毛白杨树干呼吸及其温度敏感性季节变化特征
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摘要
树干呼吸(E_s)是森林生态系统碳循环过程的重要组成部分,深入理解树干呼吸过程对未来气候变暖的响应及反馈机制有助于更加精确地估算森林生态系统碳储量。为揭示毛白杨树干呼吸及其温度敏感性的昼夜变化和季节动态规律,利用Li-Cor6400便携式光合作用测定系统及其配套使用的土壤呼吸测量气室(LI-6400-09)对冀南平原区毛白杨的树干呼吸和树干温度实施为期1年的连续监测。结果表明:(1)在生长季,毛白杨树干呼吸与树干温度之间在晚上呈现正相关的关系(R~2=0.88);相反,两者在白天为负相关的关系(R~2=0.96)。(2)整个观测期内,毛白杨树干呼吸和树干温度均呈现"钟形"的变化曲线,树干呼吸与树干温度之间存在着较好的指数函数关系(R~2=0.93),且树干呼吸的温度敏感性系数(Q_(10))为2.62;不同季节毛白杨树干呼吸的Q_(10)存在差异,生长季的Q_(10)(1.95)明显低于非生长季(3.00),表明生长呼吸和维持呼吸对温度的响应也并不相同。(3)温度矫正后的毛白杨树干呼吸(R_(15))在昼夜和季节尺度上均存在明显的变异,即夜晚的R_(15)显著高于白天(P<0.01),生长季的R_(15)明显高于非生长季(P<0.05);树干可溶性糖含量与生长季的R_(15)存在较好的相关性(R~2=0.52),而非生长季的R_(15)却主要受到树干淀粉含量的影响。研究结果表明,在生长季,毛白杨树干呼吸的在日变化主要受到温度的影响,而在季节尺度上Q_(10)的变异则与树干呼吸中维持呼吸所占比例及树干中非结构性碳水化合物(可溶性糖和淀粉)的含量及类型紧密相关。
Stem respiration plays a pivotal role in forest ecosystem carbon cycling, and understanding further its response to future climate warming has important significance for accurately estimating the carbon storage of forest ecosystems. To examine the seasonal variations of stem respiration of Chinese white poplar(Populus tomentosa), we measured stem respiration rate and stem temperature using the horizontally oriented soil chamber technique, from January 2016 to January 2017. We also conducted a 24 h measurement to examine the diurnal variation of stem respiration in May 2016. We found that(1) the stem respiration rate was positively related to stem temperature at night time(R~2 = 0.88), whereas the relationship between stem respiration rate and stem temperature featured a negative linear curve during day time(R~2 = 0.96);(2) the seasonal dynamics of the stem respiration rate followed the change of the stem temperature with a bell-shaped curve, and stem temperature explained 93% of the seasonal variation of stem respiration rate, with the temperature sensitivity(Q_(10)) of 2.62 over the entire observation period. The Q_(10) was obviously different among seasons, with a lower value found in the non-growing season(3.00), and a higher value in the growing season(1.95), indicating that the growth and maintenance respiration had different temperature responses;(3) the temperature-normalised stem respiration(R_(15)) also featured obviously diurnal and seasonal variations, where the R_(15) at night time and during the growing season were significantly higher than those of the daytime(P<0.01), and non-growing season(P<0.05). The principal factor responsible for the seasonal variation of R_(15) was the soluble sugar content in the growing season, and the starch content in the non-growing season. These results suggested that the stem respiration rate of Chinese white poplar was mainly disturbed by temperature at the diurnal scale, and closely related to the content and types of non-structural carbohydrates(soluble sugar and starch) and respiration component at seasonal scales, besides temperature.
Stem respiration plays a pivotal role in forest ecosystem carbon cycling, and understanding further its response to future climate warming has important significance for accurately estimating the carbon storage of forest ecosystems. To examine the seasonal variations of stem respiration of Chinese white poplar(Populus tomentosa), we measured stem respiration rate and stem temperature using the horizontally oriented soil chamber technique, from January 2016 to January 2017. We also conducted a 24 h measurement to examine the diurnal variation of stem respiration in May 2016. We found that(1) the stem respiration rate was positively related to stem temperature at night time(R~2 = 0.88), whereas the relationship between stem respiration rate and stem temperature featured a negative linear curve during day time(R~2 = 0.96);(2) the seasonal dynamics of the stem respiration rate followed the change of the stem temperature with a bell-shaped curve, and stem temperature explained 93% of the seasonal variation of stem respiration rate, with the temperature sensitivity(Q_(10)) of 2.62 over the entire observation period. The Q_(10) was obviously different among seasons, with a lower value found in the non-growing season(3.00), and a higher value in the growing season(1.95), indicating that the growth and maintenance respiration had different temperature responses;(3) the temperature-normalised stem respiration(R_(15)) also featured obviously diurnal and seasonal variations, where the R_(15) at night time and during the growing season were significantly higher than those of the daytime(P<0.01), and non-growing season(P<0.05). The principal factor responsible for the seasonal variation of R_(15) was the soluble sugar content in the growing season, and the starch content in the non-growing season. These results suggested that the stem respiration rate of Chinese white poplar was mainly disturbed by temperature at the diurnal scale, and closely related to the content and types of non-structural carbohydrates(soluble sugar and starch) and respiration component at seasonal scales, besides temperature.
引文
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[8] 赵广,刘刚才,朱万泽.贡嘎山峨眉冷杉树干呼吸空间特征及其对温度的响应.生态学报,2018,38(8):2732- 2742.
[9] 黄玮,朱锦懋,阮宏华,黄儒珠,王健,郑怀舟.树干CO2释放速率影响因素研究进展.生态学杂志,2010,29(4):790- 797.
[10] McGuire M A,Cerasoli S,Teskey R O.CO2 fluxes and respiration of branch segments of sycamore (Platanus occidentalis L.) examined at different sap velocities,branch diameters,and temperatures.Journal of Experimental Botany,2007,58(8):2159- 2168.
[11] 许飞,王传宽.温带4种针叶树种春、秋季节树干维持呼吸的日动态.生态学报,2015,35(10):3233- 3243.
[12] 魏国军,盛浩,杨智杰,袁一丁,杨玉盛.亚热带4种行道树树干表面CO2释放速率昼夜动态.亚热带资源与环境学报,2009,4(1):23- 31.
[13] Lavigne M B.Comparing stem respiration and growth of jack pine provenances from northern and southern locations.Tree Physiology,1996,16(10):847- 852.
[14] 马玉娥,项文化,雷丕锋.林木树干呼吸变化及其影响因素研究进展.植物生态学报,2007,31(3):403- 412.
[15] Ceschia E,Damesin C,Lebaube S,Pontailler J Y,Dufrêne é.Spatial and seasonal variations in stem respiration of beech trees (Fagus sylvatica).Annals of Forest Science,2002,59(8):801- 812.
[16] Saveyn A,Steppe K,McGuire M A,Lemeur R,Teskey R O.Stem respiration and carbon dioxide efflux of young Populus deltoides trees in relation to temperature and xylem carbon dioxide concentration.Oecologia,2008,154(4):637- 649.
[17] Damesin C,Ceschia E,Le Goff N,Ottorini J M,Dufrêne E.Stem and branch respiration of beech:from tree measurements to estimations at the stand level.New Phytologist,2002,153(1):159- 172.
[18] Yang Q P,Xu M,Chi Y G,Zheng Y P.Relationship between stem CO2 efflux and stem temperature at different measuring depths in Pinus massoniana trees.Acta Ecologica Sinica,2016,36(4):229- 235.
[19] Maier C A.Stem growth and respiration in loblolly pine plantations differing in soil resource availability.Tree Physiology,2001,21(16):1183- 1193.
[20] Brito P,Morales D,Wieser G,Jiménez M S.Spatial and seasonal variations in stem CO2 ef?ux of Pinus canariensis at their upper distribution limit.Trees,2010,24(3):523- 531.
[21] Maier C A,Johnsen K H,Clinton B D,Ludovici K H.Relationships between stem CO2 efflux,substrate supply,and growth in young loblolly pine trees.New Phytologist,2010,185(2):502- 513.
[22] Lavigne M B.Differences in stem respiration responses to temperature between balsam fir trees in thinned and unthinned stands.Tree Physiology,1987,3(3):225- 233.
[23] Davidson E A,Belk E,Boone R D.Soil water content and temperature as independent or confounded factors controlling soil respiration in a temperate mixed hardwood forest.Global Change Biology,1998,4(2):217- 227.
[24] 刘潇潇,何秋月,闫美杰,李国庆,王仕稳,杜盛.黄土丘陵区辽东栎群落优势种和主要伴生种树干液流动态特征.生态学报,2018,38(13):4744- 4751.
[25] 王淼,姬兰柱,李秋荣,肖冬梅,刘海良.长白山地区红松树干呼吸的研究.应用生态学报,2005,16(1):7- 13.
[26] 陆彬.小兴安岭典型森林群落生长季土壤呼吸及树干呼吸特征[D].哈尔滨:东北林业大学,2010.
[27] Zhu L W,Zhao P,Cai X A,Zeng X P,Ni G Y,Zhang J Y,Zou L L,Mei T T,Yu M H.Effects of sap velocity on the daytime increase of stem CO2 efflux from stems of Schima superba trees.Trees,2012,26(2):535- 542.
[28] 王秀伟,毛子军,孙涛,吴海军.春、秋季节树干温度和液流速度对东北3树种树干表面CO2释放通量的影响.生态学报,2011,31(12):3358- 3367.
[29] 许飞,王传宽.4种温带针叶树种树干CO2释放通量的季节动态及其驱动因子.植物生态学报,2017,41(4):396- 408.
[30] McGuire M A,Teskey R O.Microelectrode technique for in situ measurement of carbon dioxide concentrations in xylem sap of trees.Tree Physiology,2002,22(11):807- 811.
[31] Teskey R O,Mcguire M A.Carbon dioxide transport in xylem causes errors in estimation of rates of respiration in stems and branches of trees.Plant,Cell & Environment,2010,25(11):1571- 1577.
[32] Ryan M G.Growth and maintenance respiration in stems of Pinus contorta and Picea engelmannii.Canadian Journal of Forest Research,1990,20(1):48- 57.
[33] 石新立,王传宽,许飞,王兴昌.四个温带树种树干呼吸的时间动态及其影响因子.生态学报,2010,30(15):3994- 4003.
[34] Stockfors J.Temperature variations and distribution of living cells within tree stems:implications for stem respiration modeling and scale-up.Tree Physiology,2000,20(15):1057- 1062.
[35] Bosc A,De Grandcourt A,Loustau D.Variability of stem and branch maintenance respiration in a Pinus pinaster tree.Tree Physiology,2003,23(4):227- 236.
[36] Bader M K F,Mildner M,Baumann C,Leuzinger S,K?rner C.Photosynthetic enhancement and diurnal stem and soil carbon fluxes in a mature Norway spruce stand under elevated CO2.Environmental and Experimental Botany,2016,124:110- 119.
[37] Regier N,Streb S,Zeeman S C,Frey B.Seasonal changes in starch and sugar content of poplar (Populus deltoides × nigra cv.Dorskamp) and the impact of stem girdling on carbohydrate allocation to roots.Tree Physiology,2010,30(8):979- 987.
[2] King A W,Gunderson C A,Post W M,Weston D J,Wullschleger S D.Plant respiration in a warmer world.Science,2006,312(5773):536- 537.
[3] da Costa A C L,Metcalfe D B,Doughty C E,de Oliveira A A R,Neto G F C,da Costa M C,de Athaydes Silva Junior J,Arag?o L E O C,Almeida S,Galbraith D R,Rowland L M,Meir P,Malhi Y.Ecosystem respiration and net primary productivity after 8- 10 years of experimental through-fall reduction in an eastern Amazon forest.Plant Ecology & Diversity,2014,7(1- 2):7- 24.
[4] 涂洁,樊后保,王勇刚,李志鹏.各径级马尾松树干CO2释放速率及其温度敏感性.林业科学,2017,53(10):154- 159.
[5] Rowland L,Hill T C,Stahl C,Siebicke L,Burban B,Zaragoza-Castells J,Ponton S,Bonal D,Meir P,Williams M.Evidence for strong seasonality in the carbon storage and carbon use efficiency of an Amazonian forest.Global Change Biology,2014,20(3):979- 991.
[6] Xu M,DeBiase T A,Qi Y,Goldstein A,Liu Z G.Ecosystem respiration in a young ponderosa pine plantation in the Sierra Nevada Mountains,California.Tree Physiology,2001,21(5):309- 318.
[7] Yang Y,Zhao M,Xu X T,Sun Z Z,Yin G D,Piao S L.Diurnal and seasonal change in stem respiration of Larix principis-rupprechtii Trees,Northern China.PLoS One,2014,9(2):e89294.
[8] 赵广,刘刚才,朱万泽.贡嘎山峨眉冷杉树干呼吸空间特征及其对温度的响应.生态学报,2018,38(8):2732- 2742.
[9] 黄玮,朱锦懋,阮宏华,黄儒珠,王健,郑怀舟.树干CO2释放速率影响因素研究进展.生态学杂志,2010,29(4):790- 797.
[10] McGuire M A,Cerasoli S,Teskey R O.CO2 fluxes and respiration of branch segments of sycamore (Platanus occidentalis L.) examined at different sap velocities,branch diameters,and temperatures.Journal of Experimental Botany,2007,58(8):2159- 2168.
[11] 许飞,王传宽.温带4种针叶树种春、秋季节树干维持呼吸的日动态.生态学报,2015,35(10):3233- 3243.
[12] 魏国军,盛浩,杨智杰,袁一丁,杨玉盛.亚热带4种行道树树干表面CO2释放速率昼夜动态.亚热带资源与环境学报,2009,4(1):23- 31.
[13] Lavigne M B.Comparing stem respiration and growth of jack pine provenances from northern and southern locations.Tree Physiology,1996,16(10):847- 852.
[14] 马玉娥,项文化,雷丕锋.林木树干呼吸变化及其影响因素研究进展.植物生态学报,2007,31(3):403- 412.
[15] Ceschia E,Damesin C,Lebaube S,Pontailler J Y,Dufrêne é.Spatial and seasonal variations in stem respiration of beech trees (Fagus sylvatica).Annals of Forest Science,2002,59(8):801- 812.
[16] Saveyn A,Steppe K,McGuire M A,Lemeur R,Teskey R O.Stem respiration and carbon dioxide efflux of young Populus deltoides trees in relation to temperature and xylem carbon dioxide concentration.Oecologia,2008,154(4):637- 649.
[17] Damesin C,Ceschia E,Le Goff N,Ottorini J M,Dufrêne E.Stem and branch respiration of beech:from tree measurements to estimations at the stand level.New Phytologist,2002,153(1):159- 172.
[18] Yang Q P,Xu M,Chi Y G,Zheng Y P.Relationship between stem CO2 efflux and stem temperature at different measuring depths in Pinus massoniana trees.Acta Ecologica Sinica,2016,36(4):229- 235.
[19] Maier C A.Stem growth and respiration in loblolly pine plantations differing in soil resource availability.Tree Physiology,2001,21(16):1183- 1193.
[20] Brito P,Morales D,Wieser G,Jiménez M S.Spatial and seasonal variations in stem CO2 ef?ux of Pinus canariensis at their upper distribution limit.Trees,2010,24(3):523- 531.
[21] Maier C A,Johnsen K H,Clinton B D,Ludovici K H.Relationships between stem CO2 efflux,substrate supply,and growth in young loblolly pine trees.New Phytologist,2010,185(2):502- 513.
[22] Lavigne M B.Differences in stem respiration responses to temperature between balsam fir trees in thinned and unthinned stands.Tree Physiology,1987,3(3):225- 233.
[23] Davidson E A,Belk E,Boone R D.Soil water content and temperature as independent or confounded factors controlling soil respiration in a temperate mixed hardwood forest.Global Change Biology,1998,4(2):217- 227.
[24] 刘潇潇,何秋月,闫美杰,李国庆,王仕稳,杜盛.黄土丘陵区辽东栎群落优势种和主要伴生种树干液流动态特征.生态学报,2018,38(13):4744- 4751.
[25] 王淼,姬兰柱,李秋荣,肖冬梅,刘海良.长白山地区红松树干呼吸的研究.应用生态学报,2005,16(1):7- 13.
[26] 陆彬.小兴安岭典型森林群落生长季土壤呼吸及树干呼吸特征[D].哈尔滨:东北林业大学,2010.
[27] Zhu L W,Zhao P,Cai X A,Zeng X P,Ni G Y,Zhang J Y,Zou L L,Mei T T,Yu M H.Effects of sap velocity on the daytime increase of stem CO2 efflux from stems of Schima superba trees.Trees,2012,26(2):535- 542.
[28] 王秀伟,毛子军,孙涛,吴海军.春、秋季节树干温度和液流速度对东北3树种树干表面CO2释放通量的影响.生态学报,2011,31(12):3358- 3367.
[29] 许飞,王传宽.4种温带针叶树种树干CO2释放通量的季节动态及其驱动因子.植物生态学报,2017,41(4):396- 408.
[30] McGuire M A,Teskey R O.Microelectrode technique for in situ measurement of carbon dioxide concentrations in xylem sap of trees.Tree Physiology,2002,22(11):807- 811.
[31] Teskey R O,Mcguire M A.Carbon dioxide transport in xylem causes errors in estimation of rates of respiration in stems and branches of trees.Plant,Cell & Environment,2010,25(11):1571- 1577.
[32] Ryan M G.Growth and maintenance respiration in stems of Pinus contorta and Picea engelmannii.Canadian Journal of Forest Research,1990,20(1):48- 57.
[33] 石新立,王传宽,许飞,王兴昌.四个温带树种树干呼吸的时间动态及其影响因子.生态学报,2010,30(15):3994- 4003.
[34] Stockfors J.Temperature variations and distribution of living cells within tree stems:implications for stem respiration modeling and scale-up.Tree Physiology,2000,20(15):1057- 1062.
[35] Bosc A,De Grandcourt A,Loustau D.Variability of stem and branch maintenance respiration in a Pinus pinaster tree.Tree Physiology,2003,23(4):227- 236.
[36] Bader M K F,Mildner M,Baumann C,Leuzinger S,K?rner C.Photosynthetic enhancement and diurnal stem and soil carbon fluxes in a mature Norway spruce stand under elevated CO2.Environmental and Experimental Botany,2016,124:110- 119.
[37] Regier N,Streb S,Zeeman S C,Frey B.Seasonal changes in starch and sugar content of poplar (Populus deltoides × nigra cv.Dorskamp) and the impact of stem girdling on carbohydrate allocation to roots.Tree Physiology,2010,30(8):979- 987.