氮沉降和生物质炭对毛竹叶片光合特性的影响
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摘要
为了解氮沉降和生物质炭对毛竹Phyllostachys edulis叶片光合特性的影响,研究了4种氮沉降(0 kg·hm~(-2)·a~(-1),N_0;30 kg·hm~(-2)·a~(-1),N_(30);60 kg·hm~(-2)·a~(-1),N_(60);90 kg·hm~(-2)·a~(-1),N_(90))下施加3种不同强度(0 t·hm~(-2),BC_0;20 t·hm~(-2),BC_(20);40 t·hm~(-2),BC_(40))生物质炭处理后2龄毛竹新老叶片光合及叶绿素荧光特性的变化。结果表明:氮沉降和生物质炭均促进了毛竹新老叶的最大净光合速率(P_(max)),气孔导度(G_s)和蒸腾速率(T_r),生物质炭同时提高了老叶实际量子产量[Y(Ⅱ)]和PSⅡ潜在活性(F_v/F_0)。与不施加生物质炭相比(BC_0),N_(30)下施加20 t·hm~(-2)生物质炭的老叶及N_(60)下施加20 t·hm~(-2)生物质炭(BC_(20))的新老叶叶色值显著降低;N_(60)处理下施加生物质炭,毛竹新老叶的最大光化学效率(F_v/F_m),F_v/F_o和Y(Ⅱ)均提高,而N_(90)处理下施加生物质炭,毛竹新老叶的F_v/F_m和F_v/F_o均降低。由此认为:氮沉降条件下施加生物质炭有利于毛竹新老叶片的光合固碳能力;大气氮沉降背景下,施加生物质炭有利于提高毛竹生产力。
To understand the effects of nitrogen(N)deposition and biochar amendments on photosynthesis of new and old leaves of Moso bamboo(Phyllostachys edulis),effects of four N additions(N_0:0 kg·hm~(-2)·a~(-1),N_(30):30 kg·hm~(-2)·a~(-1),N_(60):60 kg·hm~(-2)·a~(-1),and N_(90):90 kg·hm~(-2)·a~(-1))and three biochar amendments(BC_0:0 t·hm~(-2),BC_(20):20 t·hm~(-2),and BC_(40):40 t·hm~(-2))on photosynthesis and chlorophyll fluorescence characteristics of new and old leaves in two-year-old Moso bamboo were determined using LI-6400 portable photosynthesis system,PAM-2500 chlorophyll fluorescence spectrometer and SPAD-502 chlorophyll meter in Moso bamboo plantations(20m×20 m).Results showed that N deposition and biochar amendment significantly increased(P<0.05)the maximum net photosynthetic rate(P_(max)),stomatal conductance(G_s),and transpiration rate(T_r)of new and old leaves.Biochar amendments significantly increased(P<0.001)effective quantum yield of PhotosystemⅡ(PS-Ⅱ)[Y(Ⅱ)]and potential activities of PS-Ⅱ(F_v/F_0)of old leaves.In combination with both N depositions(N_(30)and N_(60)),BC_(20)significantly decreased(P<0.001)the relative chlorophyll content(SPAD)of old leaves compared with the N depositions alone.The SPAD of new leaves for N_(60)-BC_(20)was significantly lower(P<0.001)than N_(60)-BC_0.Also,both biochar amendments in combination with N_(60)significantly increased(P<0.001)maximal potential quantum yield of PS-Ⅱ(F_v/F_m),F_v/F_o,and Y(Ⅱ)of new and old leaves compared to N depositions alone.However,biochar amendments in combination with N_(90)significantly decreased(P<0.001)both F_v/F_mand F_v/F_oof new and old leaves compared to N depositions alone.In conclusion,this study demonstrated that biochar amendments promoted photosynthesis of leaves and thus increased productivity of a Moso bamboo forest with the background of globally increasing N deposition.
To understand the effects of nitrogen(N)deposition and biochar amendments on photosynthesis of new and old leaves of Moso bamboo(Phyllostachys edulis),effects of four N additions(N_0:0 kg·hm~(-2)·a~(-1),N_(30):30 kg·hm~(-2)·a~(-1),N_(60):60 kg·hm~(-2)·a~(-1),and N_(90):90 kg·hm~(-2)·a~(-1))and three biochar amendments(BC_0:0 t·hm~(-2),BC_(20):20 t·hm~(-2),and BC_(40):40 t·hm~(-2))on photosynthesis and chlorophyll fluorescence characteristics of new and old leaves in two-year-old Moso bamboo were determined using LI-6400 portable photosynthesis system,PAM-2500 chlorophyll fluorescence spectrometer and SPAD-502 chlorophyll meter in Moso bamboo plantations(20m×20 m).Results showed that N deposition and biochar amendment significantly increased(P<0.05)the maximum net photosynthetic rate(P_(max)),stomatal conductance(G_s),and transpiration rate(T_r)of new and old leaves.Biochar amendments significantly increased(P<0.001)effective quantum yield of PhotosystemⅡ(PS-Ⅱ)[Y(Ⅱ)]and potential activities of PS-Ⅱ(F_v/F_0)of old leaves.In combination with both N depositions(N_(30)and N_(60)),BC_(20)significantly decreased(P<0.001)the relative chlorophyll content(SPAD)of old leaves compared with the N depositions alone.The SPAD of new leaves for N_(60)-BC_(20)was significantly lower(P<0.001)than N_(60)-BC_0.Also,both biochar amendments in combination with N_(60)significantly increased(P<0.001)maximal potential quantum yield of PS-Ⅱ(F_v/F_m),F_v/F_o,and Y(Ⅱ)of new and old leaves compared to N depositions alone.However,biochar amendments in combination with N_(90)significantly decreased(P<0.001)both F_v/F_mand F_v/F_oof new and old leaves compared to N depositions alone.In conclusion,this study demonstrated that biochar amendments promoted photosynthesis of leaves and thus increased productivity of a Moso bamboo forest with the background of globally increasing N deposition.
引文
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[2]李德军,莫江明,方运霆,等.模拟氮沉降对南亚热带两种乔木幼苗生物量及其分配的影响[J].植物生态学报, 2005, 29(4):543-549.LI Dejun, MO Jiangming, FANG Yunting, et al. Effects of simulated nitrogen deposition on biomass production and allocation in Schima superba and seedlings in subtropical China[J]. J Plant Ecol, 2005, 29(4):543-549.
[3] ZHANG Rui, WU Jiasheng, LI Quan, et al. Nitrogen deposition enhances photosynthesis in Moso bamboo but increases susceptibility to other stress factors[J]. Front Plant Sci, 2017. doi:10.3389/fpls.2017.01975.
[4]高培军,邱永华,周紫球,等.氮素施肥对毛竹生产力与光合能力的影响[J].浙江农林大学学报, 2014, 31(5):697-703.GAO Peijun, QIU Yonghua, ZHOU Ziqiu, et al. Productivity and photosynthetic ability of Phyllostachys edulis with nitrogen fertilization[J]. J Zhejiang A&F Univ, 2014, 31(5):697-703.
[5]宋久洋,刘领,陈明灿,等.生物质炭施用对烤烟生长及光合特性的影响[J].河南科技大学学报(自然科学版), 2014, 35(4):68-72.SONG Jiuyang, LIU Ling, CHEN Mingcan, et al. Effects of biochar addition on growth and photosynthetic characteristics of flue-cured tobacco[J]. J Henan Univ Sci Technol Nat Sci, 2014, 35(4):68-72.
[6]陈红霞,杜章留,郭伟,等.施用生物炭对华北平原农田土壤容重、阳离子交换量和颗粒有机质含量的影响[J].应用生态学报, 2011, 22(11):2930-2934.CHEN Hongxia, DU Zhangliu, GUO Wei, et al. Effects of biochar amendment on cropland soil bulk density, cation exchange capacity, and particulate organic matter content in the North China Plain[J]. J Appl Ecol, 2011, 22(11):2930-2934.
[7]吕一甲.生物炭肥对土壤性质、玉米生长及水肥利用效率影响试验研究[D].呼和浩特:内蒙古农业大学,2014.L譈Yijia. The Experimental Study of Influence of Biochar Fertilizer on Soil Properties, Maize Growth and Water and Fertilizer Use Efficiency[D]. Huhhot:Inner Mongolia Agricutural University, 2014.
[8] SOLAIMAN Z M, MURPHY D V, ABBOTT L K. Biochar influence seed germination and early growth of seedlings[J]. Plant Soil, 2012, 353(1/2):273-287.
[9]刘玉学,王耀锋,吕豪豪,等.生物质炭化还田对稻田温室气体排放及土壤理化性质的影响[J].应用生态学报, 2013, 24(8):2166-2172.LIU Yuxue, WANG Yaofeng, L譈Haohao, et al. Effects of biochar application on greenhouse gas emission from paddy soil and its physical and chemical properties[J]. J Appl Ecol, 2013, 24(8):2166-2172.
[10]钟哲科,李伟成,刘玉学,等.竹炭的土壤环境修复功能[J].竹子研究汇刊, 2009, 28(3):9-13.ZHONG Zheke, LI Weicheng, LIU Yuxue, et al. Soil environmental remediation functions of bamboo charcoal[J]. J Bamboo Res, 2009, 28(3):9-13.
[11] GALLOWAY J N, TOWNSEND A R, ERISMAN J W, et al. Transformation of the nitrogen cycle:recent trends,questions, and potential solutions[J]. Science, 2008, 320(5878):889-892.
[12] SONG Xinzhang, ZHOU Guomo, JIANG Hong, et al. Carbon sequestration by Chinese bamboo forests and their ecological benefits:assessment of potential, problems, and future challenges[J]. Environ Rev, 2011, 19:418-428.
[13]国家林业局.中国森林资源报告(2009-2013)[M].北京:中国林业出版社, 2014.
[14] SONG Xinzhang, GU Honghao, WANG Meng, et al. Management practices regulate the response of Moso bamboo foliar stoichiometry to nitrogen deposition[J]. Sci Rep, 2016, 6:24107. doi:10.10138/serp24107.
[15]金晓春,金爱武,宋艳冬,等.施肥对毛竹林换叶期冠层形成及光合能力的影响[J].浙江林学院学报,2010, 27(1):57-62.JIN Xiaochun, JIN Aiwu, SONG Yandong, et al. Canopy development and photosynthesis with fertilization during leaf-expansion period of Phyllostachys pubescens[J]. J Zhejiang For Coll, 2010, 27(1):57-62.
[16] REAY D S, DENTENER F, SMITH P, et al. Global nitrogen deposition and carbon sinks[J]. Nat Geosci, 2008, 1(7):430-437.
[17] SONG Xinzhang, LI Quan, GU Honghao. Effects of nitrogen deposition and management practices on fine root decomposition in Moso bamboo plantations[J]. Plant Soil, 2017, 410(1/2):207-215.
[18] SONG Xinzhang, ZHOU Guomo, GU Honghao, et al. Management practices amplify the effects of N deposition on leaf litter decomposition of Moso bamboo forest[J]. Plant Soil, 2015, 395(1/2):391-400.
[19] ZHANG Rui, ZHAO Yuxin, LIN Jianhong, et al. Biochar application alleviates unbalanced nutrient uptake caused by N deposition in Torreya grandis trees and seedlings[J]. For Ecol Manage, 2019, 432:319-326.
[20]吴志庄,杜旭华,熊德礼,等.不同类型竹种光合特性的比较研究[J].生态环境学报, 2013, 22(9):1523-1527.WU Zhizhuang, DU Xuhua, XIONG Deli, et al. A comparative study on photosynthetic characteristics of different types of bamboos[J]. Ecol Environ Sci, 2013, 22(9):1523-1527.
[21]张毅龙,张卫强,甘先华.低温胁迫对6种珍贵树种苗木光合荧光特性的影响[J].生态环境学报, 2014, 23(5):777-784.ZHANG Yilong, ZHANG Weiqiang, GAN Xianhua. Influence of low temperature stress on the photosynthetic fluorescence characteristics of 6 kinds of precious hardwoods seedling in winter[J]. Ecol Environ Sci, 2014, 23(5):777-784.
[22]施建敏,郭起荣,杨光耀,等.毛竹光合作用对环境因子的季节响应[J].广西植物, 2007, 27(6):923-928.SHI Jianmin, GUO Qirong, YANG Guangyao, et al. Seasonal photosynthetic responses of Phyllostachys edulis to environmental factors[J]. Guihaia, 2007, 27(6):923-928.
[23]许大全,李德耀,邱国雄,等.毛竹(Phyllostachys pubescens)叶光合作用的气孔限制研究[J].植物生理学报,1987, 13(2):44-50.XU Daquan, LI Deyao, QIU Guoxiong, et al. Studies on stomatal limitation of photosynthesis in the bamboo(Phyllostachy pubescens)leaves[J]. Acta Phytophysiol Sin, 1987, 13(2):44-50.
[24]曲道春,江洪,由美娜.氮沉降对香樟叶片光合及叶绿素荧光特性的影响研究[J].环境污染与防治,2011, 33(11):15-19.QU Daochun, JIANG Hong, YOU Meina. Effects of nitrogen deposition on photosynthesis and chlorophyll fluorescence characteristics of Cinnamomum camphora[J]. Environ Pollut Control, 2011, 33(11):15-19.
[25]周哲宇,徐超,胡策,等.毛竹快速生长期的叶绿素荧光参数特征[J].浙江农林大学学报, 2018, 35(1):75-80.ZHOU Zheyu, XU Chao, HU Ce, et al. Chlorophyll fluorescence characteristics of Phyllostachys edulis during its fast growth period[J]. J Zhejiang A&F Univ, 2018, 35(1):75-80.
[26]陈嘉琦,温国胜,王艳红,等.春季毛竹光化学效率空间异质性比较[J].福建林学院学报, 2014, 34(1):52-56.CHEN Jiaqi, WEN Guosheng, WANG Yanhong, et al. Comparison of spatial heterogeneity in photochemical efficiency of Phyllostachys pubescens in spring[J]. J Fujian Coll For, 2014, 34(1):52-56.
[27]吴志庄,王道金,厉月桥,等.施用生物炭肥对黄连木生长及光合特性的影响[J].生态环境学报, 2015, 24(6):992-997.WU Zhizhuang, WANG Daojin, LI Yueqiao, et al. Effects of biochar fertilizer application on growth properties and photosynthetic and physiological characteristics of Pistacia chinensis Bunge[J]. Ecol Environ Sci, 2015, 24(6):992-997.
[28]李艳梅,杨俊刚,孙焱鑫,等.炭基氮肥与灌水对温室番茄产量、品质及土壤硝态氮残留的影响[J].农业环境科学学报, 2015, 34(10):1965-1972.LI Yanmei, YANG Jungang, SUN Yanxin, et al. Coupling effects of biochar-based urea and irrigation on tomato yield and fruit quality and soil nitrate content in greenhouse facility[J]. J Agro-Environ Sci, 2015, 34(10):1965-1972.
[29]刘领,王艳芳,宋久洋,等.生物炭与氮肥减量配施对烤烟生长及土壤酶活性的影响[J].河南农业科学,2016, 45(2):62-66.LIU Ling, WANG Yanfang, SONG Jiuyang, et al. Effects of biochar addition combined with reducing nitrogen application rate on growth of flue-cured tobacco and soil enzyme activities[J]. J Henan Agric Sci, 2016, 45(2):62-66.
[2]李德军,莫江明,方运霆,等.模拟氮沉降对南亚热带两种乔木幼苗生物量及其分配的影响[J].植物生态学报, 2005, 29(4):543-549.LI Dejun, MO Jiangming, FANG Yunting, et al. Effects of simulated nitrogen deposition on biomass production and allocation in Schima superba and seedlings in subtropical China[J]. J Plant Ecol, 2005, 29(4):543-549.
[3] ZHANG Rui, WU Jiasheng, LI Quan, et al. Nitrogen deposition enhances photosynthesis in Moso bamboo but increases susceptibility to other stress factors[J]. Front Plant Sci, 2017. doi:10.3389/fpls.2017.01975.
[4]高培军,邱永华,周紫球,等.氮素施肥对毛竹生产力与光合能力的影响[J].浙江农林大学学报, 2014, 31(5):697-703.GAO Peijun, QIU Yonghua, ZHOU Ziqiu, et al. Productivity and photosynthetic ability of Phyllostachys edulis with nitrogen fertilization[J]. J Zhejiang A&F Univ, 2014, 31(5):697-703.
[5]宋久洋,刘领,陈明灿,等.生物质炭施用对烤烟生长及光合特性的影响[J].河南科技大学学报(自然科学版), 2014, 35(4):68-72.SONG Jiuyang, LIU Ling, CHEN Mingcan, et al. Effects of biochar addition on growth and photosynthetic characteristics of flue-cured tobacco[J]. J Henan Univ Sci Technol Nat Sci, 2014, 35(4):68-72.
[6]陈红霞,杜章留,郭伟,等.施用生物炭对华北平原农田土壤容重、阳离子交换量和颗粒有机质含量的影响[J].应用生态学报, 2011, 22(11):2930-2934.CHEN Hongxia, DU Zhangliu, GUO Wei, et al. Effects of biochar amendment on cropland soil bulk density, cation exchange capacity, and particulate organic matter content in the North China Plain[J]. J Appl Ecol, 2011, 22(11):2930-2934.
[7]吕一甲.生物炭肥对土壤性质、玉米生长及水肥利用效率影响试验研究[D].呼和浩特:内蒙古农业大学,2014.L譈Yijia. The Experimental Study of Influence of Biochar Fertilizer on Soil Properties, Maize Growth and Water and Fertilizer Use Efficiency[D]. Huhhot:Inner Mongolia Agricutural University, 2014.
[8] SOLAIMAN Z M, MURPHY D V, ABBOTT L K. Biochar influence seed germination and early growth of seedlings[J]. Plant Soil, 2012, 353(1/2):273-287.
[9]刘玉学,王耀锋,吕豪豪,等.生物质炭化还田对稻田温室气体排放及土壤理化性质的影响[J].应用生态学报, 2013, 24(8):2166-2172.LIU Yuxue, WANG Yaofeng, L譈Haohao, et al. Effects of biochar application on greenhouse gas emission from paddy soil and its physical and chemical properties[J]. J Appl Ecol, 2013, 24(8):2166-2172.
[10]钟哲科,李伟成,刘玉学,等.竹炭的土壤环境修复功能[J].竹子研究汇刊, 2009, 28(3):9-13.ZHONG Zheke, LI Weicheng, LIU Yuxue, et al. Soil environmental remediation functions of bamboo charcoal[J]. J Bamboo Res, 2009, 28(3):9-13.
[11] GALLOWAY J N, TOWNSEND A R, ERISMAN J W, et al. Transformation of the nitrogen cycle:recent trends,questions, and potential solutions[J]. Science, 2008, 320(5878):889-892.
[12] SONG Xinzhang, ZHOU Guomo, JIANG Hong, et al. Carbon sequestration by Chinese bamboo forests and their ecological benefits:assessment of potential, problems, and future challenges[J]. Environ Rev, 2011, 19:418-428.
[13]国家林业局.中国森林资源报告(2009-2013)[M].北京:中国林业出版社, 2014.
[14] SONG Xinzhang, GU Honghao, WANG Meng, et al. Management practices regulate the response of Moso bamboo foliar stoichiometry to nitrogen deposition[J]. Sci Rep, 2016, 6:24107. doi:10.10138/serp24107.
[15]金晓春,金爱武,宋艳冬,等.施肥对毛竹林换叶期冠层形成及光合能力的影响[J].浙江林学院学报,2010, 27(1):57-62.JIN Xiaochun, JIN Aiwu, SONG Yandong, et al. Canopy development and photosynthesis with fertilization during leaf-expansion period of Phyllostachys pubescens[J]. J Zhejiang For Coll, 2010, 27(1):57-62.
[16] REAY D S, DENTENER F, SMITH P, et al. Global nitrogen deposition and carbon sinks[J]. Nat Geosci, 2008, 1(7):430-437.
[17] SONG Xinzhang, LI Quan, GU Honghao. Effects of nitrogen deposition and management practices on fine root decomposition in Moso bamboo plantations[J]. Plant Soil, 2017, 410(1/2):207-215.
[18] SONG Xinzhang, ZHOU Guomo, GU Honghao, et al. Management practices amplify the effects of N deposition on leaf litter decomposition of Moso bamboo forest[J]. Plant Soil, 2015, 395(1/2):391-400.
[19] ZHANG Rui, ZHAO Yuxin, LIN Jianhong, et al. Biochar application alleviates unbalanced nutrient uptake caused by N deposition in Torreya grandis trees and seedlings[J]. For Ecol Manage, 2019, 432:319-326.
[20]吴志庄,杜旭华,熊德礼,等.不同类型竹种光合特性的比较研究[J].生态环境学报, 2013, 22(9):1523-1527.WU Zhizhuang, DU Xuhua, XIONG Deli, et al. A comparative study on photosynthetic characteristics of different types of bamboos[J]. Ecol Environ Sci, 2013, 22(9):1523-1527.
[21]张毅龙,张卫强,甘先华.低温胁迫对6种珍贵树种苗木光合荧光特性的影响[J].生态环境学报, 2014, 23(5):777-784.ZHANG Yilong, ZHANG Weiqiang, GAN Xianhua. Influence of low temperature stress on the photosynthetic fluorescence characteristics of 6 kinds of precious hardwoods seedling in winter[J]. Ecol Environ Sci, 2014, 23(5):777-784.
[22]施建敏,郭起荣,杨光耀,等.毛竹光合作用对环境因子的季节响应[J].广西植物, 2007, 27(6):923-928.SHI Jianmin, GUO Qirong, YANG Guangyao, et al. Seasonal photosynthetic responses of Phyllostachys edulis to environmental factors[J]. Guihaia, 2007, 27(6):923-928.
[23]许大全,李德耀,邱国雄,等.毛竹(Phyllostachys pubescens)叶光合作用的气孔限制研究[J].植物生理学报,1987, 13(2):44-50.XU Daquan, LI Deyao, QIU Guoxiong, et al. Studies on stomatal limitation of photosynthesis in the bamboo(Phyllostachy pubescens)leaves[J]. Acta Phytophysiol Sin, 1987, 13(2):44-50.
[24]曲道春,江洪,由美娜.氮沉降对香樟叶片光合及叶绿素荧光特性的影响研究[J].环境污染与防治,2011, 33(11):15-19.QU Daochun, JIANG Hong, YOU Meina. Effects of nitrogen deposition on photosynthesis and chlorophyll fluorescence characteristics of Cinnamomum camphora[J]. Environ Pollut Control, 2011, 33(11):15-19.
[25]周哲宇,徐超,胡策,等.毛竹快速生长期的叶绿素荧光参数特征[J].浙江农林大学学报, 2018, 35(1):75-80.ZHOU Zheyu, XU Chao, HU Ce, et al. Chlorophyll fluorescence characteristics of Phyllostachys edulis during its fast growth period[J]. J Zhejiang A&F Univ, 2018, 35(1):75-80.
[26]陈嘉琦,温国胜,王艳红,等.春季毛竹光化学效率空间异质性比较[J].福建林学院学报, 2014, 34(1):52-56.CHEN Jiaqi, WEN Guosheng, WANG Yanhong, et al. Comparison of spatial heterogeneity in photochemical efficiency of Phyllostachys pubescens in spring[J]. J Fujian Coll For, 2014, 34(1):52-56.
[27]吴志庄,王道金,厉月桥,等.施用生物炭肥对黄连木生长及光合特性的影响[J].生态环境学报, 2015, 24(6):992-997.WU Zhizhuang, WANG Daojin, LI Yueqiao, et al. Effects of biochar fertilizer application on growth properties and photosynthetic and physiological characteristics of Pistacia chinensis Bunge[J]. Ecol Environ Sci, 2015, 24(6):992-997.
[28]李艳梅,杨俊刚,孙焱鑫,等.炭基氮肥与灌水对温室番茄产量、品质及土壤硝态氮残留的影响[J].农业环境科学学报, 2015, 34(10):1965-1972.LI Yanmei, YANG Jungang, SUN Yanxin, et al. Coupling effects of biochar-based urea and irrigation on tomato yield and fruit quality and soil nitrate content in greenhouse facility[J]. J Agro-Environ Sci, 2015, 34(10):1965-1972.
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