巴郎山糙皮桦叶片光合氮利用效率的海拔响应
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摘要
为深入认识植物对环境变化的响应和适应,以分布在川西巴郎山的糙皮桦为研究对象,选择海拔2200、2500、3100和3400 m 4个分布点,测定计算了各分布点叶片光合氮利用效率(PNUE)、CO_2扩散导度(叶肉细胞导度g_m与气孔导度g_s)和氮分配比例(Rubisco氮分配比例P_R、生物力能学组分氮分配比例P_B、捕光组分氮分配比例P_L与细胞壁氮分配比例P_(CW))等参数,分析了其沿海拔的变化趋势以及叶片PNUE与其他参数的相关关系.结果表明:糙皮桦叶片PNUE、P_R和P_B在海拔2500和3100 m相对较高;叶片g_s和g_m则随海拔升高而增加,P_L随海拔升高而降低.糙皮桦叶片P_R和P_B与PNUE呈显著正相关关系,说明P_R和P_B是PNUE随海拔变异的重要内部因素.糙皮桦叶片光合系统氮分配比例P_P在海拔2500和3100 m相对较高,叶片P_(CW)随海拔升高而降低,叶片其他组分氮分配比例P_(other)随海拔升高而增加,说明随海拔的升高,糙皮桦叶片趋向将更大比例的氮分配于除光合系统和细胞壁外的其他组分中.
To better understand the response and adaptation of plants to altitudinal changes, four sites at the altitude of 2200 m, 2500 m, 3100 m and 3400 m on Balang Mountain were selected to test and calculate the eco-physiological parameters in leaves of Betula utilis, including photosynthetic nitrogen use efficiency(PNUE), CO_2 diffusion conductance(stomatal conductance g_s and mesophyll conductance g_m) and nitrogen allocation in each component(fractions of leaf nitrogen allocated to Rubisco P_R, to bioenergetics P_B, to light-harvesting components P_L, and to cell wall P_(CW)). Their changes with altitudinal variations and the relationships between leaf PNUE and the other parameters were analyzed. The results showed that PNUE, P_R, and P_B of the leaves were relatively higher at 2500 m and 3100 m. With the increases of altitude, g_s and g_m increased and P_L decreased. The correlations between P_R,_P_B and PNUE were significant, indicating that P_R and P_B were the main factors driving the changes in leaf PNUE in response to altitudinal variations. Besides, the fraction of leaf nitrogen allocated to photosynthetic apparatus(P_P) was relatively higher at 2500 m and 3100 m. With increasing altitude, P_(CW) decreased and the fraction of leaf nitrogen allocated to the other components(P_(other)) increased, which suggested that B. utilis leaves tended to allocate more nitrogen to the other components instead of the photosynthetic apparatus and cell wall with the increasing altitude to well adapt environmental changes.
To better understand the response and adaptation of plants to altitudinal changes, four sites at the altitude of 2200 m, 2500 m, 3100 m and 3400 m on Balang Mountain were selected to test and calculate the eco-physiological parameters in leaves of Betula utilis, including photosynthetic nitrogen use efficiency(PNUE), CO_2 diffusion conductance(stomatal conductance g_s and mesophyll conductance g_m) and nitrogen allocation in each component(fractions of leaf nitrogen allocated to Rubisco P_R, to bioenergetics P_B, to light-harvesting components P_L, and to cell wall P_(CW)). Their changes with altitudinal variations and the relationships between leaf PNUE and the other parameters were analyzed. The results showed that PNUE, P_R, and P_B of the leaves were relatively higher at 2500 m and 3100 m. With the increases of altitude, g_s and g_m increased and P_L decreased. The correlations between P_R,_P_B and PNUE were significant, indicating that P_R and P_B were the main factors driving the changes in leaf PNUE in response to altitudinal variations. Besides, the fraction of leaf nitrogen allocated to photosynthetic apparatus(P_P) was relatively higher at 2500 m and 3100 m. With increasing altitude, P_(CW) decreased and the fraction of leaf nitrogen allocated to the other components(P_(other)) increased, which suggested that B. utilis leaves tended to allocate more nitrogen to the other components instead of the photosynthetic apparatus and cell wall with the increasing altitude to well adapt environmental changes.
引文
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[2] Field C, Mooney HA. The photosynthesis-nitrogen relationship in wild plants// Givnish TJ, ed. On the Economy of Form and Function. Cambridge, UK: Cambridge University Press, 1986: 25-55
[3] Poorter H, Evans JR. Photosynthetic nitrogen-use efficiency of species that differ inherently in specific leaf area. Oecologia, 1998, 116: 26-37
[4] Evans JR, Seemann JR. The allocation of protein nitrogen in the photosynthetic apparatus: Costs, consequences, and control// Briggs WR, ed. Photosynthesis. New York, USA: Alan R. Liss Inc., 1989: 183-205
[5] Iqbal N, Umar S, Per TS, et al. Ethephon increases photosynthetic-nitrogen use efficiency, proline and antioxidant metabolism to alleviate decrease in photosynthesis under salinity stress in mustard. Plant Signaling & Behavior, 2017, 12: e1297000, doi: 10.1080/15592324.2017.1297000
[6] Hikosaka K. Interspecific difference in the photosynthesis-nitrogen relationship: Patterns, physiological causes, and ecological importance. Journal of Plant Research, 2004, 117: 481-494
[7] Hikosaka K. Mechanisms underlying interspecific variation in photosynthetic capacity across wild plant species. Plant Biotechnology, 2010, 27: 223-229
[8] Tang JC, Cheng RM, Shi ZM, et al. Fagaceae tree species allocate higher fraction of nitrogen to photosynthetic apparatus than Leguminosae in Jianfengling tropical montane rain forest, China. PLoS One, 2018, 13(2): e0192040
[9] Tosens T, Nishida K, Gago J, et al. The photosynthetic capacity in 35 ferns and fern allies: Mesophyll CO2 diffusion as a key trait. New Phytologist, 2016, 209: 1576-1590
[10] K?rner C. The use of ‘altitude’ in ecological research. Trends in Ecology & Evolution, 2007, 22: 569-574
[11] Cordell S, Goldstein G, Meinzer FC, et al. Allocation of nitrogen and carbon in leaves of Metrosideros polymorpha regulates carboxylation capacity and deltaC-13 along an altitudinal gradient. Functional Ecology, 1999, 13: 811-818
[12] Ran F, Zhang XL, Zhang YB, et al. Altitudinal variation in growth, photosynthetic capacity and water use efficiency of Abies faxoniana Rehd. et Wils. seedlings as revealed by reciprocal transplantations. Trees-Structure and Function, 2013, 27: 1405-1416
[13] Terashima I, Masuzawa T, Ohba H. Photosynthetic characteristics of a giant alpine plant, Rheum nobile Hook. f. et Thoms. and of some other alpine species measured at 4300 m, in the Eastern Himalaya, Nepal. Oecologia, 1993, 95: 194-201
[14] Vats SK, Kumar N, Kumar S. Gas exchange response of barley and pea cultivars to altitude variation in Himalaya. Photosynthetica, 2009, 47: 41-45
[15] Shi ZM, Liu SR, Liu XL, et al. Altitudinal variation in photosynthetic capacity, diffusional conductance and delta C-13 of butterfly bush (Buddleja davidii) plants growing at high elevations. Physiologia Plantarum, 2006, 128: 722-731
[16] Feng Q-H (冯秋红), Cheng R-M (程瑞梅), Shi Z-M (史作民), et al. Response of leaf functional traits and the relationships among them to altitude of Salix dissa in Balang Mountain. Acta Ecologica Sinica (生态学报), 2013, 33(9): 2712-2718 (in Chinese)
[17] Feng QH, Mauro C, Cheng RM, et al. Leaf functional trait responses of Quercus aquifolioides to high elevations. International Journal of Agriculture & Biology, 2013, 15: 69-75
[18] Bahar NHA, Ishida FY, Weerasinghe LK, et al. Leaf-level photosynthetic capacity in lowland Amazonian and high-elevation Andean tropical moist forests of Peru. New Phytologist, 2017, 214: 1002-1018
[19] Feng Q-H (冯秋红), Cheng R-M (程瑞梅), Shi Z-M (史作民), et al. Response of Rumex dentatus foliar nitrogen and its allocation to altitudinal gradients along Balang Mountain, Sichuan, China. Chinese Journal of Plant Ecology (植物生态学报), 2013, 37(7): 591-600 (in Chinese)
[20] Feng Q-H (冯秋红), Cheng R-M (程瑞梅), Shi Z-M (史作民), et al. Effects of altitudinal gradient on Salix atopantha foliar δ13C. Chinese Journal of Applied Ecology (应用生态学报), 2011, 22(11): 2841-2848 (in Chinese)
[21] Onoda Y, Hikosaka K, Hirose T. Allocation of nitrogen to cell walls decreases photosynthetic nitrogen-use efficiency. Functional Ecology, 2004, 18: 419-425
[22] Li D-X (李得孝), Guo Y-X (郭月霞), Yuan H-Y (员海燕), et al. Determined methods of chlorophyll from maize. Chinese Agricultural Science Bulletin (中国农学通报), 2005, 21(6): 153-155 (in Chinese)
[23] Sharkey TD. What gas exchange data can tell us about photosynthesis. Plant, Cell & Environment, 2016, 39: 1161-1163
[24] Niinemets U, Tenhunen JD. A model separating leaf structural and physiological effects on carbon gain along light gradients for the shade-tolerant species Acer saccharum. Plant, Cell & Environment, 1997, 20: 845-866
[25] Zhang SB, Zhou ZK, Hu H, et al. Gas exchange and resource utilization in two alpine oaks at different altitudes in the Hengduan Mountains. Canadian Journal of Forest Research, 2007, 37: 1184-1193
[26] Eichelmann H, Laisk A. Ribulose-1,5-bisphosphate carboxylase/oxygenase content, assimilatory charge, and mesophyll conductance in leaves. Plant Physiology, 1999, 119: 179-190
[27] Li Y, Ren BB, Ding L, et al. Does chloroplast size influence photosynthetic nitrogen use efficiency? PLoS One, 2013, 8(4): e62036
[28] Benomar L, Lamhamedi MS, Villeneuve I, et al. Fine-scale geographic variation in photosynthetic-related traits of Picea glauca seedlings indicates local adaptation to climate. Tree Physiology, 2015, 35: 864-878
[29] Harley PC, Loreto F, Dimarco G, et al. Theoretical considerations when estimating the mesophyll conduc-tance to CO2 flux by analysis of the response of photosynthesis to CO2. Plant Physiology, 1992, 98: 1429-1436
[30] Niinemets ü, Díaz-Espejo A, Flexas J. Role of mesophyll diffusion conductance in constraining potential photosynthetic productivity in the field. Journal of Expe-rimental Botany, 2009, 60: 2249-2270
[31] Bown HE, Watt MS, Mason EG, et al. The influence of nitrogen and phosphorus supply and genotype on mesophyll conductance limitations to photosynthesis in Pinus radiata. Tree Physiology, 2009, 29: 1143-1151
[32] Flexas J, Niinemets ü, Gallé A, et al. Diffusional conductances to CO2 as a target for increasing photosynthesis and photosynthetic water-use efficiency. Photosynthesis Research, 2013, 117: 45-59
[33] Gale J. Availability of carbon dioxide for photosynthesis at high altitudes: Theoretical considerations. Ecology, 1972, 53: 494-497
[34] Woodward FI, Bazzaz FA. The responses of stomatal density to CO2 partial pressure. Journal of Experimental Botany, 1988, 39: 1771-1781
[35] Nobel PS, Zaragoza LJ, Smith WK. Relation between mesophyll surface-area, photosynthetic rate, and illumination level during development for leaves of Plectranthus-parviflorus Henckel. Plant Physiology, 1975, 55: 1067-1070
[36] Long SP, Humphries S, Falkowski PG. Photoinhibition of photosynthesis in nature. Annual Review of Plant Physiology and Plant Molecular Biology, 1994, 45: 633-662
[37] Li H-M (李惠梅), Shi S-B (师生波). Photosynthetic characteristics of Gentiana straminea at different altitudes. Journal of Anhui Agricultural Sciences (安徽农业科学), 2008, 36(11): 4799-4804 (in Chinese)
[38] Lu W-M (卢文敏), Liu W-G (刘伟国), Fang X-Y (方晓雨), et al. Spectral reflectance of Betula ermanii at different altitudes in the Changbai mountains. Journal of Beijing Forestry University (北京林业大学学报), 2011, 33(1): 55-59 (in Chinese)
[39] 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)
[40] Shi Z-M (史作民), Tang J-C (唐敬超), Cheng R-M (程瑞梅), et al. A review of nitrogen allocation in leaves and factors in its effects. Acta Ecologica Sinica (生态学报), 2015, 35(18): 5909-5919 (in Chinese)
[41] Lei YB, Chen K, Hao J, et al. Contrasting responses in the growth and energy utilization properties of sympatric Populus and Salix to different altitudes: Implications for sexual dimorphism in Salicaceae. Physiologia Plantarum, 2017, 159: 30-41
[42] González-Zurdo P, Escudero A, Babiano J, et al. Costs of leaf reinforcement in response to winter cold in evergreen species. Tree Physiology, 2016, 36: 273-286
[43] Onoda Y, Wright IJ, Evans JR, et al. Physiological and structural tradeoffs underlying the leaf economics spectrum. New Phytologist, 2017, 214: 1447-1463
[44] Harrison MT, Edwards EJ, Farquhar GD, et al. Nitrogen in cell walls of sclerophyllous leaves accounts for little of the variation in photosynthetic nitrogen-use efficiency. Plant, Cell & Environment, 2009, 32: 259-270
[2] Field C, Mooney HA. The photosynthesis-nitrogen relationship in wild plants// Givnish TJ, ed. On the Economy of Form and Function. Cambridge, UK: Cambridge University Press, 1986: 25-55
[3] Poorter H, Evans JR. Photosynthetic nitrogen-use efficiency of species that differ inherently in specific leaf area. Oecologia, 1998, 116: 26-37
[4] Evans JR, Seemann JR. The allocation of protein nitrogen in the photosynthetic apparatus: Costs, consequences, and control// Briggs WR, ed. Photosynthesis. New York, USA: Alan R. Liss Inc., 1989: 183-205
[5] Iqbal N, Umar S, Per TS, et al. Ethephon increases photosynthetic-nitrogen use efficiency, proline and antioxidant metabolism to alleviate decrease in photosynthesis under salinity stress in mustard. Plant Signaling & Behavior, 2017, 12: e1297000, doi: 10.1080/15592324.2017.1297000
[6] Hikosaka K. Interspecific difference in the photosynthesis-nitrogen relationship: Patterns, physiological causes, and ecological importance. Journal of Plant Research, 2004, 117: 481-494
[7] Hikosaka K. Mechanisms underlying interspecific variation in photosynthetic capacity across wild plant species. Plant Biotechnology, 2010, 27: 223-229
[8] Tang JC, Cheng RM, Shi ZM, et al. Fagaceae tree species allocate higher fraction of nitrogen to photosynthetic apparatus than Leguminosae in Jianfengling tropical montane rain forest, China. PLoS One, 2018, 13(2): e0192040
[9] Tosens T, Nishida K, Gago J, et al. The photosynthetic capacity in 35 ferns and fern allies: Mesophyll CO2 diffusion as a key trait. New Phytologist, 2016, 209: 1576-1590
[10] K?rner C. The use of ‘altitude’ in ecological research. Trends in Ecology & Evolution, 2007, 22: 569-574
[11] Cordell S, Goldstein G, Meinzer FC, et al. Allocation of nitrogen and carbon in leaves of Metrosideros polymorpha regulates carboxylation capacity and deltaC-13 along an altitudinal gradient. Functional Ecology, 1999, 13: 811-818
[12] Ran F, Zhang XL, Zhang YB, et al. Altitudinal variation in growth, photosynthetic capacity and water use efficiency of Abies faxoniana Rehd. et Wils. seedlings as revealed by reciprocal transplantations. Trees-Structure and Function, 2013, 27: 1405-1416
[13] Terashima I, Masuzawa T, Ohba H. Photosynthetic characteristics of a giant alpine plant, Rheum nobile Hook. f. et Thoms. and of some other alpine species measured at 4300 m, in the Eastern Himalaya, Nepal. Oecologia, 1993, 95: 194-201
[14] Vats SK, Kumar N, Kumar S. Gas exchange response of barley and pea cultivars to altitude variation in Himalaya. Photosynthetica, 2009, 47: 41-45
[15] Shi ZM, Liu SR, Liu XL, et al. Altitudinal variation in photosynthetic capacity, diffusional conductance and delta C-13 of butterfly bush (Buddleja davidii) plants growing at high elevations. Physiologia Plantarum, 2006, 128: 722-731
[16] Feng Q-H (冯秋红), Cheng R-M (程瑞梅), Shi Z-M (史作民), et al. Response of leaf functional traits and the relationships among them to altitude of Salix dissa in Balang Mountain. Acta Ecologica Sinica (生态学报), 2013, 33(9): 2712-2718 (in Chinese)
[17] Feng QH, Mauro C, Cheng RM, et al. Leaf functional trait responses of Quercus aquifolioides to high elevations. International Journal of Agriculture & Biology, 2013, 15: 69-75
[18] Bahar NHA, Ishida FY, Weerasinghe LK, et al. Leaf-level photosynthetic capacity in lowland Amazonian and high-elevation Andean tropical moist forests of Peru. New Phytologist, 2017, 214: 1002-1018
[19] Feng Q-H (冯秋红), Cheng R-M (程瑞梅), Shi Z-M (史作民), et al. Response of Rumex dentatus foliar nitrogen and its allocation to altitudinal gradients along Balang Mountain, Sichuan, China. Chinese Journal of Plant Ecology (植物生态学报), 2013, 37(7): 591-600 (in Chinese)
[20] Feng Q-H (冯秋红), Cheng R-M (程瑞梅), Shi Z-M (史作民), et al. Effects of altitudinal gradient on Salix atopantha foliar δ13C. Chinese Journal of Applied Ecology (应用生态学报), 2011, 22(11): 2841-2848 (in Chinese)
[21] Onoda Y, Hikosaka K, Hirose T. Allocation of nitrogen to cell walls decreases photosynthetic nitrogen-use efficiency. Functional Ecology, 2004, 18: 419-425
[22] Li D-X (李得孝), Guo Y-X (郭月霞), Yuan H-Y (员海燕), et al. Determined methods of chlorophyll from maize. Chinese Agricultural Science Bulletin (中国农学通报), 2005, 21(6): 153-155 (in Chinese)
[23] Sharkey TD. What gas exchange data can tell us about photosynthesis. Plant, Cell & Environment, 2016, 39: 1161-1163
[24] Niinemets U, Tenhunen JD. A model separating leaf structural and physiological effects on carbon gain along light gradients for the shade-tolerant species Acer saccharum. Plant, Cell & Environment, 1997, 20: 845-866
[25] Zhang SB, Zhou ZK, Hu H, et al. Gas exchange and resource utilization in two alpine oaks at different altitudes in the Hengduan Mountains. Canadian Journal of Forest Research, 2007, 37: 1184-1193
[26] Eichelmann H, Laisk A. Ribulose-1,5-bisphosphate carboxylase/oxygenase content, assimilatory charge, and mesophyll conductance in leaves. Plant Physiology, 1999, 119: 179-190
[27] Li Y, Ren BB, Ding L, et al. Does chloroplast size influence photosynthetic nitrogen use efficiency? PLoS One, 2013, 8(4): e62036
[28] Benomar L, Lamhamedi MS, Villeneuve I, et al. Fine-scale geographic variation in photosynthetic-related traits of Picea glauca seedlings indicates local adaptation to climate. Tree Physiology, 2015, 35: 864-878
[29] Harley PC, Loreto F, Dimarco G, et al. Theoretical considerations when estimating the mesophyll conduc-tance to CO2 flux by analysis of the response of photosynthesis to CO2. Plant Physiology, 1992, 98: 1429-1436
[30] Niinemets ü, Díaz-Espejo A, Flexas J. Role of mesophyll diffusion conductance in constraining potential photosynthetic productivity in the field. Journal of Expe-rimental Botany, 2009, 60: 2249-2270
[31] Bown HE, Watt MS, Mason EG, et al. The influence of nitrogen and phosphorus supply and genotype on mesophyll conductance limitations to photosynthesis in Pinus radiata. Tree Physiology, 2009, 29: 1143-1151
[32] Flexas J, Niinemets ü, Gallé A, et al. Diffusional conductances to CO2 as a target for increasing photosynthesis and photosynthetic water-use efficiency. Photosynthesis Research, 2013, 117: 45-59
[33] Gale J. Availability of carbon dioxide for photosynthesis at high altitudes: Theoretical considerations. Ecology, 1972, 53: 494-497
[34] Woodward FI, Bazzaz FA. The responses of stomatal density to CO2 partial pressure. Journal of Experimental Botany, 1988, 39: 1771-1781
[35] Nobel PS, Zaragoza LJ, Smith WK. Relation between mesophyll surface-area, photosynthetic rate, and illumination level during development for leaves of Plectranthus-parviflorus Henckel. Plant Physiology, 1975, 55: 1067-1070
[36] Long SP, Humphries S, Falkowski PG. Photoinhibition of photosynthesis in nature. Annual Review of Plant Physiology and Plant Molecular Biology, 1994, 45: 633-662
[37] Li H-M (李惠梅), Shi S-B (师生波). Photosynthetic characteristics of Gentiana straminea at different altitudes. Journal of Anhui Agricultural Sciences (安徽农业科学), 2008, 36(11): 4799-4804 (in Chinese)
[38] Lu W-M (卢文敏), Liu W-G (刘伟国), Fang X-Y (方晓雨), et al. Spectral reflectance of Betula ermanii at different altitudes in the Changbai mountains. Journal of Beijing Forestry University (北京林业大学学报), 2011, 33(1): 55-59 (in Chinese)
[39] 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)
[40] Shi Z-M (史作民), Tang J-C (唐敬超), Cheng R-M (程瑞梅), et al. A review of nitrogen allocation in leaves and factors in its effects. Acta Ecologica Sinica (生态学报), 2015, 35(18): 5909-5919 (in Chinese)
[41] Lei YB, Chen K, Hao J, et al. Contrasting responses in the growth and energy utilization properties of sympatric Populus and Salix to different altitudes: Implications for sexual dimorphism in Salicaceae. Physiologia Plantarum, 2017, 159: 30-41
[42] González-Zurdo P, Escudero A, Babiano J, et al. Costs of leaf reinforcement in response to winter cold in evergreen species. Tree Physiology, 2016, 36: 273-286
[43] Onoda Y, Wright IJ, Evans JR, et al. Physiological and structural tradeoffs underlying the leaf economics spectrum. New Phytologist, 2017, 214: 1447-1463
[44] Harrison MT, Edwards EJ, Farquhar GD, et al. Nitrogen in cell walls of sclerophyllous leaves accounts for little of the variation in photosynthetic nitrogen-use efficiency. Plant, Cell & Environment, 2009, 32: 259-270