對大部份底棲性且採體外授精的海洋生物來說，同一物種同時大量排放精卵有其必要性。每年珊瑚大量產卵的時間準確且可以預測，但是目前卻無法準確地預測海膽同步排放精卵的時間。我們藉設計觀察方法來了解兩種海膽排放精卵的潛力，首先檢視各種有無集中排放的現象，而後再以和潮水相關的農曆日期分析排放比率，驗證牠們的排放是否有一定週期。從月間的資料沒有發現任何一種海膽有明顯的月、半月週期律動。但是經由二項式檢定發現，兩種海膽的排放不是隨機分布，而有集中在某些天的現象。一天之內，喇叭毒棘海膽白天和晚上的排放比率有顯著的差異，白天排放比率較高，經控制光線實驗發現牠的排放有內生性日週期參與調控的現象；白棘三列海膽則無。兩種海膽針對近日週期性出現的因子如：潮汐做檢定，發現高潮或低潮，海膽排放的比率沒有明顯差異，顯然潮汐不會影響大量排放精卵的時間。所以後續針對非週期性出現的因子做驗證，喇叭毒棘海膽經實驗觀察發現雄性排放的隻數與時間都較雌性來的早；隔絕海膽個體間味道的交換後配子的排放比率會有顯著差異，暗示可能有類似費洛蒙功用的物質存在，沒有發現接觸有顯著影響；當水溫升高，排放隻數也顯著上升；沒有發現水流擾動及精液、卵液對於排放比率有顯著的影響；沒有發現白棘三列海膽有上述現象。本論文結果顯示：喇叭毒棘海膽同步生殖的現象是水溫、內生性律動配合下的成果。

For marine organisms with external fertilization, it is necessary for conspecific to release sperm and eggs in the same time. But so far, we cannot accurately predict the spawning time of many sea urchins. In this investigation, we monitored the spawning potential of Tripneustes gratilla and Toxopneustes pileolus (Echinodermata: Toxopneustidae) from April 2010 to August 2010 in southern Taiwan in order to find any pattern of spawning and its possible mechanisms. No significant lunar or semilunar rhythms were found. On the other hand, spawning potential of individuals were not randomly nor evenly distributed throughout the months in both species. High frequencies of spawning individuals occurred on certain days with no obvious rhythms. Within a day, the spawning frequencies are higher in the daytime than in the nighttime, and follow up experiments suggest a possible role of an endogenous oscillator in Toxopneustes pileolus. The possible effect of tide levels was not found. In addition, we tested noncyclical factors. Males spawned earlier and in higher frequencies than females, however no inductive ability was detected in either sexes. Significantly higher frequencies of individuals spawned when water temperature was raised, whereas no role of water disturbance was detected. We didn’t find any significant connections between Tripneustes gratilla and environmental factors. So far, water temperature and light/dark condition (through entraining their biological clocks) have been found to play a role in synchronizing the spawning Toxopneustes pileolus only.

摘要................................................................................................................................. i
Abstract .......................................................................................................................... ii
壹、前言........................................................................................................................ 1
貳、材料與方法............................................................................................................ 6
参、結果...................................................................................................................... 11
肆、討論...................................................................................................................... 18
參考文獻...................................................................................................................... 27
附錄.............................................................................................................................. 66

陳國芳，1993，兩種海生搖蚊Pontomyia spp.(雙翅目：搖蚊科)生活史之初研究。國立中山大學海洋生物研究所碩士論文。台灣。中華民國。
林順康，1995，紫海膽Anthocidaris crassispina和口腮海膽Stomopneustes variolaris的生殖週期和晚期幼生飼料實驗。國立海洋大學漁業科學研究所碩士論文。台灣。中華民國。
劉德經，1996，西施舌Coelomactra antiquate(Spengler)誘導產卵的初步探討。福建水產，1:1-4
曹俊仁，1997，兩種海生搖蚊Pontomyia oceana (雙翅目：搖蚊科)同步羽化之研究。國立中山大學海洋生物研究所碩士論文。台灣。中華民國。
陳佳杏，1998，溫度對單細胞藍綠藻生長率的影響與族群大小的季節變化。國立海洋大學海洋生物研究所碩士論文。台灣。中華民國。
李品賢，1999，溫度對海生搖蚊Pontomyia oceana同步羽化之影響與其羽化週律之相關研究。國立中山大學海洋生物研究所碩士論文。台灣。中華民國。
黃貞禎，2008，墾丁沿岸海域營養鹽和浮游植物群聚之時空變異。國立東華大學海洋生物多樣性及演化研究所碩士論文。。台灣。中華民國。
林虹靜，張育菁，趙世民，2001，墾丁海域白棘三列海膽的生殖週期研究。台灣。中華民國。
Andronikov, V. B. 1975. Heat resistance of gametes of marine invertebrates in relation to temperature conditions under which the species exist. Marine Biology 30:1–12
Babcock, R. C., G. D. Bull, P. L. Harrison, A. J. Heyward, J. K. Oliver, C. C. Wallace, and B. L. Willis. 1986. Synchronous spawning of 105 scleractinian coral species on the Great Barrier Reef. Marine Biology 90:373-394
Babcock, R. C., C. N. Mundy, and D. Whitehead. 1994. Sperm diffusion models and In situ confirmation of long-distance fertilization in the free-spawning Asteroid Acanthaster planci. Biological Bulletin 186:17-28
Barnes, H. 1963. Light, temperature and the breeding of Balanus balanoids. Journal of the Marine Biological Association of the United Kingdom 43:717-727
Beatley, J.C. 1974. Phenological events and their environmental triggers in Majave desert ecosystems. Ecology 55:856-863
Boolootian, R. A., and A. R. Moore 1956. Hermaphroditism in echinoids. Biological Bulletin 111:328-335
Byrne, M., M. A. Sewell, and T. A. A. Prowse.2008. Nutritional ecology of sea urchin larvae. Influence of endogenous and exogenous nutrition on echinopluteal growth and phenotypic plasticity in Tripneustes gratilla. Functional Ecology 22:643-648
Chao, S. M. 2000. The irregular sea urchins(Echinodermata: Echinoidea) from Taiwan,with descriptions of six new records. Zoology Study 39(3): 250-265
Chen, A., and K. Soong. 2010. Uncovering behavior at spawning of the trumpet sea urchin Toxopneustes pileolus. Zoological Studies 49(1):9
Chen, C. P., and K. H. Chang.1981.Reproductive periodicity of the sea urchin, Tripreustes gratilla (L.) in Taiwan compared with other regions. International Journal of Invertebrate Reproduction 3: 309-319
Clark, A. M., and F. W. E. Rowe. 1971. Monograph of shallow-water Indo-West Pacific echinoderms. i-vii, 1-238, pls 1-31. London, Trustees of the British Museum (Natural History)
Cochran, R. C., and F. Engelmann. 1975. Environmental regulation of the annual reproductive season of Strongylocentrotus purpuratus (Stimpson). Biological Bulletin 148: 393-401
Cochran, R. C., and F. Engelmann. 1976. Characterization of spawn-inducing factors in the sea urchin, Strongylocentrotus purpuratus. General Comparative Endocrinoogy 30:189-197
Coppard, S. E., and A. C. Campbell. 2005. Distribution and abundance of regular sea urchins on two coral reefs in Fiji. Micronesica 37(2):249-269
Dai, C. F., Soong, K., and T. Y. Fan. 1993. Sexual reproduction of corals in northern and southern Taiwan. In Proceedings of the 7th International Coral Reef Symposium, Guam, Micronesia, 22-27 June 1992. Vol 1. Edited by R. H. Richmond. University of Guam Marine Laboratory, Mangilao. pp. 448-455
Danilo, T. Dy, A. Uy Frederick, and M. Coralles Christine. 2002. Feeding, respiration, and excretion by the tropical sea urchin Tripneustes gratilla (Echinodermata Echinoidea) from the Philippines. Journal of the Marine Biological Association of the United Kingdon 82:299-302
Dodge, C. A., and M. S. Edwards. 2010. Seasonal variation in the effects of food availability on gametogenesis in the purple urchin(Strongylocentrotus purpuratus). Marine Biology 159:427-433
Dworjanyn, S. A. and I. Pirozzi. 2008. Induction of settlement in the sea urchin Tripneustes gratilla by macroalgae, biofilms and conspecifics.A role for bacteria. Aquaculture 274:268-274
Farmanfarmaian, A., and A. C. Giese. 1963. Thermal tolerance and acclimation in the western purple sea urchin, Strongylocentrotus purpuratus. Physiological Zoology. 36:237-243
Fox. H. 1922. Lunar periodicity in reproduction. Nature 109:237-238
Fries, G. 1964.Uber die Einwirkung der Tugesperiodik und der Temperatur auf den Generations Wechsel, die Weibenhergrosse und die Eier von Daphnia magnus Straus. Zeitschrift fuer Morphologie und Okologie der Tiere, Vol.53,pp. 475-516
Fujisawa, H., and M. Shigei. 1990. Correlation of embryonic temperature sensitivity of sea urchins with spawning season. Journal of Experimental Marine Biology and Ecology 136:123-139
Glabe, C. G., and W. J. Lennarz. 1979. Species-specific sperm adhesion in sea urchins: a quantitative investigation of bindin-mediated egg agglutination. Journal of Cell Biology 83:595–604
Garrido, C. L., and B. J. Barber. 2001. Effects of temperature and food ration on gonad growth and oogenesis of the green sea urchin, Strongylocentrotus droebachiensis. Marine Biology 138:447-456
Giese, A. C. 1959. Reproductive cycles of some west coast invertebrates. In: R. B. Withrow, editor. Photoperiodism and related phenomena in plants and animals. American association for the advancement of science New York, pp. 625-638
Giese, A. C., and H. Kanatani. 1987. Maturation and spawning. In A. C. Giese, J. S. Pearse, and V. B. Pearse (eds.), Reproduction of marine invertebrates, Vol. IX. Seeking unity in diversity, pp. 251–329. Blackwell Scientific/Boxwood Press, Palo Alto/Pacific Grove, California.
Gordon, I. 1926. The development of the calcareous test of Echinus miliaris. Royal Society of London, Philosophical Transactions, series B 214:259-312
Guettaf, M., G. A. San, and P. Francour. 2000. Interpopulation variability of the reproductive cycle of Paracentrotus lividus(Echinodermata Echinoidea)in the south-western Mediterranean. Journal of Marine Biological Association of the United Kingdon 80:899-907
Guillou, M. and L. J. L. Lumingas. 1999. Variation in the reproductive strategy of the sea urchin Sphaerechinus granularis (Echinodermata Echinoidea) related to food availability. Journal of Marine Biological Association of the United Kingdom 79:131-136
Guzmán, H. M., C. A. Guevara, and I.C. Hernández. 2003. Reproductive cycles of two commercial species of sea cucumber (Echinodermata: Holothuroidea) from Caribbean Panama. Marine Biology 142:271-279
Gwinner, E. 1981. “Annual rhythms.” handbook of behavioural neurobiology, Volume 4: Biological Rhythms, ed. Jurgen Aschoff. New York: Plenum Press, 381-389
Hagen, N. T., and K.H. Mann. 1994. Experimental analysis of factors influencing the aggregating behaviour of the green sea urchin Strongylocentrotus droebachiensis (Muller). Journal of Marine Biological Association of the United Kingdom 176:107-126
Halberg, F., Y. L. Tong, and E. A. Johnson. 1967. Circadian system phase, an aspect of temporal morphology: procedures and illustrative examples. In: Mayersbach H von, editor. The Cellular Aspects of Biorhythms. Berlin:20-48
Harrison, P. L. Babcock, R. C., Bull, G. D., Oliver, J. K., Wallace, C. C., and B. L. Willis 1984. Mass spawning in tropical reef corals. Science 223:1187-1188
Havenhand, J. N., F-R Buttler, M. C. Thorndyke, J. E. Williamson. 2008. Near-future levels of ocean acidification reduce fertilization success in a sea urchin. Current Biology 18: 651-652
Hegarty, T. W. 1973. Temperature coefficient(Q10), seed germination and other biological processes. Nature 243:305-306
Himmelman, J. H. 1975. Phytoplankton as a stimulus for spawning in three marine invertebrates. Journal of Experimental Marine Biology and Ecology 20:199-214
Himmelman, J. H. 1978. Reproductive cycle of the green sea urchin, Strongylocentrotus droebachiensis. Canadian Journal of Zoology 56(8):1828-1836
Hinegardner, R. T. 1969. Growth and development of the laboratory cultured sea urchin. Biological Bulletin 137:465-475
Hochberg, Y. 1988. A sharper Bonferroni procedure for multiple tests of significance. Biometrika 75:800-802
Horii, T. 1997. The annual reproductive cycle and lunar spawning rhythms of the purple sea urchin Anthocidaris crassispina Nippon Suisan Gakkaishi 63(1), 17-22
Iliffe, T. M., and J. S. Pearse. 1982. Annual and lunar reproductive rhythms of the sea urchin,Diadema antillarum.philippi.in Bermuda. International Journal of Invertebrate Reproduction 5:139-148
Jones, G. M., and N. T. Hagen. 1987 Echinomermella matsi sp.,an endoparasitic nematode from the sea urchin Strongylocentrotus dorebachiensis in northern Norway.Sarsia 72:203-212)
Jones, G. M., and R. E. Scheibling. 1985. Paramoeba sp.(Amoebida, Paramoebidae)as the possible causative agent of sea urchin mass mortality off Nova Scotia. Journal of Parasitology 71:559-565
Juinio-Meñez, M. A., and H. G. P. Bangi. 2010. Extrinsic and intrinsic factors affecting the metamorphic rate of Tripneustes gratilla. Echinodermata Echinoidea Marine Ecology Progress Series 402:137-145
Kao, C. C., J. Chen, G. F. Chen, and K. Soong. 2010. Variable swarming time of an intertidal midge (Pontomyia oceana Tokunaga, 1964) controlled by a circadian clock and temperature. Marine and Freshwater Behavior and Physiology 43(1):1-9
Keckes, S., B. Ozretić, and C. Lucu. 1986. About a possible mechanism involved in the shedding of sea urchins. Experientia 22:146-147
Kimura, A., H. Nakagawa, H. Hayashi1 and K. Endo. 1984. Seasonal changes in contractile activity of a toxic substance from the pedicellaria of the sea urchin Toxopneustes pileolus. Toxicon 22(3):353-358
Kuo, K. M., and K. Soong. 2010. Post-Settlement Survival of Reef-Coral Juveniles in Southern Taiwan. Zoological Studies 49(6):724-734
Kuwabara, S. 1994. Purification and properties of peditoxin and the structure of its prosthetic group, pedoxin, from the sea urchin Toxopneustes pileolus (Lamarck). Journal of Biological Chemistry 269(43) 26734-26738
Kyorku, A. and Brady, J. 1995. Free-running Bimodal (Circadian Rhythm) in the Tsetse Fly (Glossina longipennis). Journal Insect Physiology 40:63-67
Lane, David J. W., Loisette M. Marsh, Didier Van den Spiegel, and Frank W. E. Rowe 2000. Echinoderm fauna of the south china sea: an inventory and analysis of distribution patterns. The Raffles Bulletin of Zoology Supplement 8:459-493
Lau, Dickey C. C., Stanley C. K. Lau, P. Y. Qian, and J. W. Qiu. 2009. Morphological plasticity and resource allocation in response to food limitation and hyposalinity in a sea urchin. Journal of Shellfish Research 28(2):383-388
Lessios, H. A. D.R. Robertson, and J. D. Cubit 1984. Spread of Diadema mass mortality through the Caribbean. Science 226:335-337
Lessios, H. A. 1992. Testing electrophoretic data for agreement with Hardy-Weinberg expectations. Marine Biology 112:517-523
Lessios, H. A. 1991. Presence and absence of monthly reproductive rhythms among 8 caribbean echinoids off the coast of Panama. Journal of Experimental Marine Biology and Ecology 153:27-47
Levitan, Don R. 1988. Density-Dependent Size Regulation and Negative Growth in the Sea Urchin Diadema antillarum Philippi. Oecologia 76: 627-629
Ling, S. D., C. R. Johnson, K. Ridgway, A. J. Hobday and M. Haddon.2009. Climate-driven range extension of a sea urchin: inferring future trends by analysis of recent population dynamics. Global Change Biology 15:719-731
Mayr, E. 1961. Cause and effect in biology. Science 134:1501-1506
McBride, S. C., Pinnix, W. D., Lawrence, J. M., Lawrence, A. L., and T. M. Mulligan. 1997. The effect of temperature on production of gonads by the sea urchin Strongylocentrotus francisanus fed natural and prepared diets. Journal of the World Aquaculture Society 28(4):357-365
McCarthy, Daniel A., and Craig M. Young. 2002. Gametogenesis and reproductive behavior in the echinoid Lytechinus variegates. Marine Ecology Progress Series 233:157-168
McClanahan, T. R. 1998. Predation and the distribution and abundance of tropical sea urchin populations. Journal of Experimental Marine Biology and Ecology 221:231-255
Meats, A. and K. C. Khoo, 1976. The dynamics of ovarian maturation and oocyte resorption in Queensland fruit fly, Dacus tryoni, in daily-rhythmic and constant temperature regimes. Physiological entomology 1:213-221
Meidel, S. K. and R. E. Scheibling. 1998. Annual reproductive cycle of the green sea urchin, Strongylocentrotus droebachiensis, in differing habitats in Nova Scotia, Canada. Marine Biology 131:461-478
Mendes, J. M., and J. D. Woodley. 2002. Timing of reproduction in Montastraea annularis : relationship to environmental variables. Marine Ecology Progress Series 227,241-251
Miller R. J., and A. G. Colodey 1983. Widespread mass mortalities of the green sea urchin in Nova Scotia, Canada. Marine Biology 73:263-267
Muthiga, N. A. 2005. Testing for the effects of seasonal and lunar periodicity on the reproduction of the edible sea urchin Tripneustes gratilla (L) in Kenyan coral reef lagoons. Hydrobiologia 549:57-64
Nunney, L. 1985. Female-biased sex ratios:individual or group selection? Evolution 39(2):349-361
Oliver, J. K., Babcock, R. C., Harrison, P. L., Willis, B. L. 1988 Geographic extent of mass coral spawning: Clues to ultimate causal factors. in Choat, JH, Barnes, D, Borowitzka, MA, Coll, JC, Davies, PJ, et al.(Eds), Proceedings of the 6th international Coral Reef Symposium Townsville, Australia, pp. 803-810
Orton, J. H. 1914. On the Breeding Habits of Echinus miliaris, with a Note on the Feeding Habits of Patella Uulgata. Journal of the Marine Biological Association of the United Kingdom 10(2):254-257
Ouréns, R., L. Fernández, J. Freire. 2011. Geographic, population, and seasonal patterns in the reproductive parameters of the sea urchin Paracentrotus lividus 158:793-804
Palmer, L. 1937. The shedding reaction in Arbacia punctulata. Physiological Zoology 10(3):352-367
Pennington, J. T. 1985. The ecology of fertilization of echinoid egg: the consequences of sperm dilution, adult aggregation, and synchronous spawning. Biological Bulletin 169:417-430
Poorbagher, H., M. D. Lamare, and M. F. Barker. 2010. Effects of nutrition on somatic growth and reproductive strategy of the sea urchin Pseudechinus huttoni. Marine Biology Research, 2010; 6(3): 292-301
Rahman M. S., and T. Uehara. 2009. Effects of temperature on gametes aging and fertilization success of two sea urchin species. Zoological Science 26(1):1-8
Randall, J. E., R. E. Schroeder, and W. A. Starck II. 1964. Notes on the biology of the Echinoid Diadema antillarum. Caribbean Journal of Science 4:421-433
Refinetti R. 1993. Laboratory instrumentation and computing: comparison of six methods for the determination of the period of circadian rhythms. Physiology Behavior 54: 869–875
Sebens, K. P. 1987. The ecology of indeterminate growth in animals. Annual Review of Ecology and Systematics 18:371-407
Sewell, M. A., and C. M. Young. 1999. Temperature limits to fertilization and early development in the tropical sea urchin Echinometra lucunter. Journal of Experimental Marine Biology and Ecology 236:291-305
Silva, M. M., A. M. Albuquerque, and J. F. Araujo. 2005. Light-dark cycle synchronization of circadian rhythm in blind primates. Journal of Circadian Rhythms 3:10
Soong, K., and J. C. Lang. 1992. Reproductive Integration in Reef Corals. Biological Bulletin 183:418-431
Soong, K., J. Chen, C. J. Tsao. 2006. Adaptation for accuracy or for precision? Diel emergence timing of the intertidal insect Pontomyia oceana (Chironomidae) Marine Biology 150:173-181
Soong, K. , and T. Chen. 2003. Coral transplantation regeneration and growth of Acropora Fragments in a nursery. Restoration Ecology 11(1):62-71
Siikavuopio, S. I., and J. S. Christiansen, and T. Dale. 2006. Effects of temperature and season on gonad growth and feed intake in the green sea urchin (Strongylocentrotus droebachiensis). Aquaculture 255:389-394
Takashi, H. 1997. The annual reproductive cycle and lunar spawning rhythms of the purple sea urchin Anthocidaris crassispina. Nippon Suisan Gakkaishi 63(1):17-22
Takayuki, A. and U. Shimpei. 1998. Ecological distribution and body growth of Toxopneustes pileolus in the Acropora tumida Community in Suruga Bay, Japan. Bulletin of Institute of Oceanic Research and Development, Tokai University 19:71-78
Teruya, T., K. Suenaga, T. Koyama, Y. Nakano, and D. Uemura. 2001. Arachidonic acid and α-linolenic acid, feeding attractants for the crown-of-thorns sea star Acanthaster planci, from the sea urchin Toxopneustes pileolus. Journal of Experimental Marine Biology and Ecology 266:123-134
Tkachenko, K. S. and K. Soong.2010. Protection of Habitat Types: a case study of the effectiveness of a small marine reserve and impacts of different habitats on the diversity and abundance of coral reef fishes. Zoological Studies 49(2):195-210
Ullrich-Lüter E.M., S. Dupont, E. Arboleda, H. Hausen, M. I. Arnone. 2011. Unique system of photoreceptors in sea urchin tube feet. Proceedings of the National Academy of Sciences of the United States of America 108(20):8367-72
Valentine, J. P., and G. J. Edgar. 2010. Impacts of a population outbreak of the urchin Tripneustes gratilla amongst Lord Howe Island coral communities. Coral Reefs 29:399-410
Vazquez, L. C. 2003. Effect of sperm cell density on measured toxicity from the sea urchin Tripneustes gratilla fertilization bioassay. Environmental Toxicology and Chemistry 22(9):2191-2194
Veron Jen. 1995. Corals in space and time. The biogeography and evolution of the Scleractinia. University of New South Wales Press, Sydney, p 321
Willis, B. L., R. C. Babcock, P. L. Harrison, and J. K. Oliver. 1985. Patterns in the mass spawning of corals on the Great Barrier Reef from 1981 to 1984. Proceedings of the 5th international coral reef congress, Tahiti, 4:343-348
Werren, J. H., S.K. Skinner and A. Huger. 1986. Male-killing bacteria in a parasitic wasp. Science 231:990-992
Young, M. A. L., and D. R. Bellwood. 2011. Diel patterns in sea urchin activity and predation on sea urchins on the Great Barrier Reef. Coral Reefs 30:729-736