连续2年氮添加对中金杨幼苗叶光合特性与碳氮分配的影响

doi:10.11707/j.1001-7488.20220603

摘要/Abstract

摘要：

目的: 研究不同强度氮添加对杨树叶片光合特性、各器官生物量、非结构性碳水化合物积累和分配的影响, 探讨过量氮添加下杨树的生理生态过程响应机制, 为未来氮沉降条件下杨树人工林精准施肥提供理论依据。方法: 以盆栽黑青杂交杨中'金杨7号’扦插苗为材料, 以NH₄NO₃为氮源, 设置CK(0 g·plant^-1 a^-1)、N3(3 g·plant^-1 a^-1)、N6(6 g·plant^-1 a^-1)、N9(9 g·plant^-1 a^-1)4个氮添加梯度, 连续2年测定氮添加后其功能叶光合特性、器官间生物量、碳同化物和氮分配的变化, 并分析叶片光合特性和同化物分配的调控过程。结果: 1) 连续2年氮添加后, 中金杨幼苗叶氮和叶绿素含量增加, 气孔变小变密, 瞬时水分利用效率显著提高, 光合氮利用效率显著降低。2) 中低剂量氮添加下顶梢叶片光合作用显著受到库限制, 侧枝叶净光合速率明显高于对照; 高氮处理下叶净光合速率和淀粉含量显著减小, 荧光量子产量和电子传递速率显著增加, 表明叶片中光呼吸等过程为硝酸还原过程提供大量能量。3) 氮添加持续促进侧枝生长, 顶端优势弱化, 但高氮处理下因叶片光合作用的源限制, 主茎淀粉含量和生物量积累显著降低, 根干质量和生物量分配比例与中低氮处理无显著差异; 高氮处理下各器官生长和碳氮分配结果表明, 生长素合成和运输减弱可能在碳氮关系协调过程中起到重要作用。结论: 中低剂量氮添加下中金杨侧枝叶片光合能力显著增强并促进植株生长, 叶片光合作用处于库限制; 高氮下叶片光合作用可能受到氮同化过程抑制, 顶端优势减弱, 植株生物量积累处于源限制。

关键词: 中金杨, 氮添加, 光合特性, 非结构性碳, 水化合物, 碳氮分配

Abstract:

Objective: Excessive nitrogen addition can restrain the leaf photosynthesis and growth of woody plants. However, the underlying ecophysiological mechanism is still not very clear, which is important for precision fertilization to seedlings and plantation. Poplar (Populus spp.) is an important afforestation species and biological model tree. This study aims to investigate the photosynthetic and chlorophyll fluorescence characteristics, nonstructural carbohydrate (NSC) and nitrogen distribution of hybrid poplar seedlings under nitrogen addition for 2 consecutive years, in order to reveal the ecophysiological mechanism of trees in responding and adapting to nitrogen addition. Method: In this study, the potted cuttings of P. × euramericana 'Zhongjin7' were used as experimental materials, NH₄NO₃ was used as nitrogen source and 4 gradients (CK: 0 g·plant^-1a^-1, N3: 3 g·plant^-1a^-1, N6: 6 g·plant^-1a^-1, N9: 9 g·plant^-1a^-1) were set. After nitrogen addition for 2 years, changes of leaf photosynthetic characteristics, organ biomass accumulation, non-structural carbohydrate storage and distribution were investigated. Result: After nitrogen addition for consecutive 2 years, the nitrogen content and chlorophyll content in leaves and roots significantly increased, stomata became smaller but higher density, transient water use efficiency remarkably increased, but photosynthetic nitrogen use efficiency continuously decreased. Under moderate and low dose nitrogen addition, the net photosynthesis (P_n) of leaves on top twig was suppressed by sink and high carbohydrate feedback inhibition. But the P_n of lateral branch leaves was higher than that of the control. Under high nitrogen addition, P_n and starch content were significantly decreased, but with actual fluorescent quantum yield and electron transport rate increased significantly, suggesting that photorespiration and other processes supplied large amount energy for nitrate reduction and assimilation. Nitrogen addition continued to promote lateral branch growth and weaken the apical dominance. Under N9 conditions, the starch content in main stem decreased with a lower biomass, which maybe due to the source inhibition of P_n. The root drymass and biomass distribution percentage of N9 were no significantly different with those of N3 and N6. The results of organ growth, and carbon and nitrogen distribution implied that the decline of aux in synthesis and transport might play an important role in whole plant carbon and nitrogen relationships under excessive nitrogen addition. Conclusion: Nitrogen addition with moderate and low doses can significantly enhance carbon assimilation of leaves on the lateral branch and increase plant growth of whole poplar seedlings, with a sink limit on leaf net photosynthesis. But leaf photosynthesis may be inhibited by nitrogen assimilation under high dose nitrogen addition, with a declined apical dominance. And whole plant biomass accumulation could be restricted by source.

Key words: Populus×euramericana'Zhongjin7', nitrogen addition, photosynthetic traits, nonstructural carbohydrate, carbon and nitrogen distribution,

中图分类号:

S718.43

尚佳州,高钿惠,王卫锋,周新军,宗毓铮. 连续2年氮添加对中金杨幼苗叶光合特性与碳氮分配的影响[J]. 林业科学, 2022, 58(6): 23-32.

Jiazhou Shang,Tianhui Gao,Weifeng Wang,Xinjun Zhou,Yuzheng Zong. Effect of Nitrogen Addition for Two Consecutive Years on Photosynthetic Characteristics, Carbon and Nitrogen Distribution of Populus × euramericana 'Zhongjin7' Seedlings[J]. Scientia Silvae Sinicae, 2022, 58(6): 23-32.

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参数Parameters	CK	N3	N6	N9
叶绿素a含量 Chlorophyll a content/(mg·g^-1 FW)	1.34±0.20b	1.59±0.04b	1.90±0.15a	2.05±0.13a
叶绿素b含量 Chlorophyll b content/(mg·g^-1 FW)	0.48±0.07c	0.57±0.01b	0.70±0.07a	0.75±0.02a
总叶绿素含量 Total chlorophyll content/(mg·g^-1 FW)	1.81±0.26b	2.16±0.05b	2.60±0.22a	2.80±0.14a

参数 Parameters	CK	N3	N6	N9
最大光化学量子产量 Maximal quantum yield of PSⅡ(F_v/F_m)	0.80±0.02ab	0.77±0.02b	0.80±0.00a	0.81±0.01a
实际光化学量子产量 Actual quantum yield of PSⅡ(Ф_PSⅡ)	0.51±0.02a	0.45±0.02b	0.43±0.03b	0.53±0.04a
非光化学猝灭系数 Non-photochemical quenching(NPQ)	1.02±0.12b	1.29±0.27ab	1.71±0.26a	0.91±0.23b
光化学猝灭系数 Photochemical quenching(q_P)	0.76±0.01a	0.74±0.02a	0.71±0.04a	0.74±0.02a
电子传递速率 Electron transport rate of PSⅡ(ETR)	59.05±3.14a	51.85±2.46b	50.00±4.01b	61.22±5.26a

参数 Parameters	CK	N3	N6	N9
单叶面积 Leaf area/cm²	120.42±3.23c	150.60±6.61a	135.68±10.59b	124.98±4.71c
单叶净光合速率 Net photosynthesis per leaf/(μmol·s^-1)	0.167±0.021b	0.201±0.018a	0.169±0.017b	0.147±0.016c
单叶干质量 Leaf drymass/g	0.95±0.08b	1.20±0.13a	1.10±0.04a	0.89±0.06b
比叶质量 Leaf mass per area(LMA)/(g·m^-2)	79.11±5.30a	79.85±7.67a	81.45±3.61a	71.43±2.62b
叶片厚度 Leaf thickness/μm	252.56±16.23a	226.37±9.40b	219.64±6.60c	218.86±7.67c
叶氮含量 Leaf nitrogen content/(mg·g^-1)	14.91±0.95c	15.90±0.62bc	17.18±0.89b	21.48±0.78a
单位叶面积氮含量 Leaf nitrogen per unit area/(g·m^-2)	1.14±0.01d	1.27±0.05c	1.40±0.07b	1.53±0.06a
光合氮利用效率Photosynthetic nitrogen use efficiency(PNUE)/(μmol·g^-1s^-1)	11.89±0.61a	10.36±0.46b	8.68±0.62c	7.87±0.51d
上表皮气孔密度 Upper epidermis stomatal density/mm^-2	70.35±3.08b	63.04±10.16b	81.35±17.42a	88.12±2.74a
下表皮气孔密度 Lower epidermis stomatal density/mm^-2	163.95±10.60b	170.92±5.40b	188.28±14.80a	189.65±7.16a
上表皮保卫细胞长度 Upper epidermis stomatal length/μm	25.54±0.59a	26.33±0.74a	23.92±1.10b	22.96±0.68c
下表皮保卫细胞长度 Lower epidermis stomatal length/μm	24.11±0.61a	23.92±0.49a	23.16±0.70b	21.85±0.58c

参数 Parameters	CK	N3	N6	N9
第1年株高 Plant height in the first year/cm	104.60±5.91c	128.70±2.44b	134.83±2.84ab	138.90±5.10a
第2年株高增量 Plant height increment in the second year/cm	84.60±9.10c	96.50±10.00ab	100.00±5.29a	86.30±4.35bc
第1年地径 Grand diameter in the first year/cm	0.83±0.05c	1.12±0.05ab	1.09±0.04b	1.18±0.08a
第2年地径增量 Grand diameter increment in the second year/cm	0.54±0.04a	0.55±0.05a	0.58±0.07a	0.52±0.08a
顶梢叶片数 Leaf number in tops	19.50±2.65a	23.00±1.00a	19.00±3.00a	20.00±2.65a
侧枝数 Lateral branch number	14.50±1.29c	16.67±0.58b	17.67±0.58b	20.33±1.53a
侧枝总叶片数 Total leaf number in lateral branch	79.50±14.62b	113.67±16.29ab	118.33±27.54a	121.33±22.19a
整株总叶片数 Total leaf number	99.00±16.15b	136.67±15.37a	137.33±29.02a	141.33±24.83a
根干质量 Root dry mass/g	113.28±2.52b	139.88±8.87a	138.28±19.77a	136.05±10.36a
主茎干质量 Main stem dry mass/g	84.13±4.80c	131.06±4.05a	131.94±5.48a	116.72±4.50b
侧枝干质量 Lateral branch dry mass/g	7.74±0.32b	15.03±2.41a	16.43±5.73a	15.88±2.76a
根茎总干质量 Whole-plant dry mass/g	205.15±3.24c	285.98±12.52a	286.65±11.96a	267.97±11.74b

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