大气还原态氮干湿沉降下水曲柳和兴安落叶松菌根化苗木生长响应

doi:10.17521/cjpe.2024.0474

摘要/Abstract

摘要：

研究还原态氮增加背景下东北主要造林树种水曲柳(Fraxinus mandshurica, 阔叶, 具丛枝菌根)和兴安落叶松(Larix gmelinii, 针叶, 具外生菌根)苗木对干湿还原态氮沉降的生长响应, 有助于深入了解不同类型苗木生长与大气氮沉降形态的关系, 为苗木培育精准养分管理提供理论依据。该研究以水曲柳和兴安落叶松播种苗为试验材料, 模拟不同浓度的干湿氮沉降, 即0 (CK)、35 (ND-35)、70 (ND-70)、35 (NW-35)、70 (NW-70) kg N·hm^-2·a^-1, 观测两种苗木生长、光合能力、吸收根发育及菌根侵染率的变化。主要结果: 大气干湿氮沉降下, 两种菌根类型的苗木均逐渐减少对菌根真菌的依赖性, 以提高自身光合性能或根系吸收能力作为主要响应形式。(1)具有丛枝菌根的水曲柳在氮干沉降时, 主要通过改变光合能力促进叶片及总生物量的积累, 在ND-70处理下, 净光合速率、总叶绿素含量和叶片生物量比对照分别增加49.61%、76.29%和53.84%。氮湿沉降时主要通过增加吸收根与土壤的接触面积, 在NW-35和NW-70处理下, 吸收根表面积比对照增加14.96%和16.17%, 提高了氮利用效率。(2)具有外生菌根的兴安落叶松对氮湿沉降响应更明显, 主要通过吸收根变长变细来促进根系的吸收能力, 在NW-70处理下, 吸收根长度比对照增加20.70%, 吸收根平均直径和皮层厚度比对照减少10.14%和27.25%。研究有助于深入分析菌根类型与大气还原态氮沉降之间的关系, 为不同菌根类型苗木精准养分管理提供借鉴。

关键词: 水曲柳, 兴安落叶松, 氮沉降, 苗木生长, 光合能力, 吸收根, 菌根真菌

Abstract:

Aims This study examines the growth responses of major afforestation tree species in Northeast China, specifically Fraxinus mandshurica (a broadleaf species associated with arbuscular mycorrhizal fungi) and Larix gmelinii (a coniferous species associated with ectomycorrhizal fungi), to varying levels of dry and wet deposition of reduced nitrogen. The study aims to enhance our understanding of the relationship between seedling growth patterns and atmospheric nitrogen deposition forms, thereby providing a theoretical foundation for precise nutrient management in seedling cultivation.

Methods The research utilized arbuscular mycorrhizal (AM) seedlings of F. mandshuricaand ectomycorrhizal (ECM) seedlings of L. gmelinii as test plants, and simulated varying levels of dry and wet reduced nitrogen deposition: 0 (CK), 35 (ND-35), 70 (ND-70), 35 (NW-35), and 70 (NW-70) kg N·hm^-2·a^-1. We assessed changes in seedling growth, photosynthetic capacity, root development, and mycorrhizal infection rates of the two types of seedlings.

Important findings Under conditions of atmospheric dry and wet nitrogen deposition, seedlings of both mycorrhizal types gradually reduced their mycorrhizal dependence. The primary response are the enhancement of their photosynthetic rate and an increase in root absorption capacity. (1) During nitrogen dry deposition, a significant increase in leaf and total biomass was observed for arbuscular mycorrhizal seedling of F. mandshurica, attributed to changes in photosynthetic capacity. Under the ND-70 treatment, the net photosynthetic rate, total chlorophyll content, and leaf biomass increased by 49.61%, 76.29% and 53.84% respectively, compared to the control. In contrast, during nitrogen wet deposition, nitrogen use efficiency increased primarily due to an increased contact area between absorbing roots and soil. Under NW-35 and NW-70 treatments, the surface area of absorbing roots increased by 14.96% and 16.17%, respectively, compared to the control. (2) Ectomycorrhizal seedlings of L. gmelinii exhibited a more pronounced response to nitrogen wet deposition, primarily enhanced the absorption capacity of the root system through the elongation and thinning of the absorbing roots. Under NW-70 treatment, the absorbing root length increased by 20.70% compared to the control, while the average diameter and cortical thickness of the absorbing roots decreased by 10.14% and 27.25%, respectively. The study performed an in-depth analysis of the relationship between mycorrhizal types and atmospheric reduced nitrogen deposition, serving as a reference for the precise nutrient management for tree seedlings with different mycorrhizal types.

Key words: Fraxinus mandshurica, Larix gmelinii, nitrogen deposition, seedling growth, photosynthetic capacity, absorbing root, mycorrhizal fungi

张诚航, 卫星, 吴纯泽, 王裕尧, 李浩楠. 大气还原态氮干湿沉降下水曲柳和兴安落叶松菌根化苗木生长响应. 植物生态学报, 2026, 50(3): 649-659. DOI: 10.17521/cjpe.2024.0474

ZHANG Cheng-Hang, WEI Xing, WU Chun-Ze, WANG Yu-Yao, LI Hao-Nan. Growth response of mycorrhizal Fraxinus mandshurica and Larix gmelinii seedlings to dry and wet deposition of atmospheric reduced nitrogen. Chinese Journal of Plant Ecology, 2026, 50(3): 649-659. DOI: 10.17521/cjpe.2024.0474

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2024.0474

https://www.plant-ecology.com/CN/Y2026/V50/I3/649

图/表 8

表1 大气干湿氮沉降下水曲柳和兴安落叶松苗木土壤理化性质的响应(平均值±标准误)

Table 1 Response of chemo-physical properties of Fraxinus mandshurica and Larix gmelinii soil to different dry or wet nitrogen deposition levels (mean ± SE)

树种 Tree species	处理 Treatment	铵态氮含量 NH₄⁺-N content (mg^.kg^-1)	硝态氮含量 NO₃^--N content (mg^.kg^-1)	有效氮含量 AN content (mg^.kg^-1)	土壤pH Soil pH
水曲柳 Fraxinus mandshurica	CK	14.76 ± 1.22^c	45.67 ± 1.33^c	61.10 ± 1.87^c	5.67 ± 0.09^a
	ND-35	15.15 ± 1.61^c	48.84 ± 3.55^c	63.98 ± 1.96^c	5.64 ± 0.07^a
	ND-70	17.90 ± 1.04^c	50.20 ± 1.72^c	68.10 ± 2.09^c	5.57 ± 0.06^a
	NW-35	69.16 ± 2.49^b	60.63 ± 1.61^b	119.79 ± 3.38^b	5.57 ± 0.04^a
	NW-70	136.02 ± 4.38^a	71.11 ± 2.77^a	168.41 ± 3.68^a	5.48 ± 0.03^a
兴安落叶松 Larix gmelinii	CK	13.60 ± 2.18^c	46.77 ± 1.12^c	60.37 ± 2.87^c	5.66 ± 0.08^a
	ND-35	15.14 ± 2.10^c	47.74 ± 1.28^c	62.88 ± 3.37^c	5.63 ± 0.06^a
	ND-70	15.95 ± 2.61^c	49.93 ± 1.04^c	65.89 ± 2.07^c	5.57 ± 0.04^a
	NW-35	74.53 ± 2.40^b	62.28 ± 1.24^b	136.70 ± 2.09^b	5.53 ± 0.04^a
	NW-70	107.68 ± 5.07^a	77.16 ± 1.75^a	184.84 ± 4.88^a	5.51 ± 0.06^a

图1 不同浓度干湿氮沉降下水曲柳和兴安落叶松苗木生物量分配的响应(平均值±标准误)。不同大写字母表示同种苗木的总生物量在不同处理间差异显著(p < 0.05), 不同小写字母表示同种苗木的同一器官在不同处理间差异显著(p < 0.05)。CK, 未经任何氮处理; ND-35, 35 kg N·hm-2·a-1的干沉降; ND-70, 70 kg N·hm-2·a-1的干沉降; NW-35, 35 kg N·hm-2·a-1的湿沉降; NW-70, 70 kg N·hm-2·a-1的湿沉降。

Fig. 1 Responses of biomass partitioning of Fraxinus mandshurica and Larix gmelinii under different dry or wet nitrogen deposition levels (mean ± SE). Different uppercase letters indicate significant differences in total biomass for the same seedling species across various treatments (p < 0.05), while different lowercase letters indicate significant differences within the same organ of the same seedling species across different treatments (p < 0.05). CK, without any nitrogen treatment; ND-35, dry deposition of nitrogen at rate of 35 kg N·hm-2·a-1; ND-70, dry deposition of nitrogen at rate of 70 kg N·hm-2·a-1; NW-35, wet deposition of nitrogen at rate of 35 kg N·hm-2·a-1; NW-70, wet deposition of nitrogen at rate of 70 kg N·hm-2·a-1.

表2 不同浓度干湿氮沉降下水曲柳与兴安落叶松苗木光合参数的响应(平均值±标准误)

Table 2 Response of photosynthetic parameters of Fraxinus mandshurica and Larix gmelinii to different dry or wet nitrogen deposition levels (mean ± SE)

树种 Tree species	处理 Treatment	叶面积 Leaf area (cm²)	净光合速率 P_n(μmol·m^-2·s^-1)	气孔导度 G_s(μmol·m^-2·s^-1)	蒸腾速率 T_r(μmol·m^-2·s^-1)	胞间CO₂浓度 C_i(μmol·m^-2·s^-1)
水曲柳 Fraxinus mandshurica	CK	87.05 ± 6.47^b	2.62 ± 0.26^b	0.03 ± 0.01^b	1.01 ± 0.06^c	233.20 ± 9.76^b
	ND-35	108.68 ± 5.71^ab	3.76 ± 0.22^a	0.04 ± 0.01^a	1.31 ± 0.09^ab	243.40 ± 8.82^ab
	ND-70	133.92 ± 13.01^a	3.92 ± 0.26^a	0.04 ± 0.01^a	1.52 ± 0.09^a	265.20 ± 10.24^a
	NW-35	90.29 ± 5.61^b	2.96 ± 0.12^b	0.03 ± 0.01^b	0.93 ± 0.05^c	247.40 ± 9.44^ab
	NW-70	107.02 ± 12.93^ab	3.10 ± 0.15^b	0.03 ± 0.01^b	1.11 ± 0.07^bc	265.60 ± 5.31^a
兴安落叶松 Larix gmelinii	CK	13.74 ± 1.62^a	2.07 ± 0.14^a	0.03 ± 0.01^a	0.67 ± 0.04^a	261.80 ± 15.22^b
	ND-35	14.53 ± 0.55^a	2.29 ± 0.12^a	0.03 ± 0.01^a	0.69 ± 0.03^a	257.00 ± 10.77^b
	ND-70	16.08 ± 2.30^a	2.39 ± 0.16^a	0.03 ± 0.01^a	0.72 ± 0.03^a	280.80 ± 13.26^ab
	NW-35	15.91 ± 1.40^a	2.25 ± 0.19^a	0.03 ± 0.01^a	0.71 ± 0.04^a	270.60 ± 10.62^b
	NW-70	17.07 ± 1.46^a	2.44 ± 0.13^a	0.03 ± 0.01^a	0.72 ± 0.02^a	309.40 ± 10.66^a

图2 不同浓度干湿氮沉降下水曲柳和兴安落叶松苗木叶绿素含量的响应(平均值±标准误)。不同大写字母表示同种苗木的叶绿素(a+b)含量在不同处理间差异显著(p < 0.05), 不同小写字母表示同种苗木的同一指标在不同处理间差异显著(p < 0.05)。CK, 未经任何氮处理; ND-35, 35 kg N·hm-2·a-1的干沉降; ND-70, 70 kg N·hm-2·a-1的干沉降; NW-35, 35 kg N·hm-2·a-1的湿沉降; NW-70, 70 kg N·hm-2·a-1的湿沉降。

Fig. 2 Responses of chlorophyll content of Fraxinus mandshurica and Larix gmelinii to different dry or wet nitrogen deposition levels (mean ± SE). Different uppercase letters indicate significant differences in chlorophyll (a+b) content of the same seedling under different treatments (p < 0.05), while different lowercase letters signify significant differences in the same parameter for the same seedling species across different treatments (p < 0.05). CK, without any nitrogen treatment; ND-35, dry deposition of nitrogen at rate of 35 kg N·hm-2·a-1; ND-70, dry deposition of nitrogen at rate of 70 kg N·hm-2·a-1; NW-35, wet deposition of nitrogen at rate of 35 kg N·hm-2·a-1; NW-70, wet deposition of nitrogen at rate of 70 kg N·hm-2·a-1.

表3 干湿氮沉降对水曲柳与兴安落叶松苗木吸收根形态的影响(平均值±标准误)

Table 3 Effects of dry or wet nitrogen deposition on the morphology of absorbing roots of Fraxinus mandshurica and Larix gmelinii seedlings (mean ± SE)

树种 Tree species	处理 Treatment	根尖数 Root tips	根长度 Root length (cm)	根表面积 Root surface area (cm²)	根体积 Root volume (cm³)	根平均直径 Root average diameter (μm)
水曲柳 Fraxinus mandshurica	CK	3608.33 ± 212.69^b	1394.02 ± 34.07^ab	133.27 ± 3.96^c	1.08 ± 0.02^a	985.16 ± 12.52^c
	ND-35	4068.67 ± 105.66^ab	1372.66 ± 35.26^b	137.71 ± 2.13^c	1.13 ± 0.03^a	1020.39 ± 18.39^bc
	ND-70	4200.00 ± 186.58^ab	1515.32 ± 54.77^ab	140.27 ± 2.99^bc	1.12 ± 0.02^a	1041.25 ± 13.46^b
	NW-35	4278.00 ± 291.16^ab	1480.63 ± 46.04^ab	153.21 ± 6.21^ab	1.12 ± 0.02^a	1034.18 ± 34.13^bc
	NW-70	4577.00 ± 264.68^a	1548.23 ± 68.31^a	154.83 ± 4.88^a	1.14 ± 0.02^a	1123.01 ± 12.92^a
兴安落叶松 Larix gmelinii	CK	539.33 ± 21.30^a	112.50 ± 3.99^b	11.64 ± 0.29^a	0.11 ± 0.01^b	978.77 ± 25.63^a
	ND-35	535.33 ± 18.62^a	121.67 ± 6.60^ab	11.71 ± 0.13^a	0.12 ± 0.01^ab	976.02 ± 35.47^ab
	ND-70	547.67 ± 12.14^a	125.97 ± 7.78^ab	11.68 ± 0.11^a	0.12 ± 0.01^ab	923.84 ± 19.36^ab
	NW-35	587.67 ± 34.38^a	127.48 ± 8.17^ab	12.20 ± 0.10^a	0.12 ± 0.01^ab	919.01 ± 16.56^ab
	NW-70	583.67 ± 25.56^a	135.79 ± 3.91^a	11.84 ± 0.18^a	0.13 ± 0.01^a	879.49 ± 34.22^b

表4 干湿氮沉降对水曲柳与兴安落叶松苗木吸收根解剖结构和菌根侵染率的影响(平均值±标准误)

Table 4 Effects of dry or wet nitrogen deposition on anatomic structure and mycorrhizal infection rates of absorbing root of Fraxinus mandshurica and Larix gmelinii seedlings (mean ± SE)

树种 Tree species	处理 Treatment	皮层厚度 Root cortex thickness (μm)	中柱直径 Root stele diameter (μm)	菌根侵染率 Mycorrhizal colonization rate (%)
水曲柳 Fraxinus mandshurica	CK	340.16 ± 14.63^b	645.01 ± 15.43^a	60.00 ± 5.74^ab
	ND-35	351.47 ± 22.02^ab	668.92 ± 17.74^a	58.67 ± 3.33^ab
	ND-70	364.71 ± 15.82^ab	676.54 ± 13.88^a	56.67 ± 5.78^ab
	NW-35	360.35 ± 16.17^ab	673.50 ± 18.92^a	75.00 ± 2.89^a
	NW-70	412.09 ± 15.29^a	694.24 ± 17.71^a	48.33 ± 4.41^b
兴安落叶松 Larix gmelinii	CK	285.35 ± 13.10^a	693.42 ± 16.64^a	38.33 ± 4.41^a
	ND-35	259.22 ± 11.87^ab	686.80 ± 36.23^a	36.67 ± 1.67^ab
	ND-70	239.62 ± 23.74^ab	684.22 ± 30.88^a	31.67 ± 1.67^ab
	NW-35	247.11 ± 16.98^ab	671.90 ± 30.62^a	28.34 ± 4.41^ab
	NW-70	207.59 ± 19.18^b	671.90 ± 15.60^a	26.67 ± 3.33^b

图3 干湿氮沉降下两种苗木生长及光合能力的相关性。*, p ≤ 0.05。ChlT, 总叶绿素含量; Ci, 胞间CO2浓度; Gs, 气孔导度; LA, 叶面积; LB, 叶生物量; NC, 氮浓度; Pn, 净光合速率; TB, 总生物量; Tr, 蒸腾速率。

Fig. 3 Correlation between growth and photosynthetic capacity of two seedling species under dry and wet nitrogen deposition. *, p ≤ 0.05. ChlT, total chlorophyll content; Ci, intercellular CO2 concentration; Gs, stomatal conductance; LA, leaf area; LB, leaf biomass; NC, nitrogen concentration; Pn, net photosynthetic rate; TB, total biomass; Tr, transpiration rate.

图4 干湿氮沉降下两种苗木吸收根和菌根发育的相关性。*, p ≤ 0.05。ARB, 吸收根生物量; ARD, 吸收根平均直径; ARL, 吸收根长度; ARS, 吸收根表面积; ART, 吸收根根尖数; ARV, 吸收根体积; CT, 皮层厚度; MIR, 菌根侵染率; NC, 氮浓度; SD, 中柱直径。

Fig. 4 Correlation between absorbing root and mycorrhizal development of two seedling species under dry and wet nitrogen deposition. *, p ≤ 0.05. ARB, absorbing root biomass; ARD, average absorbing root diameter; ARL, absorbing root length; ARS, absorbing root surface area; ART, number of absorbing root tips; ARV, absorbing root volume; CT, cortex thickness; MIR, mycorrhizal infection rate; NC, nitrogen concentration; SD, stele diameter.

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