颱風是台灣最常見的自然擾動，颱風會造成大量的落葉，落葉會伴隨營養流失及光合作用效能下降。林木在大量落葉後，是否會藉由調整再吸收效率，來補償颱風所造成的營養損失？ 本研究自2010年8月進行至2010年12月，在福山試驗林一號集水區選擇山龍眼、長葉木薑子、綠樟與鋸葉長尾栲等四個樹種，追蹤分析颱風前後其成熟葉、落葉營養動態，以了解颱風如何影響林木再吸收效率的變動。研究期間有兩個中度颱風影響台灣，依次為9/19登陸的凡那比颱風與10月下旬接近台灣的梅姬颱風，兩個颱風皆帶來豪雨，但僅有凡那比颱風有較大風勢。研究結果顯示，兩個颱風對落葉量與再吸收效率造成的影響並不一致，9月的凡那比颱風對福山生態系帶來的影響大於10月的梅姬颱風。非颱風侵襲的11月有另一波落葉量高峰，山龍眼與長葉木薑子在9月凡那比侵襲期間落葉量較大，而綠樟與鋸葉長尾栲在11月的落葉量較大。在營養再吸收率方面，凡那比颱風造成N、K的再吸收效率下降，Ca、Mg再吸收效率上升，但對P再吸收率影響不大；另外梅姬颱風對營養再吸收效率沒有明顯的影響。追蹤颱風過後的數週，N再吸收效率沒有回升，顯示福山林木對應頻繁擾動，不會以耗費能量增加再吸收效率的方式來補償生長，以避免枝葉又受到不可預期擾動打落而失去更大量的營養。不同樹種在颱風與非颱風期間的落葉量有很大差異，且即使在非颱風期間亦有很大的差異，因此在討論颱風與非颱風期間的再吸收時，宜針對不同樹種分別採樣，才不會因物種間落葉量及再吸收的差異模糊了颱風對再吸收的影響。P再吸收效率與再吸收度皆高於N，顯示P是福山生態系的限制因子。

Typhoon is the most common natural disturbance in Taiwan. Typhoon disturbance often caused high quantity of leaflitter accompanied by the loss of nutrients and reduced photosynthesis. Will the damaged trees adjust nutrient retranslocation to enhance photosynthesis and as such compensate typhoon damage? We studied the pattern of retranslocation of Helicia formosana Hemsl、Litsea acuminata (Bl.) Kurata , Meliosma squamulata Hance , and Castanopsis cuspidata (Thunb. ex Murray) Schottky var. carlesii (Hemsl.)Yamazaki cf. sessilis (Nakai) Liao in Watershed #1 of Fushan Experimental Forest between August and December 2010 which includes period before and after typhoon disturbance. Two typhoons affected Taiwan during the study period, one is typhoon Fanapi landed Taiwan in 19 September 2010 and typhoon Megi approached Taiwan in late October 2010. Both typhoons brought large amount of precipitation but high winds only occurred when typhoon Fanapi affected Taiwan. The result indicates that the two typhoons affected Fushan Experimental Forest differently with typhoon Fanapi had a greater impact than typhoon Megi. There was another leaflitter peak in November unassociated with typhoon disturbance. Helicia formosana and Litsea acuminata had high quantity of leaflitter following typhoon Fanapi whereas leaflitter of Meliosma squamulata and Castanopsis cuspidata peaked in November. In terms of nutrient retranslocation, following typhoon Fanapi the retranslocation efficiency of N and K declined, the efficiency of Ca and Mg increased and the efficiency of P showed little change. No clear pattern of changes in retranslocation was observed following Typhoon Megi. Several weeks after typhoon Fanapi, N retranslocation efficiency did not return to levels before the typhoon. Possibly the energy consuming retranslocaiton is not preferred at Fushan where typhoon disturbance may occur several times a year as such investment in retranslocation may lead to large nutrient and energy loss. Tree species differed significantly in typhoon versus non-typhoon induced leaflitter and even in different collections in non-typhoon period. The result indicates that species-specific leaf-litter collection is required for a full understanding of typhoon effects on nutrient retranslocation. Both the retranslocaiton efficiency and retranslocation proficiency of P were greater than N indicating that P is a more limiting element at Fushan Ecosystem.

王雪卿。2009。台灣中部低海拔天然林與人工林枯落物、枯落物分解、土壤養分之比較。國立彰化師範大學地理學系碩士論文。
朱珮綺。2005。台大實驗林神木溪保護林兩相鄰林份枯落物動態及其養分之研究。國立台灣大學森林環境暨資源學系碩士論文。
林則桐、馬復京、張乃航。1995。福山試驗林的植物社會與天然更新之研究。林業試驗所百週年慶學術研討會論文集 71-82。
林國銓。1997。福山闊葉林落葉物及枝葉層之動態變化。台灣林業科學 12:135-144。
林國銓。2006。颱風干擾對福山天然闊葉林枯落物的影響。林業研究專訊 13:29-31。
林國銓、黃吳清標、劉哲政。1997。福山試驗林天然闊葉樹之物候現象。台灣林業科學12:347-353。
洪富文與程煒兒。1996。 福山闊葉林生態系的土壤養分庫及其有效養分動態。台灣林業科學 11:465-473。
張雅惠。2001。風對福山試驗林林木結構的影響。國立彰化師範大學地理學系碩士論文。
簡龍祥。2002。台灣東北部瑪鍊溪流域值群生態之研究。國立台灣大學森林學研究所資源保育組碩士論文。
劉育延。2006。福山三種地形主要樹種葉部養分濃度之季節變化。國立台灣大學森林環境暨資源學系碩士論文。
劉俊毅。2008。棲蘭山區臺灣扁柏老熟林及次生林枯落物養分動態。國立台灣大學森林環境暨資源學系碩士論文。
Aerts, R. 1995. The advantages of being evergreen. Trends in Ecology and Evolution 10: 402-407.
Aerts, R. 1996. Nutrient retranslocation from senescing leaves of perennials: are there general patterns? Journal of Ecology 84: 597-608.
Aerts, R. and F. S. Chapin Ⅲ. 2000. The mineral nutrition of wild plants revisited: a re-evaluation of processes and patterns. Advanced in Ecological Research 30.
Aide, T. M., J. K. Zimmerman, M. Rosario, and H. Marcano. 1996. Forest recovery in abandoned cattle pastures along an elevational gradient in northeastern Puerto Rico. Biotropica 28: 537-548.
Basnet, K., G. E. Likens, F. N. Scatena, and A. E. Lugo. 1992. Hurricane Hugo: damage to a tropical rainforest in Puterto Rico. Journal of Tropical Ecology 8: 47-55.
Berendse, F. and R. Aerts. 1987. Nitrogen use efficiency: a biologically meaningful definition? Functional Ecology 1: 293-296.
Birk, E. M. and P. M. Vitousek. 1986. Nitrogen availability and nitrogen use efficiency in loblolly pine stands. Ecology 67: 69-79.
Caldwell, M. M., J. H. Richards, D. A. Johnson, R. S. Nowak, and R. S. Dzurec. 1981. Coping with herbivory: photosynthetic capacity and resource allocation in two semiarid Agropyron bunchgrasses. Oecologia 50: 14-24.
Chapin, F. S. Ⅲ, D. A. Johnson, and J. D. McKendrick. 1980. Seasonal movement of nutrients in plants of differing growth form in an Alaskan tundra ecosystem: implications for herbivor. Journal of Ecology 68: 189-209.
Chapin, F. S. Ⅲ and R. A. Kedrowski. 1983. Seasonal changes in nitrogen and phosphorus fractions and autumn retranslocation in evergreen and deciduous taiga trees. Ecology 64: 376-391.
Cole, D. W. and M. Rapp. 1981. Elemental cycling in forest ecosystems. Reichle, D.E.(ed), dynamic properties of forest ecosystems,. pp.341-409. Cambridge university press, New York, New York, USA.
Craine, J. M. and M. C. Mack. 1998. Nutrients in senesced leaves: comment. Ecology 79: 1818-1820.
Feinsinger, P., Beach J. H., Y. B. Linhart, W. H.Busby, and K. G. Murray. 1987. Disturbance pollinator predictability, and pollination success among Costa Rican cloud forest plants. Ecology 68: 1294-1305.
Field, C. and H. A. Mooney 1986. The phptosynthesis-nitrogen relationship in wild plants. Givnish TJ.(ed), On the economy of plant form and function. pp. 25-49.Cambridge University Press, New York.
Fife, D. N. and E. K. S. Nambiar, and E. Saur. 2008. Retranslocation of foliar nutrients in evergreen tree species planted in a mediterranean environment. Tree Physiology 28: 187-196.
Fisher, S. G., N. B. Grimm, E. Marti, R. M. Holmes, and J. B. Jones Jr. 1998. Material spiraling in stream corridors: a telescoping ecosystem model. Ecosystems 1: 19-34.
Herbert, D. A., J. H. Fownes, and P. M. Vitousek. 1999. Hurricane damage to a Hawaiian forest: nutrient supply rate affects resistance and resilience. Ecology 80: 908-920.
Heichel, G. H. and N. C. Turner. 1983. CO2 assimilation of primary and regrowth foliage of red maple (Acer rubrum L.) and red oak (Quercus rubra L.): response to defoliation. Oecologia 57: 14-19.
Karban, R. and S. Y. Strauss. 1993. Effects of herbivores on growth and reproduction of their perennial host, Erigeron glaucus. Ecology 74: 39-46.
Killingbeck, K. T. 1996. Nutrients in Senesced Leaves: Keys to the Search for Potential Retranslocation and Retranslocation Proficiency. Ecology 77: 1716-1727.
Lin, T. C. and J. M. Chiang. 2002. Applications of hemispherical photographs in studies of forest ecology. Taiwan Journal of Forest Science 17: 387-400.
Lin, T. C., S. P. Hamburg, Y. J. Hsia, T. T. Lin, H. B. King, L. J. Wang, and K. C. Lin. 2003. Influence of typhoon disturbances on the understory light regime and stand dynamics of a subtropical rain forest in northeastern Taiwan. The Japanese Forest Society 8: 139-145.
Lin, T. C., S. P. Hamburg, K. C. Lin, L. J. Wang, C. T. Chang, Y. J. Hsia, M. A. Vadeboncoeur, C. M. M. McMullen, and C. P. Liu. 2010. Typhoon disturbance and forest dynamics: Lessons from a northwest pacific subtropical forest. Ecosystems 14: 127-143.
Lin, K. C., S. P. Hamburg, S. L. Tang, Y. J. Hsia, and T. C. Lin. 2003. Typhoon effects on litterfall in a subtropical forest. Canadian Journal of Forest Research 33: 2184-2192.
Lodge, D. J., F. N. Scatena, C. E. Asbury, and M. J. Sanchez. 1991. Fine litterfall and related nutrient inputs resulting from hurricane Hugo in subtropical wet and lower mountaine rain forests of Puerto Rico. Biotropica 23: 336-342.
Lodge, D. J. and W. H. McDowell. 1991. Summary of ecosystem-level effects of Caribbean hurricanes. Biotropica 23: 373-378.
Lovett , G. M. and P. Tobiessen. 1993. Carbon and nitrogen assimilation in red oaks (Quercus rubra L.) subject to defoliation and nitrogen stress. Tree Physiology 12: 259-269
Lovett, G. M., L. M. Christenson, P. M. Groffman, C. G. Jones, J. E. Hart, and M. J. Mitchell. 2002. Insect defoliation and nitrogen cycling in forests. Bioscience 52: 335-341.
Mabry, C. M., S. P. Hamburg, T. C. Lin, F. W. Horng, H. B. King, and Y. J. Hsia. 1998. Typhoon disturbance and stand-level damage patterns at a subtropical forest in Taiwan. Biotropica 30: 238-250.
May, J. D. and K. T. Killingbeck. 1992. Effects of Preventing Nutrient Retranslocation on Plant Fitness and Foliar Nutrient Dynamics. Ecology 73: 1868-1878.
McNaughton, S. J. 1983. Compensatory plant growth as a response to herbivory. Oikos 40: 329-336.
Miller, H. G. 1986. Carbon X nutrient interactions¬¬─the limitations to productivity. Tree Physiology 2: 373-385.
Nambiar, E. K. S. and D. N. Fife. 1987. Growth and nutrient retranslocation in needles of radiata pine in relation to nitrogen Supply. Annals of Botany 60: 147-156.
Nambiar, E. K. S. and D. N. Fife. 1991. Nutrient retranslocation in temperate conifers. Tree Physiology 9: 185-207.
Palma, R. M., R. L. Defrieri, M. F. Tortarolo, J. Prause. and J. F. Gallardo. 2000. Seasonal changes of bioelements in the litter and their potential return to green leaves in four Species of the Argentine subtropical forest. Annals of Botany 85: 181-186.
Perry, D. A., R. Oren, and S. C. Hart. 2008. Forest ecosystem 2nd edition. The Johns Hopkins University Press, Baltimore and London, USA.
Potter, D. A. and C. T. Redmond. 1989. Early spring defoliation, secondary leaf flush, and leafminer outbreaks on American holly. Oecologia 81: 192-197.
Reich, P. B., M. B. Walters, S. C. Krause, D. W. Vanderklein, K. F. Raffa, and T. Tabone. 1993. Growth, nutrition and gas exchange of Pinus resinosa following artificial defoliation. Trees 7: 67-77.
Robertson, G.P., D.C. Coleman, C.S. Bledsoe, and P. Sollins. 1999. Standard soil methods for long-term ecological research. Oxford University Press.

Sanford, R. L., Jr., W. J. Parton, D. S. Ojima, and D. J. Lodge. 1991. Hurricane effects on soil organic matter dynamics and forest production in the Luquillo experimental forest, Puetro Rico. Biotropica 23: 317-323.
Small, E. 1972. Photosynthetic rates in relation to nitrogen recycling as an adaptation to nutrient deficiency in peat bog plants. Canadian Journal of Botany 50: 2227-2233.
Thomas, H. and J. L. Stoddart. 1980. Leaf senescence. Annual Review of Plant Physiology 31: 83-111.
Toit, J. T. D., J. P. Bryant, and K. Frisby. 1990. Regrowth and palatability of Acacia shoots following pruning by African savanna browsers. Ecology 71: 149-154.
Ukonmaanaho, L. and M. Starr. 2001. The importance of leaching from litter collected in litterfall traps. Environmental Monitoring and Assessment 66: 129-146.
Vitousek, P. M. 1982. Nutrient cycling and nutrient use efficiency. The American Naturalist 119: 553-572.
Vitousek, P.M. 1984. Litterfall, nutrient cycling, and nutrient limitation in tropical forests. Ecology 65: 285-298.
Wright, I. J. and M. Westoby. 2003. Nutrient concentration, retranslocation and lifespan: leaf trait of Australian sclerophyll species. Functional Ecology 17: 10-19.
Xu, X., E. Hirata, and H. Shibata . 2004. Effect of typhoon disturbance on fine litterfall and related nutrient input in a subtropical forest on Okinawa Island. Japan. Basic and Applied Ecology 5: 271-282.
Xu, X. 2006. Nutrient dynamics in decomposing needles of Pinus luchuensis after typhoon disturbance in a subtropical environment. Annals of Forest Science 63: 707-713.