施肥对闽楠幼苗光合碳固定的影响

doi:10.11707/j.1001-7488.20220505

摘要/Abstract

摘要：

目的: 研究不同氮(N)、磷(P)和钾(K)施肥量和施肥方式影响闽楠叶片光合特性、内含物含量, 酶活性及植株养分吸收量, 揭示施肥调控光合碳固定机制, 为闽楠壮苗培育和科学施肥提供理论依据。方法: 采用"3414"田间试验方法, 分析不同N、P和K施肥处理对闽楠幼苗光合色素含量、光合作用参数、叶片内含物及相应酶活性的影响; 通过极差分析, 筛选最佳的氮磷钾施肥方式; 通过逐步回归分析、相关分析和通径分析, 构建施肥调控光合碳固定途径。结果: 1) N、P、K施肥提高闽楠叶片光合色素含量, 光合、蒸腾和水分利用能力, 抑制叶片可溶性糖, 可溶性蛋白和丙二醛含量, 提高叶片硝酸还原酶、谷氨酰胺合成酶和酸性磷酸酶活性, 抑制多酚氧化酶活性, 进而提高叶片淀粉含量, 促进碳固定。在光合碳固定中, 叶片酸性磷酸酶、水分利用效率和淀粉含量是调控的关键点。2) N元素增强硝酸还原酶活性, 提高叶绿素a含量, 增强水分利用效率, 促进碳固定。3) P元素提高叶绿素b含量, 降低丙二醛含量, 保护细胞完整性, 提高碳同化效率, 但对于光合产物碳代谢无显著影响。4) K元素提高叶绿素a和b含量, 增大气孔导度, 提高气孔CO₂和水分利用效率, 提高淀粉含量, 促进光合碳固定。5) 在N、P、K配施下, N, P和K吸收量最大, 与酸性磷酸酶呈极显著相关, 最有利于光合碳固定; 其次为N、P配施, 最差的为单施磷肥, 以T6(N: 0.532 g·plant^-1, P₂O₅: 0.133 g·plant^-1, K₂O: 0.356 g·plant^-1)处理效果最好。结论: 施肥后, 闽楠幼苗吸收N、P和K, 加大叶片气孔通气, 显著提高气孔导度和蒸腾速率, 提高水分利用效率, 增强闽楠幼苗的净光合速率, 促进光合产物的输出, 提高蛋白质合成, 在酸性磷酸酶的作用下调控碳代谢, 产生淀粉, 为其生长提供物质和能量。

关键词: 光合碳固定, 氮磷钾施肥, 酸性磷酸酶, 水分利用效率, 淀粉含量, 闽楠幼苗

Abstract:

Objective: In this paper, the effects of different fertilization rates and models of nitrogen (N), phosphorus (P) and potassium (K) on photosynthetic characteristics, organic compound content and enzyme activities in leaves, and nutrients of Phoebe bournei seedlings were investigated, to reveal the mechanism of photosynthetic carbon fixation regulated by fertilization, which would provide a theoretical basis for the cultivation of strong seedlings and scientific fertilization of P. bournei. Objective: The "3414" field experiment was conducted to analyze the effects of different N, P, K fertilization treatments on photosynthetic pigment contents, photosynthetic indexes, leaf organic compound content and enzyme activities of P. bournei seedlings. The range analysis method was used to screen the best N, P, K fertilization model. The stepwise regression, correlation and path analysis method were used to construct the photosynthetic carbon fixation pathway regulated by fertilization. Result: 1) N, P, K fertilization increased the photosynthetic pigment content, photosynthesis, transpiration and water use capacity, reduced the contents of soluble sugars, soluble protein and malondialdehyde (MDA), increased the activities of nitrate reductase (NR), glutamine synthetase (GS) and acid phosphatase (ACP), inhibited polyphenol oxidase (PPO) activity, increased leaf starch content, and then promoted leaf carbon fixation of P. bournei seedlings. In the photosynthetic carbon metabolism, ACP, WUE and starch content were the key regulation points. 2) N element enhanced NR activity, increased chlorophyll a content, water use efficiency, and leaf carbon fixation of P. bournei seedlings. 3) P element increased chlorophyll b content, decreased MDA content, protected cell integrity, and improved carbon assimilation efficiency, but had no significant effect on the metabolism of photosynthate of P. bournei seedlings. 4) K element increased the contents of chlorophyll a and b, stomatal conductance, stomatal CO₂ and water use efficiency, starch content, and leaf carbon fixation of P. bournei seedlings. 5) Under the combined application of N, P and K fertilizer, the absorption of N, P and K was the most, which was significantly correlated with ACP activities, and was the most beneficial to photosynthetic carbon fixation, followed by the combined application of N and K fertilizer, and the smallest was single application of P fertilizer. The treatment effect of T6 (N: 0.532 g·plant^-1, P₂O₅: 0.133 g·plant^-1, K₂O: 0.356 g·plant^-1) was the best. Conclusion: After fertilization, P. bournei seedlings absorb N, P and K element, thereby increase the stomata ventilation, significantly improve the stomatal conductance, transpiration rate, and water use efficiency, increase net photosynthetic rate, photosynthetic product, and protein synthesis. Under the action of ACP enzyme, fertilization regulates carbon metabolism, and promotes the production of starch which can provide material and energy for growth of P. bournei seedlings.

Key words: photosynthetic carbon fixation, N, P, K fertilization, acid phosphatase, water use efficiency, starch content, Phoebe bournei seedling

中图分类号:

S723.7
S794

王妍,冯金玲,吴小慧,黄蓝明,吴娟,陈宇,杨志坚. 施肥对闽楠幼苗光合碳固定的影响[J]. 林业科学, 2022, 58(5): 40-52.

Yan Wang,Jinling Feng,Xiaohui Wu,Lanming Huang,Juan Wu,Yu Chen,Zhijian Yang. Effects of Fertilization on Photosynthetic Carbon Fixation of Phoebe bournei Seedlings[J]. Scientia Silvae Sinicae, 2022, 58(5): 40-52.

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参考文献 0

	陈隆升, 陈永忠, 彭邵锋, 等. 油茶对低磷胁迫的生理生化效应研究. 林业科学研究, 2010, 23 (5): 782- 786.
	Chen L S , Chen Y Z , Peng S F , et al. Study on physiological and biochemical responses of Camellia oleifera to low phosphorus stress. Forest Research, 2010, 23 (5): 782- 786.
	楚秀丽, 刘青华, 范辉华, 等. 不同生境、造林模式闽楠人工林生长及林分分化. 林业科学研究, 2014, 27 (4): 445- 453.
	Chu X L , Liu Q H , Fan H H , et al. Growth and structure differentiation of Phoebe bournei plantation with different sites and modes of afforestation. Forest Research, 2014, 27 (4): 445- 453.
	代庭伟, 吕雪艳, 陈放, 等. 过表达本氏烟基因NbPAP2对转基因植物生长发育的影响. 四川大学学报(自然科学版), 2020, 57 (5): 197- 203.
	Dai T W , Lü X Y , Chen F , et al. Overexpression of purple phosphatase gene NbPAP2 in tabacco promotes the growth and development of transgenic plants. Journal of Sichuan University (Natural Science Edition), 2020, 57 (5): 197- 203.
	杜俊卿, 云兴福, 马立国, 等. 氯钾离子共体诱导后黄瓜叶片内酶活性的变化. 华北农学报, 2004, 19 (2): 32- 36. doi: 10.3321/j.issn:1000-7091.2004.02.009
	Du J Q , Yun X F , Ma L G , et al. The change of enzyme activities induced with copolymer of chlorine and potassium ions in cucumber leaf. Acta Agriculturae Boreali-Sinica, 2004, 19 (2): 32- 36. doi: 10.3321/j.issn:1000-7091.2004.02.009
	郝龙飞, 王庆成, 刘婷岩, 等. 指数施肥对斑叶稠李苗木生物量分配、光合作用及根系形态的影响. 林业科学, 2014, 50 (11): 175- 181.
	Hao L F , Wang Q C , Liu T Y , et al. Effect of exponential fertilization on biomass allocation, photosynthesis and root morphology of Padus maackii seedlings. Scientia Silvae Sinicae, 2014, 50 (11): 175- 181.
	贺维, 胡庭兴, 王锐, 等. 施肥对桢楠幼苗光合生理及生长特性的影响. 西北植物学报, 2014, 34 (6): 1187- 1197.
	He W , Hu T X , Wang R , et al. Effect of fertilization on photosynthetic physiology and growth characteristics of Phoebe zhennan seedlings. Acta Botanica Boreali-Occidentalia Sinica, 2014, 34 (6): 1187- 1197.
	贾剑波, 刘文娜, 文仕知, 等. 水碳控制条件对闽楠叶片气孔特征和气体交换参数的影响. 中国水土保持科学, 2019, 17 (3): 1- 7.
	Jia J B , Liu W N , Wen S Z , et al. Effects of CO₂ concentration and soil moisture on leaf stomatal traits and gas exchange parameters of Phoebe bournei. Science of Soil and Water Conservation, 2019, 17 (3): 1- 7.
	阚学飞, 赵明辉, 隋阳辉, 等. 氮肥对超绿水稻齐穗期光合特性的影响. 作物杂志, 2010, 192 (4): 24- 29. doi: 10.3969/j.issn.1001-7283.2010.04.006
	Kan X F , Zhao M H , Sui Y H , et al. Effects of nitrogen fertilizer on photosynthetic characteristics in super-green rice at full heading stage. Crops, 2010, 192 (4): 24- 29. doi: 10.3969/j.issn.1001-7283.2010.04.006
	李彬, 洪滔, 陈欣凡, 等. 氮梯度加载下闽楠幼苗生长与叶绿素荧光变化. 森林与环境学报, 2019, 39 (3): 241- 247.
	Li B , Hong T , Chen X F , et al. Growth and chlorophyll fluorescence characteristics of Phoebe bournei seedlings under nitrogen gradient loading. Journal of Forest and Environment, 2019, 39 (3): 241- 247.
	李亚姝, 曾文芳, 胡桂馨, 等. 钾元素对苜蓿碳水化合物的分配及抗蓟马的影响. 草地学报, 2020, 28 (6): 1580- 1587.
	Li Y S , Zeng W F , Hu G X , et al. Effect of potassium application on carbohydrate distribution and alfalfa resistance to Thrips (Thysanoptera: Thripidae). Acta Agrestia Sinica, 2020, 28 (6): 1580- 1587.
	林启美, 黄德明. 应用酸性磷酸酶进行番茄磷素诊断. 华北农学报, 1991, 6 (2): 73- 83.
	Lin Q M , Huang D M . Evaluation of an acid phosphatase assay for detection of phosphorus deficiency in tomato leaves. Acta Agriculturae Boreali-Sinica, 1991, 6 (2): 73- 83.
	刘宝, 陈存及, 林达定, 等. 21个闽楠种源叶片光合色素含量及叶绿素荧光参数分析. 江西农业大学学报, 2014, 36 (1): 115- 121. doi: 10.3969/j.issn.1000-2286.2014.01.018
	Liu B , Chen C J , Lin D D , et al. Analyses of photosynthetic pigment content and chlorophyll fluorescence parameter in leaves of 21 provenances of Phoebe bournei. Acta Agrivulturae Universitatis Jiangxiensis, 2014, 36 (1): 115- 121. doi: 10.3969/j.issn.1000-2286.2014.01.018
	刘建福, 范燕萍, 王明元, 等. 发光二级管不同光质对姜黄光合特性和姜黄素含量的影响. 生态学杂志, 2014, 33 (3): 631- 636.
	Liu J F , Fan Y P , Wang M Y , et al. Effects of different LED light qualities on photosynthetic characteristics and curcumin contents of Curcuma longa L. Chinese Journal of Ecology, 2014, 33 (3): 631- 636.
	刘连涛, 李存东, 孙红春, 等. 氮素营养水平对棉花不同部位叶片衰老的生理效应. 植物营养与肥料学报, 2007, 13 (5): 910- 914. doi: 10.3321/j.issn:1008-505x.2007.05.023
	Liu L T , Li C D , Sun H C , et al. Physiological effects of nitrogen nutrition on the senescence of cotton leaves at different positions. Plant Nutrition and Fertilizer Science, 2007, 13 (5): 910- 914. doi: 10.3321/j.issn:1008-505x.2007.05.023
	刘敏, 李絮花, 刘文博, 等. 腐植酸对番茄苗期氮素代谢的影响. 水土保持学报, 2019, 33 (3): 327- 331.
	Liu M , Li X H , Liu W B , et al. Effect of humic acid on nitrogen metabolism at tomato seedling stage. Journal of Soil and Water Conservation, 2019, 33 (3): 327- 331.
	刘攀道, 黄睿, 许文茸, 等. 植物紫色酸性磷酸酶的研究进展. 热带作物学报, 2019, 4 (2): 410- 416.
	Liu P D , Huang R , Xu W R , et al. Research progress of purple acid phosphatase in plants. Chinese Journal of Tropical Crops, 2019, 4 (2): 410- 416.
	逯红栋, 巩振辉, 黄炜, 等. 9个辣椒雄性不育材料花蕾生理生化特性研究. 西北植物学报, 2006, 26 (4): 832- 835. doi: 10.3321/j.issn:1000-4025.2006.04.032
	Lu H D , Gong Z H , Huang W , et al. Physiological and biochemical characteristics of flower buds in nine male sterile lines of pepper (Capsicum annuum). Acta Botanica Boreali-Occidentalia Sinica, 2006, 26 (4): 832- 835. doi: 10.3321/j.issn:1000-4025.2006.04.032
	陆志峰, 鲁剑巍, 潘勇辉, 等. 钾素调控植物光合作用的生理机制. 植物生理学报, 2016, 52 (12): 1773- 1784.
	Lu Z F , Lu J W , Pan Y F , et al. Physiological mechanisms in potassium regulation of plant photosynthesis. Plant Physiology Journal, 2016, 52 (12): 1773- 1784.
	罗凡, 龚雪蛟, 张厅, 等. 氮磷钾对春茶光合生理及氨基酸组分的影响. 植物营养与肥料学报, 2015, 21 (1): 147- 155.
	Luo F , Gong X J , Zhang T , et al. Effects of nitrogen, phosphorus and potassium on photo-biological characteristics and amino acid components of tea plants in spring. Plant Nutrition and Fertilizer Science, 2015, 21 (1): 147- 155.
	罗凡, 张厅, 龚雪蛟, 等. 不同施肥方式对茶树新梢氮磷钾含量及光合生理的影响. 应用生态学报, 2014, 25 (12): 3499- 3506.
	Luo F , Zhang T , Gong X J , et al. Effects of different fertilization ways on the contents of N, P, K in new shoots and photo-biological characters of tea tree. Chinese Journal of Applied Ecology, 2014, 25 (12): 3499- 3506.
	罗欢, 司建华, 赵春彦, 等. 荒漠河岸林胡杨光合参数变化特征及影响因子研究. 高原气象, 2020, 39 (2): 393- 401.
	Luo H , Si J H , Zhao C Y , et al. Study on the variation characteristics of photosynthetic parameters and environmental influencing factors of Populus euphratica in desert riparian forest. Plateau Meteorology, 2020, 39 (2): 393- 401.
	罗杰, 陈洪, 申玲, 等. 不同肥料种类及其施肥水平对1年生桢楠幼苗光合生理及生长特性的影响. 应用与环境生物学报, 2017, 23 (5): 826- 836.
	Luo J , Chen H , Shen L , et al. Effect of different fertilizers and N levels on the photosynthetic physiology and growth of one-year-old Phoebe zhenan S. Lee seedlings. Chinese Journal of Applied & Environmental Biology, 2017, 23 (5): 826- 836.
	马长青, 柏素花, 戴洪义. '嘎拉'苹果多酚氧化酶基因MdPPO的克隆与表达分析. 植物生理学报, 2013, 49 (8): 803- 810.
	Ma C Q , Bai S H , Dai H Y . Cloning and expression analysis of polyphenol oxidase gene (PPO) identified from 'Gala' apple. Plant Physiology Journal, 2013, 49 (8): 803- 810.
	施福军, 粟春青, 韦艺, 等. 光氮互作对闽楠幼苗叶片光合生理特性的影响. 西北植物学报, 2020, 40 (4): 667- 675.
	Shi F J , Su C Q , Wei Y , et al. Effect of light and nitrogen interaction on photosynthetic physiological characteristics in leaves of Phoebe bournei seedlings. Acta Botanica Boreali-Occidentalia Sinica, 2020, 40 (4): 667- 675.
	苏兰茜, 白亭玉, 鱼欢, 等. 有机肥与化肥配施对菠萝蜜幼苗光合及养分吸收的影响. 中国土壤与肥料, 2020, 289 (5): 117- 123.
	Su L X , Bai T Y , Yu H , et al. Effects of combined application of organic fertilizer and chemical fertilizer on photosynthesis and nutrient uptake of jackfruit seedlings. Soil and Fertilizer Sciences in China, 2020, 289 (5): 117- 123.
	唐晓清, 肖云华, 王康才, 等. 氮素营养对苗期菘蓝叶中硝酸还原酶活性与矿质元素吸收的影响. 西北植物学报, 2013, 33 (9): 1851- 1858.
	Tang X Q , Xiao Y H , Wang K C , et al. Effect of nitrogen nutrition on nitrate reductase activity and mineral elements absorption of Isatis indigotica Fort. at seedling stage. Acta Botanica Boreali-Occidentalia Sinica, 2013, 33 (9): 1851- 1858.
	王东光, 杨锦昌, 李荣生, 等. 不同磷素供给闽楠苗木对自然低温的生理响应. 华南农业大学学报, 2016, 37 (2): 101- 106.
	Wang D G , Yang J C , Li R S , et al. Physiological response of Phoebe bournei seedlings with different phosphorus supply levels to natural low temperature. Journal of South China Agricultural University, 2016, 37 (2): 101- 106.
	王平荣, 张帆涛, 高家旭, 等. 高等植物叶绿素生物合成的研究进展. 西北植物学报, 2009, 29 (3): 629- 636.
	Wang P R , Zhang F T , Gao J X , et al. An overview of chlorophyll biosynthesis in higher plants. Acta Botanica Boreali-Occidentalia Sinica, 2009, 29 (3): 629- 636.
	汪顺义, 刘庆, 史衍玺, 等. 施钾对甘薯氮素转移分配及氮代谢酶活性的影响. 应用生态学报, 2016, 27 (11): 3569- 3576.
	Wang S Y , Liu Q , Shi Y X , et al. Effects of potassium on nitrogen translocation and distribution and nitrogen metabolism enzyme activities of sweet potato. Chinese Journal of Applied Ecology, 2016, 27 (11): 3569- 3576.
	王阳, 刘春花, 胡凯红, 等. 不同供磷水平对核桃实生幼苗叶片营养、光合及叶绿素荧光特性的影响. 中国果树, 2021, 212 (6): 13- 18.
	Wang Y , Liu C H , Hu K H , et al. Effects of different phosphorus levels on leaf nutrition, photosynthesis and chlorophyll fluorescence characteristics of walnut seedlings. China Fruits, 2021, 212 (6): 13- 18.
	王月福, 于振文, 李尚霞, 等. 小麦开花后不同器官k中硝酸还原酶和谷氨酰胺合成酶活性比较. 植物生理学通讯, 2003, 39 (3): 209- 210.
	Wang Y F , Yu Z W , Li S X , et al. Comparison of nitrate reductase and glutamine synthetase activities in different organs of wheat after flowering. Plant Physiology Communications, 2003, 39 (3): 209- 210.
	王振兴, 朱锦懋, 王健, 等. 闽楠幼树光合特性及生物量分配对光环境的响应. 生态学报, 2012, 32 (12): 3841- 3848.
	Wang Z X , Zhu J M , Wang J , et al. The response of photosynthetic characters and biomass allocation of P. bournei young trees to different light regimes. Acta Ecologica Sinica, 2012, 32 (12): 3841- 3848.
	向芬, 李维, 刘红艳, 等. 氮素水平对茶树叶片氮代谢关键酶活性及非结构性碳水化合物的影响. 生态学报, 2019, 39 (24): 9052- 9057.
	Xiang F , Li W , Liu H Y , et al. Effects of nitrogen levels on key enzyme activities and non-structural carbohydrates in nitrogen metabolism in tea leaves. Acta Ecologica Sinica, 2019, 39 (24): 9052- 9057.
	许改平, 吴兴波, 刘芳, 等. 高温胁迫下毛竹叶片色素含量与反射光谱的相关性. 林业科学, 2014, 50 (5): 41- 48.
	Xu G P , Wu X B , Liu F , et al. The correlation between the pigment content and reflectance spectrum in Phyllostachys edulis leaves subjected to high temperature. Scientia Silvae Sinicae, 2014, 50 (5): 41- 48.
	徐晓迪, 陈永滨, 罗梅秀, 等. 闽楠不同季节光合特性日进程变化研究. 江西农业大学学报, 2017, 39 (5): 913- 920.
	Xu X D , Chen Y B , Luo M X , et al. A study of the changes of photosynthetic characteristics of Phoebe bournei in different seasons. Acta Agriculturae Universitatis Jiangxiensis, 2017, 39 (5): 913- 920.
	杨佳骏, 吴永波, 张燕红. 高温与干旱胁迫对'南林895杨'扦插苗生长和超微结构的影响. 林业科学, 2020, 56 (5): 176- 183.
	Yang J J , Wu Y B , Zhang Y H . Effects of high temperature and drought stresses on growth and ultrastructure of Populus × euramericana 'Nanlin-895' cutting seedlings. Scientia Silvae Sinicae, 2020, 56 (5): 176- 183.
	杨志坚, 冯金玲, 吴小慧, 等. 氮磷钾施肥对闽楠幼苗营养元素吸收与利用的影响. 生态学杂志, 2021, 40 (4): 998- 1011.
	Yang Z J , Feng J L , Wu X H , et al. Effects N, P and K fertilization on nutrient uptakes and utilizations of Phoebe bournei seedlings. Chinese Journal of Ecology, 2021, 40 (4): 998- 1011.
	姚任科, 张国显, 杨丽娟. 不同灌溉、施钾方式对番茄光合特性和产量、品质的影响. 土壤通报, 2016, 47 (4): 909- 913.
	Yao R K , Zhang G X , Yang L J . Effect of various irrigation and potassium application methods on tomato photosynthetic characteristics, yield and quality in greenhouse. Chinese Journal of Soil Science, 2016, 47 (4): 909- 913.
	于浩, 陈展, 曹吉鑫, 等. O₃浓度升高对桢楠和闽楠幼苗光合作用, 抗氧化能力及生物量的影响. 林业科学研究, 2018, 31 (5): 33- 41.
	Yu H , Chen Z , Cao J X , et al. Effects of elevated O₃level on photosynthesis, antioxidant capacity and biomass of Phoebe zhennan and Phoebe bournei seedlings in subtropical China. Forest Research, 2018, 31 (5): 33- 41.
	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. 生态环境学报, 2014, 23 (5): 777- 784.
	Zhang Y L , Zhang W Q , Gan X H . Influence of low temperature stress on the photosynthetic fluorescence characteristics of 6 kinds of precious hardwoods seedling in winter. Ecology and Environment Sciences, 2014, 23 (5): 777- 784.
	郑春芳, 陈威, 刘伟成, 等. 低温胁迫后红树植物秋茄幼苗光合特性及蔗糖代谢的恢复机制. 生态学杂志, 2020, 39 (12): 4048- 4056.
	Zheng C F , Chen W , Liu W C , et al. Recovery mechanisms of photosynthesis and sucrose metabolism in Kandelia obovate seedlings from low temperature stress. Chinese Journal of Ecology, 2020, 39 (12): 4048- 4056.
	郑元, 唐军荣, 高柱, 等. 不同施肥处理对木棉叶片光合特性和幼苗生长的影响. 植物资源与环境学报, 2016, 25 (2): 55- 64.
	Zheng Y , Tang J R , Gao Z , et al. Effects of different fertilization treatments on photosynthetic characteristics of leaf and seedling growth of Bombax ceiba. Journal of Plant Resources and Environment, 2016, 25 (2): 55- 64.
	周佳民, 宋荣, 曹亮, 等. 不同百合(品)种生长发育特性、光合特性的比较分析及综合评价. 中国中药杂志, 2019, 44 (21): 4581- 4587.
	Zhou J M , Song R , Cao L , et al. Comparative analysis and comprehensive evaluation from growth and photosynthetic characteristics different lily species and varieties. China Journal of Chinese Materia Medica, 2019, 44 (21): 4581- 4587.
	Cheminant S , Wild M , Bouvier F , et al. DELLAs regulate chlorophyll and carotenoid biosynthesis to prevent photooxidative damage during seedling deetiolation in Arabidopsis. The Plant Cell, 2011, 23 (5): 1849- 1860.
	De Souza Kulmann M S , Arruda W S , Vitto B B , et al. Morphological and physiological parameters influence the use efficiency of nitrogen and phosphorus by Eucalyptus seedlings. New Forests, 2021, 7, 17.
	Fuerst E P , Okubara P A , Anderson J V , et al. Polyphenol oxidase as a biochemical seed defense mechanism. Frontiers in Plant Science, 2014, 5, 689.
	Gao D P , Zhang Z S , Ding C , et al. Increasing phosphate fertilizer application to improve photosynthetic capacity and yield of summer soybean in weak light environment. Acta Agronomica Sinica, 2020, 46 (2): 249.
	Geigenberger P . Regulation of starch biosynthesis in response to a fluctuating environment. Plant Physiology, 2011, 155 (4): 1566- 1577.
	Gu X , Liu Y , Li N , et al. Effects of the foliar application of potassium fertilizer on the grain protein and dough quality of wheat. Agronomy, 2021, 11 (9): 1749.
	Kuai J , Li X , Xie Y , et al. Leaf characteristics at recovery stage affect seed oil and protein content under the interactive effects of nitrogen and waterlogging in rapeseed. Agriculture, 2020, 10 (6): 207.
	Li S , Tian Y , Wu K , et al. Modulating plant growth-metabolism coordination for sustainable agriculture. Nature, 2018, 56 (772): 595- 600.
	Liu J , Sun C , Zhai F F , et al. Proteomic insights into the photosynthetic divergence between bark and leaf chloroplasts in Salix matsudana. Tree Physiology, 2021, 41 (11): 2142- 2152.
	McClain A M , Sharkey T D . Triose phosphate utilization and beyond: from photosynthesis to end product synthesis. Journal of Experimental Botany, 2019, 70 (6): 1755- 1766.
	Namvar A , Khandan T . Inoculation of rapeseed under different rates of inorganic nitrogen and sulfur fertilizer: impact on water relations, cell membrane stability, chlorophyll content and yield. Archives of Agronomy and Soil Science, 2015, 61 (8): 1137- 1149.
	Wang Y , Zhang Z , Liang Y , et al. High potassium application rate increased grain yield of shading-stressed winter wheat by improving photosynthesis and photosynthate translocation. Frontiers in Plant Science, 2020, 11, 134.
	Xiong D , Chen J , Yu T , et al. SPAD-based leaf nitrogen estimation is impacted by environmental factors and crop leaf characteristics. Scientific Reports, 2015, 5, 13389.
	Zang H , Xiao M , Wang Y , et al. Allocation of assimilated carbon in paddies depending on rice age, chase period and N fertilization: Experiment with ¹³CO₂ labelling and literature synthesis. Plant and Soil, 2019, 445, 113- 123.
	Zhang X , Li K , Xing R , et al. Metabolite profiling of wheat seedlings induced by chitosan: revelation of the enhanced carbon and nitrogen metabolism. Frontiers in Plant Science, 2017, 8, 2017.

编号No.	处理Treament	N/(g·plant^-1)	P₂O₅/(g·plant^-1)	K₂O/(g·plant^-1)
T1	N₀P₀K₀	0(-)	0(-)	0(-)
T2	N₀P₂K₂	0(-)	2(0.133)	2(0.356)
T3	N₁P₂K₂	1(0.266)	2(0.133)	2(0.356)
T4	N₂P₀K₂	2(0.532)	0(-)	2(0.356)
T5	N₂P₁K₂	2(0.532)	1(0.067)	2(0.356)
T6	N₂P₂K₂	2(0.532)	2(0.133)	2(0.356)
T7	N₂P₃K₂	2(0.532)	3(0.200)	2(0.356)
T8	N₂P₂K₀	2(0.532)	2(0.133)	0(-)
T9	N₂P₂K₁	2(0.532)	2(0.133)	1(0.178)
T10	N₂P₂K₃	2(0.532)	2(0.133)	3(0.534)
T11	N₃P₂K₂	3(0.798)	2(0.133)	2(0.356)
T12	N₁P₁K₂	1(0.266)	1(0.067)	2(0.356)
T13	N₁P₂K₁	1(0.266)	2(0.133)	1(0.178)
T14	N₂P₁K₁	2(0.532)	1(0.067)	1(0.178)

处理 Treatment	N增量 N increment/(mg·plant^-1)	P增量 P increment/(mg·plant^-1)	K增量 K increment/(mg·plant^-1)	C增量 C increment/(mg·plant^-1)
T1	5.04±2.05 h	6.67±0.56 ef	52.80±8.91 g	82.62±4.73 h
T2	13.48±4.31 g	9.33±1.39 cdef	104.07±11.24 def	78.96±24.11 h
T3	29.11±4.24 de	11.12±3.01 bcd	130.46±3.14 bcd	125.49±20.29 ef
T4	36.93±1.22 bc	11.44±0.61 bc	117.34±24.99 cde	187.38±7.06 b
T5	38.58±0.84 bc	12.44±3.57 bc	150.75±13.87 bc	145.46±2.64 de
T6	56.07±4.85 a	17.92±5.72 a	265.83±46.88 a	287.94±19.52 a
T7	32.46±4.84 cde	12.18±1.00 bc	92.47±5.48 ef	174.81±11.30 bc
T8	28.00±0.70 e	7.64±0.47 def	56.06±30.95 g	147.84±2.23 de
T9	34.11±2.47 bcd	10.28±1.13 cde	91.23±9.86 ef	157.03±10.22 cd
T10	18.34±5.19 fg	6.33±2.39 f	139.35±7.97 bc	109.57±11.81 fg
T11	21.34±3.81 f	6.02±1.24 f	72.11±16.19 fg	91.62±10.03 gh
T12	36.08±1.05 bc	14.60±0.48 ab	152.20±2.11 b	164.24±1.17 bcd
T13	39.38±3.05 b	16.29±1.11 a	147.77±7.18 bc	153.82±29.28 cd
T14	37.65±1.24 bc	9.58±1.04 cdef	105.77±7.29 def	155.68±6.32 cd

施肥方式 Fertilization mode	叶绿素a含量 Chlorophyll a content/(mg·g^-1)	叶绿素b含量 Chlorophyll b content/(mg·g^-1)	类胡萝卜含量 Carotenoids content/(mg·g^-1)	叶绿素总含量 Total chlorophyll content/(mg·g^-1)	叶绿素a/b比值 Chlorophyll a/b ratio
NPK	5.80±0.19 a	2.01±0.18 a	0.59±0.05 a	7.39±0.25 a	2.60±0.08 a
NP	2.47±0.15 d	2.01±0.19 a	0.42±0.05 b	3.19±0.17 d	2.08±0.09 b
NK	3.89±0.18 b	2.01±0.25 a	0.56±0.06 a	4.93±0.22 b	2.08±0.11 b
PK	3.50±0.21 c	1.48±0.15 b	0.45±0.05 b	4.98±0.27 b	0.35±0.06 c
N	2.47±0.12 d	2.01±0.30 a	0.38±0.07 bc	3.14±0.11 d	2.08±0.13 b
P	2.08±0.14 e	1.11±0.08 c	0.31±0.04 c	3.19±0.14 d	0.20±0.03 c
K	2.66±0.23 d	1.13±0.22 bc	0.44±0.07 b	3.79±0.28 c	0.28±0.09 c

施肥方式 Fertilization mode	C_i/ (mmol·mol^-1)	Tr/ (mmol·m^-2s^-1)	g_s/ (mol·m^-2s^-1)	P_n/ (μmol·m^-2s^-1)	WUE
NPK	0.217±0.010 a	0.80±0.03a	0.17±0.01 a	4.70±0.16 a	8.30±0.23 a
NP	0.137±0.011 c	0.48±0.02 c	0.12±0.01 b	3.46±0.08 c	6.12±0.17 b
NK	0.124±0.007 c	0.51±0.03 c	0.16±0.01 a	4.31±0.20 b	3.58±0.30 d
PK	0.172±0.010 b	0.61±0.04 b	0.05±0.01 c	3.68±0.19 c	7.99±0.27 a
N	0.124±0.008 c	0.30±0.02 d	0.12±0.01 b	2.81±0.08 d	3.49±0.18 d
P	0.137±0.011 c	0.48±0.02 c	0.04±0.02 c	2.29±0.07 e	4.52±0.12 c
K	0.091±0.007 d	0.51±0.05 c	0.04±0.01 c	3.62±0.29 c	1.86±0.39 e

施肥方式 Fertilization mode	可溶性糖含量 Soluble sugar content/ (mg·g^-1)	淀粉含量 Starch content/ (mg·g^-1)	可溶性蛋白含量 Soluble protein content/ (mg·g^-1)	MDA含量 MDA content/ (mmol·g^-1)
NPK	0.133±0.022 a	0.050±0.004 a	0.203±0.017 a	0.049±0.003 a
NP	0.112±0.022 ab	0.049±0.002 a	0.164±0.025 b	0.033±0.005 b
NK	0.112±0.020 ab	0.035±0.005 b	0.183±0.016 ab	0.020±0.003 d
PK	0.097±0.021 bc	0.049±0.006 a	0.160±0.013 b	0.030±0.002 b
N	0.112±0.016 ab	0.035±0.009 b	0.097±0.008 c	0.015±0.001 e
P	0.076±0.022 cd	0.049±0.008 a	0.098±0.006 c	0.025±0.002 c
K	0.045±0.020 d	0.020±0.002 c	0.160±0.034 b	0.018±0.001d e