbicides on corals. Marine Pollution Bulletin 51(5): 495-506.
Jones, R. J., Hoegh-Guldberg, O., Larkum, A. W. D. & Schreiber, U. (1998). Temperature-induced bleaching of corals begins with impairment of the CO2 fixation mechanism in zooxanthellae. Plant Cell and Environment 21(12): 1219-1230.
Jones, R. J., Ward, S., Yang Amri, A. & Hoegh-Guldberg, I. O. (2000). 63
Changes in quantum efficiency of Photosystem II of symbiotic dinoflagellates of corals after heat stress, and of bleached corals sampled after the 1998 Great Barrier Reef mass bleaching event Marine and Freshwater Research 51(1): 63-71.
Krause, G.H. & Weis E. (1991). Chlorophyll fluorescence and photosynthesis: the basic. Annual Review Plant Physiology and Plant Molecular Biology 42: 313-349.
Kiers, E. T., Rousseau1, R. A., West, S. A. & Denison, R. F. (2003). Host sanctions and the legume–rhizobium mutualism. Nature 425: 78-81.
Lesser, M. P. & Farrell, J. H. (2004). Exposure to solar radiation increases
damage to both host tissues and algal symbionts of corals during thermal stress. Coral Reefs 23(3): 367-377.
Lewis, J. B. (1977). Processes of organic production on coral reefs. Biological
Reviews of the Cambridge Philosophical Society 52(3): 305-347.
Lin, K.L., Wang, J.T. & Fang, L.S. (2000). Participation of glycoproteins on zooxanthellal cell walls in the establishment of a symbiotic relationship with the sea anemone, Aiptasia pulchella. Zoological Studies. 39: 172-178.
MacDonald, M. J. (1982). Evidence for the malate aspartate shuttle in pancreatic islets. Archives of Biochemistry and Biophysics 213(2): 643-649.
Martin-Jézéquel, V., Sournia, A. & Birrien, J.-L. (1992). A daily study of the
diatom spring bloom at Roscoff (France) in 1985. III. Free amino acids composition studied by HPLC analysis. Journal of Plankton Research 14(3): 409-421.
Maxwell K. & Johnson G.N. (2000). Chlorophyll fluorescence － a practical guide. Journal of experimental botany 51: 659-668.
Mieog, J. C., Olsen, J. L., Berkelmans, R., Bleuler-Martinez, S. A., Willis, B.
L. & van Oppen, M. J. (2009). The roles and interactions of symbiont, host and environment in defining coral fitness. Public Library of Science ONE 4(7): 63-64.
Moore, S. J. (1989). Narcotizing sea anemones. Marine biological association of the united kingdom 69: 803-811.
Mullinax, T. R., Mock, J. N., McEvily, A. J. & Harrison, J. H. (1982). Regulation of mitochondrial malate dehydrogenase. Evidence for an allosteric citrate-binding site. Biological Chemistry 257(22): 33-39.
Muscatine, L. (1967). Glycerol excretion by symbiotic algae from corals and Tridacna and its control by the host. Science 156(37): 516-519.
Muscatine, L. (1990). The role of symbiotic algae in carbon and energy flux in reef corals. In: Dubinsky Z. (ed), Ecosystems of the world: Coral reefs. Ecosystems of the World 25: 45-87.
Muscatine L., Falkowski P. G., Porter J. W. & Dubinsky Z. (1984). Fate of photosynthetically fixed carbon in light and shade-adapted colonies of the symbiotic coral, Stylophora pistillata. Proceedings of the Royal Society of London Series B-Biological Sciences 222: 181-202.
Muscatine, L. & Pool, R. R. (1979). Regulation of numbers of intracellular algae. Proceedings of the Royal Society of London Series B-Biological Sciences 204: 131–139.
Marlow, H.Q. & Martindale, M.Q. (2007). Embryonic development in two species of scleractinian coral embryos: Symbiodinium localization and mode of gastrulation. TRENDS ECOL EVOL 9: 355–367.
Muscatine, L., Falkowski, P.G., Dubinsky, Z., Cook, P.A. & McCloskey, L.R. (1989). The effect of external nutrient resources on the population dynamics of zooxanthellae in a reef coral. Proceedings of the Royal Society of London Series B-Biological Sciences 236: 311–324.
Pochon, X., Montoya-Burgos, J. I., Stadelmann, B. & Pawlowski, J. (2006). Molecular phylogeny, evolutionary rates, and divergence timing of the symbiotic dinoflagellate genus Symbiodinium. Molecular Phylogenetics and Evolution 38(1): 20-30.
Pochon, X. & Gates, R. D. (2010). A new Symbiodinium clade (Dinophyceae) from soritid foraminifera in Hawai'i. Molecular Phylogenetics and Evolution 56:492–497.
Rahav, O., Dubinsky, Z., Achituv, Y. & Falkowski, P. G. (1989). Ammonium metabolism in the zooxanthellate Coral, Stylophora pistillata.
Proceedings of the Royal Society of London Series B-Biological Sciences 236(1284): 325-337.
Rees, T. A. V. (1986). The Green Hydra Symbiosis and Ammonium I. The Role of the Host in Ammonium Assimilation and its Possible Regulatory Significance. Proceedings of the Royal Society of London Series B-Biological Sciences 229(1256): 299-314.
Perez S. & Weis V. (2006). Nitric oxide and cnidarian bleaching: an eviction notice mediates breakdown of a symbiosis. Experimental Biology 209: 2804-2810.
Souter, D. W. & Lindén, O. (2000). The health and future of coral reef systems. Ocean & Coastal Management 43(8): 657-688.
Sawyer, S. J. & Muscatine, L. (2001). Cellular mechanisms underlying temperature-induced bleaching in the tropical sea anemone Aiptasia pulchella. Experimental biology 204: 34－43.
Saldarriaga, J. F., Taylor, F. J. R., Keeling, P. J. & Cavalier-S mith, T. (2001). Dinoflagellate nuclear SSU rRNA phylogeny suggests multiple plastid losses and replacements. Molecular Evolution 53:204–213 .
Schreiber U., Bilger W. & Nubauer C. (1994). Chlorophyll fluorescence as a nonintrusive indicator for rapid assessment of in vivo photosynthesis. Springer-Verlag, Berlin. pp 49-70.
Schreiber U. (2004). Pulse－Amplitude (PAM) fluorometry and saturation pulse method. In: Papageorgiou G, Govindjee (eds) Chlorophyll Fluorescence: A signature of Photosynthesis. Kluwer Academic Publishers, Dordrecht, The Netherlands pp 279-319.
Simon K. D. & Clayton B. C. (2001). The influence of ｀host release factor＇ on carbon release by zooxanthellae isolated from fed and starved Aiptasia pallida. Comprative Biochemistry and Physiology. Part A, Molecular & Integrative Physiology 129(2):487-494.
Stat, M., Morris,E. & Gates, R. D. (2008). Functional diversity in coral–dinoflagellate symbiosis. Proceedings of the National Academy of Sciences 105: 9256.
Stat, M., Carter, D. & Hoegh-Guldberg, O. (2006). The evolutionary history of Symbiodinium and scleractinian hosts—symbiosis, diversity, and the effect of climate change. Perspectives Plant Ecology Evolution Systematics 8: 23–43.
Taylor, D. L. (1974). Symbiotic marine algae: taxonomy and biological fitness. In: Vernberg, W. B. (ed.), Symbiosis in the Sea. University of South Carolina Press, South Carolina. pp 245–262.
Venn, A. A., Loram, J. E. & Douglas, A. E. (2008). Photosynthetic symbioses in animals. Experimental Botany 59(5): 1069-1080.
Vidal-Dupiol, J., Adjeroud, M., Roger, E., Foure, L., Duval, D., Mone, Y., Ferrier-Pages, C., Tambutte, E., Tambutte, S. & Zoccola, D. (2009). Coral bleaching under thermal stress: putative involvement of host/symbiont recognition mechanisms. BioMed Centrol physiology 9: 14.
Wang, J. T. & Douglas, A. E. (1997). Nutrients, signals, and photosynthate release by symbiotic algae: the impact of taurine on the dinoflagellate alga Symbiodinium from the sea anemone Aiptasia pulchella. Plant Physiology 114: 213-221.
Wang, J. T. & Douglas, A. E. (1998). Nitrogen recycling or nitrogen conservation in an alga-invertebrate symbiosis? Experimental Biology 201(16): 2445-2453.
Wang, J. T. & Douglas, A. E. (1999). Essential amino acid synthesis and nitrogen recycling in an alga–invertebrate symbiosis. Marine Biology 135(2): 219-222.
Warner, M.E., Fitt, W.K. and Schmidt, G.W. (1999). Damage to photosystem II in symbiotic dinoflagellates: a determinant of coral bleaching. Proceedings of the National Academy of Sciences 96: 8007.
Weis, V. M. (2008). Cellular mechanisms of Cnidarian bleaching: stress causes the collapse of symbiosis. Experimental Biology 211(19): 3059-3066.
Wilkinson, C. R. (1999). Global and local threats to coral reef functioning and
existence: review and predictions. Marine and Freshwater Research 50(8): 867-878.
Whitehead L.F. & Douglas A.E. (2003). Metabolite comparisons and the identity of nutrients translocated from symbiotic algae to an animal host. Journal of experimental Biology 206: 3149-3157.
Yamashiro, H., Oku, H. & Onaga, K. (2005). Effect of bleaching on lipid content and composition of Okinawan corals. Fisheries Science 71(2): 448-453.
Yellowlees, D., Rees, T. A. & Leggat, W. (2008). Metabolic interactions between algal symbionts and invertebrate hosts. Plant Cell and environment 31(5): 679-694.