香樟幼龄林不同叶龄叶片的光合特征和单萜释放规律
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摘要
通过调查亚热带地区重要树种香樟(Cinnamomum camphora)叶片生长发育中幼龄、成熟和老龄3个阶段的光合生理与植物挥发性有机化合物(Plant biogenic volatile organic compounds,BVOCs)单萜释放规律,研究光合光电子输运、CO2固定对单萜释放的影响和调控规律.光合生理特征参数调查采用非直角双曲线模型和Farquhar模型,单萜释放特征参数调查采用Guenther BVOCs排放模型.结果显示:(1)香樟叶片释放的单萜以蒈烯和罗勒烯为主.在本研究的基础状态下(温度25℃,光照强度800μmol m~(-2) s~(-1)和CO_2浓度400×10~(-6)),净光合速率(P_n)、气孔导度(G_s)、蒸腾速率(T_r)和胞间CO_2浓度(C_i)大小规律为成熟叶>幼龄叶>衰老叶,总单萜释放速率(E)为幼龄叶>衰老叶>成熟叶.(2)叶片3个叶龄阶段的单萜-光响应曲线变化趋势与光合的光响应曲线类似,但受CO2浓度的影响不明显.成熟叶具有最高的初始量子效率(α)、最大净光合速率(P_(nmax))、Rubiscom酶最大羧化速率(V_(cmax))、光下呼吸速率(R_p)和光饱和点(LSP),但是幼龄叶的光补偿点(LCP)、暗呼吸速率(Rd)和响应曲线的曲角系数(θ)最大.衰老叶具有最高的最大电子传递速率(J_(max))和最低V_(cmax),因此其Jmax/Vcmax比率为3个阶段最高,意味着衰老叶光能利用率最低,强光下光能过剩,这很可能也是常绿植物冬天利用过剩光能保持一定温度越冬的一个调控机制.(3)幼龄叶的最大单萜排放速率(E_(max))最大,其次为衰老叶,成熟叶最小.在3个阶段中,成熟叶的表观量子效率(β)和真实量子效率(βT)均是最大,幼龄叶的β最小,但βT却仅次于成熟叶.综上所述,香樟叶片单萜的释放受叶片叶龄不同影响显著,具有显著的光依赖特性,对CO_2浓度的响应不明显;香樟叶片单萜释放水平很可能受自身生长过程中能量和碳源供应的调控,也受不同生长季节环境因子变化的影响,结果可为区域性BVOCs释放模型提供有益的参数借鉴.
Plant biogenic volatile organic compounds(BVOCs) emission is driven by environmental factors and ontogeny stages. Cinnamomum camphora, an evergreen tree species widely distributed and used for urban greening in south of China, emits multiple BVOCs, including monoterpenes. However, the regulation of how photosynthetic physiology affects monoterpenes emission is poorly understood. In this study, we investigated the photosynthetic characteristics and monoterpenes emission of C. camphora under three foliar ontogeny stages—young, mature, and old—to gain insight into whether a direct regulatory relationship exists between monoterpenes emission and photosynthetic CO_2 assimilation. The photosynthetic characteristics were estimated by a light/CO_2 response curve using the non-rectangular hyperbola and Farquhar photosynthesis model. All monoterpenes emission parameters were estimated by using the Guenther BVOCs emission model.The main monoterpenes emitted from C. camphora were careen and ocimene. The parameters of net photosynthesis rate(P_n),stomatal conductance(G_s), transpiration rate(T_r), and intercellular CO_2 concentration(C_i) were significantly different in the three foliar ontogeny stages, when measured under standard conditions, and the highest and lowest values of these parameters were found in the mature and old foliage, respectively. However, the highest total monoterpenes emission rate(E) was found in the young foliage and the lowest emission rate in the mature foliage. The response tendency of total monoterpenes emission with light intensity was similar to photosynthesis-light intensity response curves under all three foliar ontogeny stages, while the response to CO_2 concentration were no significant impacts on total monoterpenes emission. The mature foliage had the highest initial quantum efficiency(α), maximum net photosynthetic rate(P_(nmax)), maximum carboxylase activity of Rubisco(V_(cmax)), respiration rate under light(Rp), and light saturation point(LSP). However, the highest values of the light compensation point(LCP) and dark respiration rate(Rd) were found in the young foliage. The highest value of the capacity of photosynthetic electron transport(J_(max)) and the lowest value of the capacity of Vcmax were found in old foliage. This meant that old foliage had the lowest light energy utilization and there was excess light energy under strong light, which was probably also a regulatory mechanism for the evergreen plants to use excess light energy to maintain a certain temperature to overwinter. The highest maximum monoterpenes emission rate(E_(max)) and the lowest apparent quantum efficiency(β) for total monoterpenes emission were found in young foliage; however, the lowest Emax and the highest apparent quantum efficiency(β) and true quantum yield(β_T) and for total monoterpenes emission were found in mature foliage. We conclude that monoterpenes emission from C.camphora is mainly dependent on foliage ontogeny; however, the emission rate was significantly affected by light response,but not by CO_2 concentration, indicating that the total monoterpene emission could be regulated by energy and carbon source supply during the growth process and also influenced by the changes of environmental factors in different seasons. This study will be useful in precisely determining the emission parameters for a regional model in BVOCs emission estimation.
Plant biogenic volatile organic compounds(BVOCs) emission is driven by environmental factors and ontogeny stages. Cinnamomum camphora, an evergreen tree species widely distributed and used for urban greening in south of China, emits multiple BVOCs, including monoterpenes. However, the regulation of how photosynthetic physiology affects monoterpenes emission is poorly understood. In this study, we investigated the photosynthetic characteristics and monoterpenes emission of C. camphora under three foliar ontogeny stages—young, mature, and old—to gain insight into whether a direct regulatory relationship exists between monoterpenes emission and photosynthetic CO_2 assimilation. The photosynthetic characteristics were estimated by a light/CO_2 response curve using the non-rectangular hyperbola and Farquhar photosynthesis model. All monoterpenes emission parameters were estimated by using the Guenther BVOCs emission model.The main monoterpenes emitted from C. camphora were careen and ocimene. The parameters of net photosynthesis rate(P_n),stomatal conductance(G_s), transpiration rate(T_r), and intercellular CO_2 concentration(C_i) were significantly different in the three foliar ontogeny stages, when measured under standard conditions, and the highest and lowest values of these parameters were found in the mature and old foliage, respectively. However, the highest total monoterpenes emission rate(E) was found in the young foliage and the lowest emission rate in the mature foliage. The response tendency of total monoterpenes emission with light intensity was similar to photosynthesis-light intensity response curves under all three foliar ontogeny stages, while the response to CO_2 concentration were no significant impacts on total monoterpenes emission. The mature foliage had the highest initial quantum efficiency(α), maximum net photosynthetic rate(P_(nmax)), maximum carboxylase activity of Rubisco(V_(cmax)), respiration rate under light(Rp), and light saturation point(LSP). However, the highest values of the light compensation point(LCP) and dark respiration rate(Rd) were found in the young foliage. The highest value of the capacity of photosynthetic electron transport(J_(max)) and the lowest value of the capacity of Vcmax were found in old foliage. This meant that old foliage had the lowest light energy utilization and there was excess light energy under strong light, which was probably also a regulatory mechanism for the evergreen plants to use excess light energy to maintain a certain temperature to overwinter. The highest maximum monoterpenes emission rate(E_(max)) and the lowest apparent quantum efficiency(β) for total monoterpenes emission were found in young foliage; however, the lowest Emax and the highest apparent quantum efficiency(β) and true quantum yield(β_T) and for total monoterpenes emission were found in mature foliage. We conclude that monoterpenes emission from C.camphora is mainly dependent on foliage ontogeny; however, the emission rate was significantly affected by light response,but not by CO_2 concentration, indicating that the total monoterpene emission could be regulated by energy and carbon source supply during the growth process and also influenced by the changes of environmental factors in different seasons. This study will be useful in precisely determining the emission parameters for a regional model in BVOCs emission estimation.
引文
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10曹雪丹,李文华,鲁周民,张忠良,吴万兴.北缘地区枇杷春季光合特性研究[J].西北林学院学报,2008,23(6):33-37[Cao XD,Li WH,Lu ZM,Zhang ZL,Wu WX.The photosynthetic characteristics of Eriobotrya japonica in northern marginal cultivation area in spring[J].JNW For Univ,2008,23(6):33-37]
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