贝加尔针茅光合特征与叶片功能特性对长期氮添加的响应

doi:10.11686/cyxb2016110

摘要/Abstract

摘要： 以贝加尔针茅草甸草原建群种贝加尔针茅为研究对象,通过一个连续6年氮添加野外控制试验,设置5个氮素处理:N₀(0 kg N/hm²)、N₃₀ (30 kg N/hm²)、N₅₀ (50 kg N/hm²)、N₁₀₀ (100 kg N/hm²)、N₁₅₀ (150 kg N/hm²),探讨氮沉降变化对贝加尔针茅种群光合特性与叶片功能特性变化规律及其影响因子。结果表明,氮素添加变化显著影响贝加尔针茅的净光合速率等光合特性,氮素添加处理净光合速率(net photosynthetic rate, P_n)、气孔导度(stomatal conductance, G_s)、蒸腾速率(transpiration rate, T_r)、光合氮利用效率(photosynthetic nitrogen use efficiency, PNUE)和光合能量利用效率(photosynthetic energy use efficiency, PEUE)均低于或显著低于无氮素添加处理。氮素添加处理叶片比叶面积(specific leaf area, SLA)、叶片N含量(leaf N content, N_mass)、建成成本(leaf construction cost, CC_mass)、叶片N∶P均高于或显著高于无氮素添加处理。相关分析表明,5个氮素处理的P_n与G_s、T_r、PNUE、PEUE、叶片P含量(leaf P content, P_mass)、土壤N∶P呈显著正相关,与叶片比叶面积(SLA)、N_mass、CC_mass、土壤含水量、土壤pH值呈极显著负相关;叶片N_mass与SLA、CC_mass、土壤N∶P呈显著正相关,与PNUE、PEUE、P_mass、土壤含水量、土壤pH值呈显著负相关。综合以上研究表明,长期氮添加会降低贝加尔针茅净光合速率、养分利用效率,提高叶片建成成本和叶片N∶P,土壤含水量和土壤pH值是引起这种改变发生的主要因子。

Abstract: Atmospheric nitrogen deposition is an important component phenomenon of global change, and induces a series of ecological problems. We conducted a field manipulation experiment to study the photosynthetic response and change in leaf functional traits after long-term nitrogen addition in an S.baicalensis grassland in Inner Mongolia. This study was conducted from 2010-2015. Five treatments including N₀ (0 kg N/ha), N₃₀ (30 kg N/ha), N₅₀ (50 kg N/ha), N₁₀₀ (100 kg N/ha) and N₁₅₀ (150 kg N/ha) were set up and forty-eight plots, sized 8 m×8 m, were established with 2 m strips between each plot. The objective was to determine how photosynthetic characteristics and leaf traits vary with nitrogen addition and what causes these differences. Photosynthetic traits and leaf traits differed among different nitrogen addition treatments. Net photosynthetic rate (P_n), stomatal conductance (G_s), transpiration rate (T_r), photosynthetic nitrogen use efficiency (PNUE), photosynthetic energy use efficiency (PEUE) under N₃₀, N₅₀, N₁₀₀, and N₁₅₀ treatments were lower or significantly lower than those under N₀. Specific leaf area (SLA), leaf N content (N_mass), leaf construction cost (CC_mass), leaf N∶P under N₃₀, N₅₀, N₁₀₀, N₁₅₀ treatments tended to be higher or were significantly higher than those under N₀. Correlation analysis showed that P_n was positively correlated with G_s, T_r, PNUE, PEUE, leaf P content (P_mass), N_mass and soil N, P while P_n was negatively correlated with SLA, N_mass, CC_mass, soil water content and soil pH; N_mass was positively correlated with SLA, CC_mass and soil N, P, and N_mass was negatively correlated with PNUE, PEUE, P_mass, soil water content and soil pH. In conclusion, long-term nitrogen addition induced decreases in P_n and nutrient use efficiency of S. baicalensis, and increases in leaf CC_mass and N∶P. Soil water content and pH decreased with nitrogen addition. This suggests that water content and soil pH are two important factors affecting photosynthetic characteristics and leaf traits of S.baicalensis under different nitrogen addition regimes.

刘红梅, 李洁, 皇甫超河, 陈新微, 杨殿林. 贝加尔针茅光合特征与叶片功能特性对长期氮添加的响应[J]. 草业学报, 2016, 25(11): 76-85.

LIU Hong-Mei, LI Jie, HUANGFU Chao-He, CHEN Xin-Wei, YANG Dian-Lin. Effects of long-term nitrogen addition on photosynthetic characteristics and leaf traits of Stipa baicalensis in Inner Mongolia, China[J]. Acta Prataculturae Sinica, 2016, 25(11): 76-85.

参考文献

[1] Galloway J N, Dentener F J, Capone D G, et al . Nitrogen cycles: past, present, and future. Biogeochemistry, 2004, 70(2): 152-226.
[2] Liu X J, Zhang Y, Han W X, et al . Enhanced nitrogen deposition over China. Nature, 2013, 494: 459-462.
[3] Li W J, Liu H M, Zhao J N, et al . Effects of nitrogen and water addition on plant species diversity and biomass of common species in the Stipa baicalensis Steppe, Inner Mongolia, China. Acta Ecologiga Sinica, 2015, 35(19): 6460-6469.
[4] Clark C M, David T. Loss of plant species after chronic low-level nitrogen deposition to prairie grasslands. Nature, 2008, 451: 712-715.
[5] Chalcraft D R, Cox S B, Clark C, et al . Scale-dependent responses of plant biodiversity to nitrogen enrichment. Ecology, 2008, 89(8): 2165-2171.
[6] Huang J Y, Yu H L, Yuan Z Y, et al . Effects of long-term increased soil N on leaf traits of several species in typical Inner Mongolian grassland. Acta Ecologiga Sinica, 2012, 32(5): 1419-1427.
[7] Wan H W, Yang Y, Bai S Q, et al . Variations in leaf functional traits of six species along a nitrogen addition gradient in Leymus chinensis steppe in Inner Mongolia. Journal of Plant Ecology, 2008, 32(3): 611-621.
[8] Ceulemans R, Janssens I A, Jach M E. Effects of CO 2 enrichment on trees and forests: lessons to be learned in view of future ecosystem studies. Annuals of Botany, 1999, 84(5): 577-590.
[9] Maroco J P, Breia E, Faria T, et al . Effects of long-term exposure to elevated CO 2 and N fertilization on the development of photosynthetic capacity and biomass accumulation in Quercus suber L. Plant Cell and Environment, 2002, 25(1): 105-113.
[10] Zhang X C, Shangguan Z P. The responses of photosynthetic electron transport and partition in the winter wheat leaves of different drought resistances to nitrogen levels. Plant Physiology Communications, 2009, 45(1): 13-18.
[11] Nakaji T, Fukami M, Dokiya Y, et al . Effects of high nitrogen load on growth, photosynthesis and nutrient status of Cryptomeria japonica and Pinus densiflora seedlings. Trees, 2001, 15(8): 453-461.
[12] Neff J C, Townsend A R, Gleixner G, et al . Variable effects of nitrogen additions on the stability and turnover of soil carbon. Nature, 2002, 419: 915-917.
[13] Guo J H, Liu X J, Zhang Y, et al . Significant acidification in major Chinese croplands. Science, 2010, 327: 1008-1010.
[14] Li B. Influence of Warming and N Deposition on Physiological and Ecological Characteristic of Leymus chinesis in the Songnen Grassland[D]. Changchun: Northeast Normal University, 2015.
[15] Bin Z J, Wang J J, Zhang W P, et al . Effects of N addition on ecological stoichiometric characteristics in six dominant plant species of alpine meadow on the Qinghai-Xizang Plateau, China. Chinese Journal of Plant Ecology, 2014, 38(3): 231-237.
[16] Hu J Y, Zhu J X, Zhou Z, et al . Effects of experimental nitrogen additions on understory species diversity in four forests in China. Acta Scientiarum Naturalium Universitatis Pekinensis, 2014, 50(5): 904-910.
[17] Huenneke L F, Hamburg S P, Koide R, et al . Effects of soil resources on plant invasion and community structure in Californian serpentine grassland. Ecology Usa, 1990, 71(2): 478-491.
[18] Zhao S, Zhang J N, Lai X, et al . Analysis of microbial biomass C, N and soil microbial community structure of Stipa steppes using PLFA at grazing and fenced in Inner Mongolia, China. Journal of Agro-Environment Science, 2011, 30(6): 1126-1134.
[19] Zhang N L, Wan S Q, Li L H. Impacts of urea N addition on soil microbial community in a semiarid temperate steppe in northern China. Plant and Soil, 2008, 311(1/2): 19-28.
[20] Deng Y, Liu X N, Xin X P, et al . Diurnal dynamics of photosynthetic characteristics of Leymus chinensis under different grazing intensities taking the Hulunber meadow steppe as an example. Acta Prataculturae Sinica, 2012, 21(3): 308-313.
[21] Bao S D. Soil and Agricultural Chemistry Analysis[D]. Third Edition. Beijing: China Agricultural Press, 2000.
[22] Williams K, Percival F, Merinao J, et al . Estimation of tissue construction cost from heat of combustion and organic nitrogen content. Plant, Cell & Environment, 1987, 10(9): 725-734.
[23] Vries F W, Brunsting A H, Laar H H V. Products, requirements and efficiency of biosynthesis: A quantitative approach. Journal of Theoretical Biology, 1974, 45(2): 339-377.
[24] Inclan R, Alonso R, Pujadas M, et al . Ozone and drought stress: Interactive effects on gas exchange in Aleppo pine ( Pinus halepensis Mill). Chemosphere, 1998, 36(4/5): 685-690.
[25] Wan S, Xia J, Liu W, et al . Photosynthetic overcompensation under nocturnal warming enhances grassland carbon sequestration. Ecology, 2009, 90: 2700-2710.
[26] Bai Y F, Wu J G, Xing Q, et al . Primary production and rain use efficiency across a precipitation gradient on the Mongolia Plateau. Ecology, 2008, 89(8): 2140-2153.
[27] Craine J M, Morrow C, Stock W D. Nutrient concentration ratios and co-limitation in South African grasslands. New Phytologist, 2008, 179(3): 829-836.
[28] Asner G P, Townsend A R, Riley W J, et al . Physical and biogeochemical controls over terrestrial ecosystem responses to nitrogen deposition. Biogeochemistry, 2001, 54(1): 1-39.
[29] Xu D Q. Some problems in stomatal limitation analysis of photosynthesis. Plant Physiology Communications, 1997, 33(4): 241-244.
[30] Xiao S S, Dong Y S, Qi Y C, et al . Effects of mineral fertilizer addition on leaf functional traits and photosynthetic characteristics of Leymus chinensis from a temperate grassland in Inner Mongolia in China. Acta Scientiae Circumstantiae, 2010, 30(12): 2535-2543.
[31] An H, Shangguan Z P. Specific leaf area, leaf nitrogen content, and photosynthetic acclimation of Trifolium repens L. seedlings grown at different irradiances and nitrogen concentrations. Photosynthetica, 2008, 46(1): 143-147.
[32] Yang H, Luo Y C. Responses of the functional traits in Cleistogenes squarrosa to nitrogen addition and drought. Chinese Journal of Plant Ecology, 2015, 39(1): 32-42.
[33] Novriyanti E, Watanabe M, Makoto K, et al . Photosynthetic nitrogen and water use efficiency of acacia and eucalypt seedlings as afforestation species. Photosynthetica, 2012, 50(2): 273-281.
[34] Castellanos A E, Martinez M J, Llano J M, et al . Successional trends in Sonoran desert abandoned agricultural fields in northern Mexico. Journal of Arid Environments, 2005, 60(3): 437-455.
[35] Mahajan S, Tuteja T. Cold, salinity and drought stresses: An overview. Archives of Biochemistry & Biophysics, 2005, 444(2): 139-158.
[36] Dou J X, Liu J S, Wang Y, et al . Ecophysiological responses of Calamagrostis angustifolia to nitrogen deposition. Wetland Science, 2009, 7(1): 40-46.
[37] Lambers H, Chapin III F S, Pons T L. Plant Physiological Ecology[M]. Second Edition. New York: Springer-Verlag, 2008.
[38] Li L Z, Zhang D G, Xin X P, et al . Photosynthetic characteristics of Leymus chinensis under different soil moisture grades in Hulunber prairie. Acta Ecologiga Sinica, 2009, 29(10): 5271-5279.
[39] Bergstrom A K, Faithfull C, Karlsson D, et al . Nitrogen deposition and warming-effects on phytoplankton nutrient limitation in subarctic lakes. Global Change Biology, 2013, 19(8): 2557-2568.
[3] 李文娇, 刘红梅, 赵建宁, 等. 氮素和水分添加对贝加尔针茅草原植物多样性及生物量的影响. 生态学报, 2015, 35(19): 6460-6469.
[6] 黄菊莹, 余海龙, 袁志友, 等. 长期 N 添加对典型草原几个物种叶片性状的影响. 生态学报, 2012, 32(5): 1419-1427.
[7] 万宏伟, 杨阳, 白世勤, 等. 羊草草原群落 6 种植物叶片功能特性对氮素添加的响应. 植物生态学报, 2008, 32(3): 611-621.
[14] 李博. 温度升高和氮沉降对松嫩草地羊草种群生理与生态特性的影响[D]. 长春: 东北师范大学, 2015.
[15] 宾振钧, 王静静, 张文鹏, 等. 氮肥添加对青藏高原高寒草甸6个群落优势种生态化学计量学特征的影响. 植物生态学报, 2014, 38(3): 231-237.
[16] 胡钧宇, 朱剑霄, 周璋, 等. 氮添加对4种森林类型林下植物多样性的影响. 北京大学学报, 2014, 50(5): 904-910.
[18] 赵帅, 张静妮, 赖欣, 等. 放牧与围栏内蒙古针茅草原土壤微生物生物量碳、氮变化及微生物群落结构 PLFA 分析. 农业环境科学学报, 2011, 30(6): 1126-1134.
[20] 邓钰, 柳小妮, 辛晓平, 等. 不同放牧强度下羊草的光合特性日动态变化. 草业学报, 2012, 21(3): 308-313.
[21] 鲍士旦. 土壤农化分析[D]. 第三版. 北京: 中国农业出版社, 2000.
[29] 许大全. 光合作用气孔限制分析中的一些问题. 植物生理学通讯, 1997, 33(4): 241-244.
[30] 肖胜生, 董云社, 齐玉春, 等. 内蒙古温带草原羊草叶片功能特性与光合特征对外源氮输入的响应. 环境科学学报, 2010, 30(12): 2535-2543.
[32] 杨浩, 罗亚晨. 糙隐子草功能性状对氮添加和干旱的响应. 植物生态学报, 2015, 39(1): 32-42.
[36] 窦晶鑫, 刘景双, 王洋, 等. 小叶章对氮沉降的生理生态响应. 湿地科学, 2009, 7(1): 40-46.
[38] 李林芝, 张德罡, 辛晓平. 等. 呼伦贝尔草甸草原不同土壤水分梯度下羊草的光合特性. 生态学报, 2009, 29(10): 5271-5279.