杉木幼苗光合及叶绿素荧光特征对氮沉降的短期响应

doi:10.11924/j.issn.1000-6850.casb2021-1035

摘要/Abstract

摘要：

为了探究模拟氮沉降条件下杉木幼苗光合及叶绿素荧光特征变化规律,从光合生理的角度探讨不同季节杉木幼苗对氮沉降的短期响应。选取杉木幼苗为实验对象,模拟氮沉降实验,设置4个处理水平对照(N0)、低氮（N30:30 kg/(hm²·a)、中氮（N60:60 kg/(hm²·a)、高氮(N90:90 kg/(hm²·a)。结果表明：添加氮以后,杉木幼苗的净光合速率(Pn),初始荧光产量(F₀)、最大荧光产量(F_m)、最大PSII光能转换效率(F_v/F_m)、PSII潜在活性(F_v/F₀)显著降低,水分利用率(Wue)显著增高(P<0.05)。随着氮沉降水平的增加,气孔导度(Gs)与蒸腾速率(Tr)呈现先升高后下降趋势,水分利用率呈现先下降后升高趋势。在中氮和高氮处理下,F_v/F_m、F_v/F₀值显著降低(P<0.05)。在冬季,净光合速率与气孔导度、蒸腾速率和胞间二氧化碳浓度呈极显著正相关(P<0.01),春季,净光合速率与气孔导度和蒸腾速率呈极显著正相关(P<0.01),与胞间二氧化碳浓度呈负相关。氮沉降累积达一定阈值后,增加蒸腾失水,从而降低植株的水分利用效率。在冬季,净光合速率主要受到气孔限制的影响,热耗散增加,PSII反应中心出现光抑制现象,到春季,过量的氮沉降累积破坏了幼苗的光合结构,导致胞间二氧化碳浓度上升而净光合速率下降,非气孔限制成为影响净光合速率的主要原因。中氮处理可能使叶绿体中PSII潜在活性减弱,光反应中心光合电子的传递能力降低,导致光合效率下降。

关键词: 氮沉降, 杉木幼苗, 气体交换参数, 叶绿素荧光

Abstract:

To explore the change law of the photosynthesis and chlorophyll fluorescence characteristics of fir seedlings under simulated nitrogen deposition condition, the short-term response of fir seedlings to nitrogen deposition in different seasons was discussed from the perspective of photosynthesis physiology. The fir seedlings were selected as experimental subjects, and the nitrogen deposition experiment was simulated with four treatment levels of the control (N0), low nitrogen (N30:30 kg/(hm².a)), medium nitrogen (N60:60 kg/(hm².a)), and high nitrogen(N90:90 kg/(hm².a)). The results showed that after adding nitrogen, the net photosynthesis rate (Pn), initial fluorescence yield (F₀), maximum fluorescence yield (F_m), maximum PSII light energy conversion efficiency (F_v/F_m) and PSII potential activity (F_v/F₀) were significantly reduced, while water utilization rate (Wue) increased significantly (P<0.05). With the increase of nitrogen deposition level, stomatal conductance (Gs) and transpiration rate (Tr) showed a tendency of increase first and then decrease, and the water use efficiency showed a trend of first decrease then increase. With the medium and high nitrogen treatment, F_v/F_m and F_v/F₀ values were significantly reduced (P<0.05). In winter, the net photosynthesis rate was significantly and positively correlated with stomatal conductance, transpiration rate and intercellular CO2 concentration (P<0.01); and in spring, the net photosynthesis rate was significantly and positively correlated with stomatal conductance and transpiration rate (P<0.01), and negatively correlated with intercellular CO2 concentration. After the accumulation of nitrogen deposition reached a certain threshold, the water loss was increased, which reduced the water utilization efficiency of the plant. In winter, the net photosynthesis rate was mainly affected by the stomatal limit, heat dissipation increased, and the PSII reaction center showed photo-suppression phenomenon; when it turned to spring, the excessive accumulation of nitrogen deposition destroyed the photosynthesis structure of seedlings, resulting in an increase in the concentration of carbon dioxide between cells and a decrease in the net photosynthesis rate. Then the non-stomatal limits became the main reason for the impact on the net photosynthesis rate. The medium nitrogen treatment could reduce the potential activity of PSII in chlorophyll, and the photosynthetic electron transfer capacity of the photoreaction center decreases, resulting in the decrease of photosynthesis efficiency.

Key words: nitrogen deposition, fir seedling, gas exchange parameters, chlorophyll fluorescence parameters

中图分类号:

赵佼娇, 哀建国. 杉木幼苗光合及叶绿素荧光特征对氮沉降的短期响应[J]. 中国农学通报, 2022, 38(29): 67-73.

ZHAO Jiaojiao, AI Jianguo. Short-term Response of Photosynthesis and Chlorophyll Fluorescence Characteristics of Fir Seedlings to Nitrogen Deposition[J]. Chinese Agricultural Science Bulletin, 2022, 38(29): 67-73.

图/表 8

图1. 不同梯度水平氮沉降处理下冬季与春季杉木幼苗的净光合速率注：不同大写字母表示冬季不同氮沉降水平处理间的差异性显著(P<0.05)。不同小写字母表示夏季不同氮沉降水平处理间的差异性显著(P<0.05),CK为对照组,LN为低氮组,MN为中氮组,HN为高氮组。

图2. 不同梯度水平氮沉降处理下冬季与春季杉木幼苗的胞间二氧化碳浓度注：不同大写字母表示冬季不同氮沉降水平处理间的差异性显著(P<0.05)。不同小写字母表示夏季不同氮沉降水平处理间的差异性显著(P<0.05),CK为对照组,LN为低氮组,MN为中氮组,HN为高氮组。

图3. 不同梯度水平氮沉降处理下冬季与春季杉木幼苗的气孔导度注：不同大写字母表示冬季不同氮沉降水平处理间的差异性显著(P<0.05)。不同小写字母表示夏季不同氮沉降水平处理间的差异性显著(P<0.05),CK为对照组,LN为低氮组,MN为中氮组,HN为高氮组。

图4. 不同梯度水平氮沉降处理下冬季与春季杉木幼苗的蒸腾速率注：不同大写字母表示冬季不同氮沉降水平处理间的差异性显著(P<0.05)。不同小写字母表示夏季不同氮沉降水平处理间的差异性显著(P<0.05),CK为对照组,LN为低氮组,MN为中氮组,HN为高氮组。

图5. 不同氮沉降处理下冬季与春季杉木幼苗的水分利用率注：不同大写字母表示冬季不同氮沉降水平处理间的差异性显著(P<0.05)。不同小写字母表示夏季不同氮沉降水平处理间的差异性显著(P<0.05),CK为对照组,LN为低氮组,MN为中氮组,HN为高氮组。

指标	Pn	Ci	Gs	Tr	Wue
Pn	1
Ci	0.617**	1
Gs	0.9**	0.427	1
Tr	0.907**	0.407	0.977**	1
Wue	-0.516*	0.054	-0.79**	0.824**	1

表1. 冬季氮沉降处理下杉木幼苗光合气体参数间的Pearson相关性分析

指标	Pn	Ci	Gs	Tr	Wue
Pn	1
Ci	-0.221	1
Gs	0.927**	0.044	1
Tr	0.930**	0.092	0.986**	1
Wue	0.014	0.805**	-0.323	-0.349	1

表2. 春季氮沉降处理下杉木幼苗光合气体参数间的Pearson相关性分析

图6. 不同梯度水平氮沉降处理下冬季与春季杉木幼苗的各荧光参数注：不同大写字母表示冬季不同氮沉降水平处理间的差异性显著(P<0.05)。不同小写字母表示夏季不同氮沉降水平处理间的差异性显著(P<0.05),CK为对照组,LN为低氮组,MN为中氮组,HN为高氮组。

参考文献 50

[1]	LU Me, YANG. Responses of ecosystem nitrogen cycle to nitrogen addition: a meta-analysis[J]. New phytologist, 2010, 189(4):1040-1050. doi: 10.1111/j.1469-8137.2010.03563.x URL
[2]	于美佳, 叶彦辉, 韩艳英, 等. 氮沉降对森林生态系统影响的研究进展[J]. 安徽农业科学, 2021, 49(3):19-24,27.
[3]	赵娜, 贾力, 刘洪来, 等. 氮肥的环境风险及管理研究进展[J]. 草原与草坪, 2011, 31(1):89-93,96.
[4]	李洁, 薛立. 氮磷沉降对森林土壤生化特性影响研究进展[J]. 世界林业研究, 2017, 30(2):14-19.
[5]	PALANI S, TKALICH P, et al. ANN application for prediction of atmospheric nitrogen deposition to aquatic ecosystems[J]. Marine pollution bulletin, 2011, 62(6):1198-1206. doi: 10.1016/j.marpolbul.2011.03.033 pmid: 21481425
[6]	CORNELL S E, et al. Atmospheric nitrogen deposition: Revisiting the question of the importance of the organic component[J]. Environmental Pollution, 2011, 159(10):2214-2222. doi: 10.1016/j.envpol.2010.11.014 pmid: 21131113
[7]	GUAN L L, WEN D Z. More nitrogen partition in structural proteins and decreased photosynthetic nitrogen-use efficiency of Pinus massoniana under in situ polluted stress[J]. Journal of plant research, 2011, 124(6):663-673. doi: 10.1007/s10265-011-0405-2 URL
[8]	KATTGE J, KNORR W, RADDATZ T, et al. Quantifying photosynthetic capacity and its relationship to leaf nitrogen content for global-scale terrestrial biosphere models[J]. Global Change biology, 2009, 15(4):976-991. doi: 10.1111/j.1365-2486.2008.01744.x URL
[9]	顾鸿昊, 翁俊, 孔佳杰, 等. 粗放和集约经营毛竹林叶片的生态化学计量特征[J]. 浙江农林大学学报, 2015, 32(5):661-667.
[10]	LIU X J, Ying, et al. Enhanced nitrogen deposition over China[J]. Nature, 2013, 494(Feb.1TN.7438):459-459.
[11]	LI Q, SONG X, CHANG S X, et al. Nitrogen depositions increase soil respiration and decrease temperature sensitivity in a Moso bamboo forest[J]. Agricultural and forest meteorology, 2019, 268:48-54. doi: 10.1016/j.agrformet.2019.01.012 URL
[12]	BOXMAN A W, BLANCK K, BRANDRUD T E. Vegetation and soil biota response to experimentally-changed nitrogen inputs in coniferous forest ecosystems of the NIREX project[J]. Forest ecology and management, 1998, 101(1):65-79. doi: 10.1016/S0378-1127(97)00126-6 URL
[13]	LÜ C, TIAN H, et al. Spatial and temporal patterns of nitrogen deposition in China: Synthesis of observational data[J]. Journal of geophysical research atmospheres, 2007, 112(22):1-10.
[14]	BLANES M C, EMMETT B A. Alleviation of P limitation makes tree roots competitive for N against microbes in a N-saturated conifer forest: A test through P fertilization and 15N labelling[J]. Soil biology & biochemistry, 2012, 48(none):51-59. doi: 10.1016/j.soilbio.2012.01.012 URL
[15]	DENG M, LIU L, SUN Z, et al. Increased phosphate uptake but not resorption alleviates phosphorus deficiency induced by nitrogen deposition in temperate Larix principis-rupprechtii plantations[J]. New phytologist, 2016, 212(4):1019-1029. doi: 10.1111/nph.14083 pmid: 27400237
[16]	FANG X M, ZHANG X L, ZONG Y Y, et al. Soil phosphorus functional fractions and tree tissue nutrient concentrations influenced by stand density in subtropical Chinese fir plantation forests[J]. PLoS one, 2017, 12(10):e0186905.
[17]	涂利华, 戴洪忠, 胡庭兴, 等. 模拟氮沉降对华西雨屏区撑绿杂交竹林土壤呼吸的影响[J]. 应用生态学报, 2011, 22(4):829-836.
[18]	朱敏, 张振华, 于君宝, 等. 氮沉降对黄河三角洲芦苇湿地土壤呼吸的影响[J]. 植物生态学报, 2013, 37(6):517-529. doi: 10.3724/SP.J.1258.2013.00053
[19]	林雨萱, 哀建国, 宋新章, 等. 模拟氮沉降和磷添加对杉木林土壤呼吸的影响[J]. 浙江农林大学学报, 2021, 38(3):494-501.
[20]	沈芳芳, 袁颖红, 樊后保, 等. 氮沉降对杉木人工林土壤有机碳矿化和土壤酶活性的影响[J]. 生态学报, 2012, 32(2):517-527.
[21]	刘滔, 尹光彩, 刘菊秀, 等. 酸沉降对南亚热带森林土壤主要元素的影响[J]. 应用与环境生物学报, 2013, 19(2):255-261.
[22]	黄婷, 包和林, 吴承祯, 等. 氮-硫沉降对邓恩桉及杉木人工林凋落物C和N残留率的影响[J]. 植物资源与环境学报, 2013, 22(4):11-19.
[23]	王巧, 张君波, 雷赵枫, 等. 模拟氮沉降和磷添加对杉木生态化学计量学特征的影响[J]. 生态学杂志, 2019, 38(2):368-375.
[24]	杨春菊. 临安市马尾松林分因子与地形关系的研究[D]. 杭州: 浙江农林大学, 2016.
[25]	REAY D S, F DENTENER, SMITH P, et al. Global nitrogen deposition and carbon sinks[J]. Nature Geoscience, 2008, 1(7):430-437. doi: 10.1038/ngeo230 URL
[26]	GRASSI G, MEIR P, CROMER R, et al. Photosynthetic parameters in seedlings of Eucalyptus grandis as affected by rate of nitrogen supply[J]. Plant cell & environment, 2010, 25(12):1677-1688.
[27]	张维, 贺亚玲, 吴泽昂, 等. 模拟增温对梭梭光合生理生态特征的影响[J]. 草地学报, 2017, 25(2):296-302.
[28]	田艳丽, 种培芳, 陆文涛, 等. 不同光合途径植物红砂和珍珠猪毛菜幼苗对氮沉降及降水变化的光合响应[J]. 草地学报, 2021, 29(1):121-130.
[29]	赵静, 刘羽霞, 许嘉巍, 等. 模拟氮沉降对长白山苔原灌草混合群落中植物光合特性的影响[J]. 植物科学学报, 2020, 38(5):678-686.
[30]	付晓萍, 田大伦, 闫文德. 模拟酸雨对樟树光合日变化的影响[J]. 中南林业科技大学学报, 2006, 26(6):38-43.
[31]	ZHANG D Y, WANG X H, CHEN Y, et al. Determinant of photosynthetic capacity in rice leaves under ambient air conditions[J]. Photosynthetica, 2005, 43(2):273-276. doi: 10.1007/s11099-005-0044-8 URL
[32]	CHANTIGNY M H. Dissolved and water-extractable organic matter in soils: a review on the influence of land use and management practices[J]. Geoderma, 2003, 113(3-4):357-380. doi: 10.1016/S0016-7061(02)00370-1 URL
[33]	翟占伟, 龚吉蕊, 罗亲普, 等. 氮添加对内蒙古温带草原羊草光合特性的影响[J]. 植物生态学报, 2017, 41(2):196-208. doi: 10.17521/cjpe.2016.0128
[34]	ZHANG T, YANG S, GUO R, et al. Warming and Nitrogen Addition Alter Photosynthetic Pigments, Sugars and Nutrients in a Temperate Meadow Ecosystem[J]. Plos one, 2016, 11(5):e0155375.
[35]	钱树玥, 王巧, 哀建国, 等. 氮沉降和磷添加对杉木光合及叶绿素荧光特征的影响[J]. 生态科学, 2018, 37(5):113-121.
[36]	MOLLIER, PELLERIN. Maize root system growth and development as influenced by phosphorus deficiency[J]. Journal of experimental botany, 1999, 50(333):487-497. doi: 10.1093/jxb/50.333.487 URL
[37]	吴楚, 范志强, 王政权. 磷胁迫对水曲柳幼苗叶绿素合成、光合作用和生物量分配格局的影响[J]. 应用生态学报, 2004, 15(6):935-940.
[38]	LI D, MO J M, FANG Y T, et al. Ecophysiological responses of woody plants to elevated nitrogen deposition[J]. Journal of tropical and subtropical botany, 2004, 12(5):482-488.
[39]	潘晓华, 刘水英, 李锋, 等. 低磷胁迫对不同水稻品种幼苗光合作用的影响[J]. 作物学报, 2003, 29(5):770-774.
[40]	张海伟, 徐芳森. 不同磷水平下甘蓝型油菜光合特性的基因型差异研究[J]. 植物营养与肥料学, 2010, 16(5):1196-1202.
[41]	王朋朋, 王丹, 王昊, 等. 长期氮、磷添加对青藏高原2种高寒草甸植物光合特性的影响[J]. 江苏农业科学, 2019, 47(13):325-329.
[42]	董莉莉, 张慧东, 魏文俊, 等. 长白落叶松人工林生理及光合特性对模拟氮沉降的响应[J]. 辽宁林业科技, 2018(6):1-4.
[43]	武传兰, 隆小华, 金善钊, 等. 盐胁迫对不同品系杨树幼苗生长和叶绿素荧光的效应[J]. 生态学杂志, 2012, 31(6):1347-1352.
[44]	李月灵, 金则新. 玉兰叶片光合速率和叶绿素荧光参数的日变化[J]. 江苏农业科学, 2012, 40(9):164-168.
[45]	SARIJEVA G, KNAPP M, LICHTENTHALER H K. Differences inphotosynthetic activity, chlorophyll and carotenoid levels, and inchlorophyll fluorescence parameters in green sun and shade leaves of Ginkgo and Fagus[J]. Journal of plant physiology, 2007, 164(7):950 -955. doi: 10.1016/j.jplph.2006.09.002 URL
[46]	李俊贞, 何乐祖, 赵春梅, 等. 盐胁迫对黄果厚壳桂幼苗荧光和生理特性的影响[J]. 山西农业科学, 2021, 49(8):919-923.
[47]	单提波, 徐正进. 不同气孔密度水稻叶片光合及叶绿素荧光参数的日变化特征[J]. 沈阳农业大学学报, 2015, 46(2):129-134.
[48]	巩擎柱, 吕金印, 徐炳成, 等. 水分胁迫和种植方式对小麦叶绿素荧光参数及水分利用效率的影响[J]. 西北农林科技大学学报, 2006, 34(5):83-92.
[49]	翁小航, 李慧, 周永斌, 等. 氮钙协同对杨树生长、光合特性及叶绿素荧光的影响[J]. 沈阳农业大学学报, 2021, 52(3):356-361.
[50]	张羽, 孙浩钊, 刘燕飞, 等. 氮添加和干旱对亚热带两个树种生长、光合及挥发性有机碳释放的影响[J]. 福建农林大学学报(自然科学版), 2021, 50(4):524-532.