Progresses of alpine treeline formation mechanism.

doi:10.13287/j.1001-9332.201609.010

Abstract

Abstract: Alpine treelines represent one of the most distinct vegetation boundaries between canopy closed montane forest and treeless alpine vegetation. This transitional ecotone is highly sensitive to global and regional climate change and is considered as an ideal indicator of such changes. Treeline studies have evolved from morphological description to various hypotheses of treeline formation. Although individual hypothesis may provide reasonable explanation locally, a generalized hypothesis that is applicable on the global scale is still lacking. Temperature is considered the limited factor controlling the distribution of alpine treeline as low temperature restricts biochemical processes of tree growth. However, which particular biochemical processes are affected by low temperature remains unknown. This paper summarized the mechanisms of treeline formation with a focus on how low temperature affects photosynthesis characteristics, nutrient characteristics, non-structural carbohydrate (NSC) and antioxidant system. We also reviewed the key issues and future perspectives in treeline research.

CONG Yu, HE Hong-shi, GU Xiao-nan, XU Wen-ru, LIU Kai, ZONG Sheng-wei, DU Hai-bo. Progresses of alpine treeline formation mechanism.[J]. Chinese Journal of Applied Ecology, 2016, 27(9): 3035-3041.

References

[1] Körner C. A re-assessment of high elevation treeline positions and their explanation. Oecologia, 1998, 115: 445-459
[2] Li M-H (李迈和), Kräuchi N. The state of knowledge on alpine treeline and suggestions for future research. Journal of Sichuan Forestry Science and Technology (四川林业科技), 2005, 26(4): 36-42 (in Chinese)
[3] Song H-T (宋洪涛), Cheng S (程颂), Sun S-Q (孙守琴). Hypotheses of alpine timberline formation mechanism: A review. Chinese Journal of Ecology (生态学杂志), 2009, 28(11): 2393-2402 (in Chinese)
[4] Körner C, Paulsen J. A world-wide study of high altitude treeline temperatures. Journal of Biogeography, 2004, 31: 713-732
[5] Larcher W. Winter stress in high mountains// Turner H, Tranquillini W, eds. Establishment and Tending of Subalpine Forest: Research and Management. Birmensdorf: Swiss Federal Institute of Forestry Research, 1985: 258
[6] Sakai A, Larcher W. Frost Survival of Plants: Responses and Adaptation to Freezing Stress. Berlin, Heidelberg: Springer Science & Business Media, 1987
[7] Fang J-Q (方近圻), Wu N (吴宁), Luo P (罗鹏), et al. Impacts of distances on alpine timberline reformation. Chinese Journal of Ecology (生态学杂志), 2005, 24(12): 1493-1498
[8] Tranquillini W. Physiological Ecology of the Alpine Timberline. New York: Springer-Verlag, 1979
[9] Wardle P. Is the alpine timberline set by physiological tolerance, reproductive capacity, or biological interactions. Proceedings of the Ecological Society of Australia,1981, 11: 53-66
[10] Sveinbjörnsson B, Kauhanen H, Nordell O. Treeline ecology of mountain birch in the Torneträsk area. Ecolo-gical Bulletins, 1996, 45: 65-70
[11] Bliss LC. Alpine. Amsterdam, The Netherlands: Sprin-ger, 1985: 41-65
[12] Barrick KA, Schoettle AW. A comparison of the foliar nutrient status of elfinwood and symmetrically formed tall trees, Colorado Front Range, USA. Canadian Journal of Botany, 1996, 74: 1461-1475
[13] Richardson AD. Foliar chemistry of balsam fir and red spruce in relation to elevation and the canopy light gra-dient in the mountains of the northeastern United States. Plant and Soil, 2004, 260: 291-299
[14] Stevens GC, Fox JF. The causes of treeline. Annual Review of Ecology and Systematics, 1991, 22: 177-191
[15] Grace J. Cuticular water loss unlikely to explain tree-line in Scotland. Oecologia, 1990, 84: 64-68
[16] Yu D, Wang Q, Liu J, et al. Formation mechanisms of the alpine Erman’s birch (Betula ermanii) treeline on Changbai Mountain in Northeast China. Trees, 2014, 28: 935-947
[17] Takahashi K. Effects of wind and thermal conditions on timberline formation in central Japan: A lattice model. Ecological Research, 2014, 29: 121-131
[18] Körner C. Alpine Plant Life: Functional Plant Ecology of High Mountain Ecosystems. Berlin: Springer, 1999
[19] Jobbagy EG, Jackson RB. Global controls of forest line elevation in the northern and southern hemispheres. Global Ecology and Biogeography, 2000, 9: 253-268
[20] Shi P, Körner C, Hoch G. A test of the growth-limitation theory for alpine tree line formation in evergreen and deciduous taxa of the eastern Himalayas. Functional Ecology, 2008, 22: 213-220
[21] James JC, Grace J, Hoad SP. Growth and photosynthesis of Pinus sylvestris at its altitudinal limit in Scotland. Journal of Ecology, 1994: 297-306
[22] Körner C, Diemer M. In situ photosynthetic responses to light, temperature and carbon dioxide in herbaceous plants from low and high altitude. Functional Ecology, 1987: 179-194
[23] Liu H-Y (刘鸿雁), Gu H-T (谷洪涛), Tang Z-Y (唐志尧), et al. Tree photosynthesis on alpine timberline and its relationships to environmental factors in the eastern part of temperate China. Journal of Mountain Science (山地学报), 2002, 20(1): 32-36 (in Chinese)
[24] Wang X-C (王晓春), Zhou X-F (周晓峰), Sun Z-H (孙志虎). Research advances in the relationship between alpine timberline and climate change. Chinese Journal of Ecology (生态学杂志), 2005, 24(3): 301-305 (in Chinese)
[25] Zhu W-Z (朱万泽), Wu Y-B (吴永波), Xue J-H (薛建辉). Photosynthetic characteristics of Quercus pannosa in Gongga Mountain Region. Journal of Nanjing Forestry University (Natural Science) (南京林业大学学报: 自然科学版), 2006, 30(1): 25-28 (in Chinese)
[26] Tranquillini W, Havranek WM, Ecker P. Effects of atmospheric humidity and acclimation temperature on the temperature response of photosynthesis in young Larix decidua Mill. Tree Physiology, 1986, 1: 37-45
[27] Chapin FS. The mineral nutrition of wild plants. Annual Review of Ecology and Systematics, 1980, 11: 233-260
[28] Reich PB, Walters MB, Ellsworth DS. From tropics to tundra: Global convergence in plant functioning. Proceedings of the National Academy of Sciences of the Uni-ted States of America, 1997, 94: 13730-13734
[29] Körner C. Carbon limitation in trees. Journal of Ecology, 2003, 91: 4-17
[30] Thébault A, Clément JC, Ibanez S, et al. Nitrogen limitation and microbial diversity at the treeline. Oikos, 2014, 123: 729-740
[31] Koerselman W, Meuleman AF. The vegetation N:P ratio: A new tool to detect the nature of nutrient limitation. Journal of Applied Ecology, 1996: 1441-1450
[32] Güsewell S. N:P ratios in terrestrial plants: Variation and functional significance. New Phytologist, 2004, 164: 243-266
[33] Reich PB, Oleksyn J. Global patterns of plant leaf N and P in relation to temperature and latitude. Procee-dings of the National Academy of Sciences of the United States of America, 2004, 101: 11001-11006
[34] Oleksyninst J, Reich PB, Zytkowiak R, et al. Needle nutrients in geographically diverse Pinus sylvestris L. populations. Annals of Forest Science, 2002, 59: 1-18
[35] Loomis PF, Ruess RW, Sveeinbjörnsson B, et al. Nitrogen cycling at treeline: Latitudinal and elevational patterns across a boreal landscape. Ecoscience, 2006, 13: 544-556
[36] Li MH, Xiao WF, Shi P, et al. Nitrogen and carbon source-sink relationships in trees at the Himalayan treelines compared with lower elevations. Plant, Cell & Environment, 2008, 31: 1377-1387
[37] Farquhar G, von Caemmerer S, Berry J. A biochemical model of photosynthetic CO₂ assimilation in leaves of C3 species. Planta, 1980, 149: 78-90
[38] Ögren E, Nilsson T, Sundblad LG. Relationship between respiratory depletion of sugars and loss of cold hardiness in coniferous seedlings over-wintering at raised temperatures: Indications of different sensitivities of spruce and pine. Plant, Cell & Environment, 1997, 20: 247-253
[39] Ögren E. Maintenance respiration correlates with sugar but not nitrogen concentration in dormant plants. Physiologia Plantarum, 2000, 108: 295-299
[40] Li M, Hoch G, Körner C. Source/sink removal affects mobile carbohydrates in Pinus cembra at the Swiss treeline. Trees, 2002, 16: 331-337
[41] Li MH, Xiao WF, Wang SG, et al. Mobile carbohydrates in Himalayan treeline trees. Ⅰ. Evidence for carbon gain limitation but not for growth limitation. Tree Physiology, 2008, 28: 1287-1296
[42] Pan Q-M (潘庆民), Han X-G (韩兴国), Bai Y-F (白永飞), et al. Advances in physiology and ecology studies on stored non-structure carbohydrates in plants. Chinese Bulletin of Botany (植物学通报), 2002, 19(1): 30-38 (in Chinese)
[43] Koch K. Carbohydrate-modulated gene expression in plants. Annual Review of Plant Biology, 1996, 47: 509-540
[44] Martin M, Miceli F, Morgan J, et al. Synthesis of osmotically active substances in winter wheat leaves as related to drought resistance of different genotypes. Journal of Agronomy and Crop Science, 1993, 171: 176-184
[45] Sheen J, Zhou L, Jang JC. Sugars as signaling molecules. Current Opinion in Plant Biology, 1999, 2: 410-418
[46] Morin X, Améglio T, Ahas R, et al. Variation in cold hardiness and carbohydrate concentration from dormancy induction to bud burst among provenances of three European oak species. Tree Physiology, 2007, 27: 817-825
[47] Sakai A, Yoshida S. The role of sugar and related compounds in variations of freezing resistance. Cryobiology, 1968, 5: 160-174
[48] Kaurin A, Junttila O, Hanson J. Seasonal changes in frost hardiness in cloudberry (Rubus chamaemorus) in relation to carbohydrate content with special reference to sucrose. Physiologia Plantarum, 1981, 52: 310-314
[49] Zhu WZ, Cao M, Wang SG, et al. Seasonal dynamics of mobile carbon supply in Quercus aquifolioides at the upper elevational limit. PLoS One, 2012, 7: e34213
[50] Würth MK, Pelaez-Riedl S, Wright SJ, et al. Non-structural carbohydrate pools in a tropical forest. Oecologia, 2005, 143: 11-24
[51] Hoch G, Körner C. The carbon charging of pines at the climatic treeline: A global comparison. Oecologia, 2003, 135: 10-21
[52] Hoch G, Popp M, Körner C. Altitudinal increase of mobile carbon pools in Pinus cembra suggests sink limitation of growth at the Swiss treeline. Oikos, 2002, 98: 361-374
[53] Sala A, Woodruff DR, Meinzer FC. Carbon dynamics in trees: Feast or famine? Tree Physiology, 2012, 32: 764-775
[54] Cavieres LA, Rada F, Azócar A, et al. Gas exchange and low temperature resistance in two tropical high mountain tree species from the Venezuelan Andes. Acta Oecologica, 2000, 21: 203-211
[55] Bacelar EA, Santos DL, Moutinho-Pereira JM, et al. Immediate responses and adaptative strategies of three olive cultivars under contrasting water availability regimes: Changes on structure and chemical composition of foliage and oxidative damage. Plant Science, 2006, 170: 596-605
[56] Runion G, Entry J, Prior S, et al. Tissue chemistry and carbon allocation in seedlings of Pinus palustris subjected to elevated atmospheric CO₂ and water stress. Tree Physiology, 1999, 19: 329-335
[57] Li N, He N, Yu G, et al. Leaf non-structural carbohydrates regulated by plant functional groups and climate: Evidences from a tropical to cold-temperate forest transect. Ecological Indicators, 2016, 62: 22-31
[58] Poorter L, Kitajima K. Carbohydrate storage and light requirements of tropical moist and dry forest tree species. Ecology, 2007, 88: 1000-1011
[59] Asao S, Ryan MG. Carbohydrate regulation of photosynthesis and respiration from branch girdling in four species of wet tropical rain forest trees. Tree Physiology, 2015, 35: 608-620
[60] Wang S-G (王三根) . Plant Physiology and Biochemistry. Beijing: China Agriculture Press, 2013 (in Chinese)
[61] Zhang T (张涛), An L-Z (安黎哲), Chen T (陈拓), et al. Antioxidative system in leaves of Picea crassifolia and Sabina przewalskii along an altitudinal gra-dient. Chinese Journal of Plant Ecology (植物生态学报), 2009, 33(4): 802-811 (in Chinese)
[62] Holmberg N, Bülow L. Improving stress tolerance in plants by gene transfer. Trends in Plant Science, 1998, 3: 61-66
[63] Swanson S, Gilroy S. ROS in plant development. Physiologia Plantarum, 2010, 138: 384-392
[64] Jaspers P, Kangasjärvi J. Reactive oxygen species in abiotic stress signaling. Physiologia Plantarum, 2010, 138: 405-413
[65] Joo JH, Yoo HJ, Hwang I, et al. Auxin-induced reactive oxygen species production requires the activation of phosphatidylinositol 3-kinase. Febs Letters, 2005, 579: 1243-1248
[66] Müller K, Linkies A, Vreeburg RA, et al. In vivo cell wall loosening by hydroxyl radicals during cress seed germination and elongation growth. Plant Physiology, 2009, 150: 1855-1865
[67] Shi SQ, Shi Z, Jiang ZP, et al. Effects of exogenous GABA on gene expression of Caragana intermedia roots under NaCl stress: Regulatory roles for H₂O₂ and ethy-lene production. Plant, Cell & Environment, 2010, 33: 149-162
[68] Mur LA, Kenton P, Lloyd AJ, et al. The hypersensitive response: The centenary is upon us but how much do we know? Journal of Experimental Botany, 2008, 59: 501-520
[69] Kang H-J (康华靖), Liu P (刘鹏), Xu G-D (徐根娣), et al. Ecophysiological response of Emmenopterys henryi at different altitudes in China’s Dapan Mountain natural reserve. Chinese Journal of Plant Ecology (植物生态学报), 2008, 32(4): 865-872 (in Chinese)
[70] Yang S-S (杨淑慎), Gao J-F (高俊凤). Influence of active oxygen and free radicals on plant senescence. Acta Botanica Boreali-Occidentalia Sinica (西北植物学报), 2001, 21(2): 215-220 (in Chinese)
[71] Gechev T, Willekens H, van Montagu M, et al. Diffe-rent responses of tobacco antioxidant enzymes to light and chilling stress. Journal of Plant Physiology, 2003, 160: 509-515
[72] Hu Q-P (胡启鹏), Guo Z-H (郭志华), Sun L-L (孙玲玲), et al. Response of leaf functional traits of Betula ermanii saplings to the altitudinal variation. Acta Ecolo-gica Sinica (生态学报), 2013, 33(12): 3594-3601 (in Chinese)
[73] Liang J-P (梁建萍), Niu Y (牛远), Xie J-S (谢敬斯), et al. Antioxidase activities and photosynthetic pigment contents in Larix principis-rupprechtii leaves along an altitudinal gradient. Chinese Journal of Applied Ecology (应用生态学报), 2007, 18(7): 1414-1419 (in Chinese)