浮游生物悖論

在水生生物學中，浮游生物悖論（英語：paradox of the plankton）描述的是有限的資源卻意外地支持了大量浮游生物物種的情況，這顯然挑戰了競爭排除原則，即兩個物種競爭相同的資源時，有一個會佔劣勢而滅絕。

生態悖論

浮游生物悖論源於觀察到的浮游生物多樣性與競爭排除原則^[1]（也稱為高斯定律^[2]）之間的矛盾，後者認為，當兩個物種競爭相同的資源時，最終只有一個物種能夠存續，而另一個會滅絕。這樣的兩個物種之間是不可能共存的^[3]，因為占主導地位的物種將不可避免地耗盡共同的資源，從而壓制劣勢種。儘管浮游植物相互競爭的資源種類（例如光、硝酸鹽、磷酸鹽、硅酸、鐵）有限，但它們的生命在所有系統發生學水平上都是高度多樣的。

浮游生物悖論最初由喬治·伊夫林·哈欽森於1961年指出，他提出藉助光或湍流的垂直梯度、共生或偏利共生、差異捕食，或者環境條件不斷變化等因素，可以解決這一悖論。^[4]近來的文獻提出，考慮以下這些因素可以解決該悖論：混沌流體運動^[5]、尺度選擇性進食（size-selective grazing）^[6]、時空異質性^[7]和環境波動^[8]。更一般地說，一些研究人員認為，生態和環境因素不斷相互作用，使得浮游生物的棲息地永遠不會達到有利於單一物種的平衡。^[9]在Mitchell等人（2008）的論文中，研究人員發現，對浮游生物分布的小規模分析結果呈現出斑塊聚集的特徵，數量級為10 cm，它們存在的時間足夠長（>10分鐘），允許浮游生物進食、競爭和傳播。^[10]

參考文獻

^ Hardin, G. The Competitive Exclusion Principle. Science. 1960, 131 (3409): 1292–1297. Bibcode:1960Sci...131.1292H. PMID 14399717. doi:10.1126/science.131.3409.1292.
^ Gause, G. F. Experimental Studies on the Struggle for Existence - I. Mixed Population of Two Species of Yeast. Journal of Experimental Biology. 1932, 9: 389–402. doi:10.1242/jeb.9.4.389.
^ Johnson, Christopher A.; Bronstein, Judith L. Coexistence and competitive exclusion in mutualism. Ecology. 2019, 100 (6): e02708. ISSN 1939-9170. PMID 30924140. doi:10.1002/ecy.2708 （英語）.
^ Hutchinson, G. E. (1961) The paradox of the plankton. American Naturalist 95, 137-145.
^ Károlyi, G., Péntek, Á., Scheuring, I., Tél, T., Toroczkai, Z. (2000) Chaotic flow: the physics of species coexistence （頁面存檔備份，存於網際網路檔案館）. Proceedings of the National Academy of Sciences 97, 13661-13665.
^ Wiggert, J.D., Haskell, A.G.E., Paffenhofer, G.A., Hofmann, E.E. and Klinck, J.M. (2005) The role of feeding behavior in sustaining copepod populations in the tropical ocean 網際網路檔案館的存檔，存檔日期2008-09-05.. Journal of Plankton Research 27, 1013-1031.
^ Miyazaki, T., Tainaka, K., Togashi, T., Suzuki, T. and Yoshimura, J. (2006) Spatial coexistence of phytoplankton species in ecological timescale Archived copy. [2007-06-06]. （原始內容存檔於2007-09-27）. . Population Ecology 48(2), 107-112.
^ Descamps-Julien, B.; Gonzalez, A. Stable coexistence in a fluctuating environment: An experimental demonstration (PDF). Ecology. 2005, 86 (10): 2815–2824 [18 October 2014]. doi:10.1890/04-1700. （原始內容 (PDF)存檔於17 November 2006）.
^ Scheffer, M., Rinaldi, S., Huisman, J. and Weissing, F.J. (2003) Why plankton communities have no equilibrium: solutions to the paradox. Hydrobiologia 491, 9-18.
^ Mitchell, J.G., Yamazaki, H., Seuront, L., Wolk, F., Li, H. (2008) Phytoplankton patch patterns: Seascape anatomy in a turbulent ocean. Journal of Marine Systems 69, 247-253.

外部連結

The Paradox of the Plankton by Klaus Rohde

[1] Hardin, G. The Competitive Exclusion Principle. Science. 1960, 131 (3409): 1292–1297. Bibcode:1960Sci...131.1292H. PMID 14399717. doi:10.1126/science.131.3409.1292.

[2] Gause, G. F. Experimental Studies on the Struggle for Existence - I. Mixed Population of Two Species of Yeast. Journal of Experimental Biology. 1932, 9: 389–402. doi:10.1242/jeb.9.4.389.

[3] Johnson, Christopher A.; Bronstein, Judith L. Coexistence and competitive exclusion in mutualism. Ecology. 2019, 100 (6): e02708. ISSN 1939-9170. PMID 30924140. doi:10.1002/ecy.2708 （英語）.

[4] Hutchinson, G. E. (1961) The paradox of the plankton. American Naturalist 95, 137-145.

[5] Károlyi, G., Péntek, Á., Scheuring, I., Tél, T., Toroczkai, Z. (2000) Chaotic flow: the physics of species coexistence （頁面存檔備份，存於網際網路檔案館）. Proceedings of the National Academy of Sciences 97, 13661-13665.

[6] Wiggert, J.D., Haskell, A.G.E., Paffenhofer, G.A., Hofmann, E.E. and Klinck, J.M. (2005) The role of feeding behavior in sustaining copepod populations in the tropical ocean 網際網路檔案館的存檔，存檔日期2008-09-05.. Journal of Plankton Research 27, 1013-1031.

[7] Miyazaki, T., Tainaka, K., Togashi, T., Suzuki, T. and Yoshimura, J. (2006) Spatial coexistence of phytoplankton species in ecological timescale Archived copy. [2007-06-06]. （原始內容存檔於2007-09-27）. . Population Ecology 48(2), 107-112.

[8] Descamps-Julien, B.; Gonzalez, A. Stable coexistence in a fluctuating environment: An experimental demonstration (PDF). Ecology. 2005, 86 (10): 2815–2824 [18 October 2014]. doi:10.1890/04-1700. （原始內容 (PDF)存檔於17 November 2006）.

[9] Scheffer, M., Rinaldi, S., Huisman, J. and Weissing, F.J. (2003) Why plankton communities have no equilibrium: solutions to the paradox. Hydrobiologia 491, 9-18.

[10] Mitchell, J.G., Yamazaki, H., Seuront, L., Wolk, F., Li, H. (2008) Phytoplankton patch patterns: Seascape anatomy in a turbulent ocean. Journal of Marine Systems 69, 247-253.

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