叶重比及株高质量比解释亚高寒草甸禾本科对氮素添加的积极响应

张仁懿; 徐当会; 袁建立; 李文金; 艾得协措

doi:10.11829/j.issn.1001-0629.2019-0199

叶重比及株高质量比解释亚高寒草甸禾本科对氮素添加的积极响应

兰州大学生命科学学院 / 草地农业生态系统国家重点实验室，甘肃兰州 730000

基金项目: 国家自然科学基金(31600336)；兰州大学中央高校基本科研业务费专项资金(lzujbky-2017-159)

摘要: 草地生态系统的氮素添加往往增强禾本科的优势度，而该过程的作用机制却鲜有报道。本文以青藏高原亚高寒草甸为研究对象，通过分析不同功能群植物对氮素添加的响应及形态特征差异，探讨禾本科植物对氮素添加的适应机制。结果显示，对照群落代表物种的叶重比、株高质量比与植株高度显著相关(P < 0.05)，其中禾本科植物的叶重比和株高质量比分别为77.86%和34.32 cm·g^–1，显著高于杂类草的66.12%和7.20 cm·g^–1 (P < 0.05)。氮素添加15 g·m^–2时显著提高群落生物量(P < 0.001)、植株高度(P < 0.01)、盖度(P < 0.01)以及叶绿素含量(P < 0.05)，各功能群中仅禾本科和莎草科(即禾草类)生物量比例显著上升(均为P < 0.001)、仅禾本科的盖度显著上升(P < 0.01)；添加氮素与否，禾本科植物的平均植株高度均高于其他功能群植物(P < 0.001)。因此，禾本科植物较高的叶重比和株高质量比，提高了其地上资源的竞争能力，使其能够对氮素添加作出快速而有效的响应。

关键词:

English

Using leaf mass fractions and height/mass ratios to explain the positive response of Gramineae to nitrogen addition on a sub-alpine meadow

State Key Laboratory of Grassland Agro-ecosystems / School of Life Science, Lanzhou University, Lanzhou 730000, Gansu, China

Received Date: 2019-04-15
Accepted Date: 2019-06-13
Available Online: 2019-10-31
Publish Date: 2019-09-30

Abstract: Nitrogen addition tends to increase the dominance of Gramineae in grassland ecosystems. However, the underlying mechanism is rarely reported. We analyzed the morphological characteristics and responses of functional groups to nitrogen addition to explore the adaptation mechanism of Gramineae to nitrogen addition on a sub-alpine meadow on the Qinghai-Tibet plateau. Results showed that the leaf mass fraction (LMF) and height/mass ratio (HMR) were significantly correlated with the plant height for a representative species (P < 0.05). The LMF and HMR of Gramineae were 77.86% and 34.32 cm·g^–1, respectively, which is significantly higher (P < 0.05) than those of forbs (66.12% and 7.20 cm·g^–1, respectively). Nitrogen addition (15 g·m^–2) significantly increased the aboveground biomass (P < 0.001), plant height (P < 0.01), coverage (P < 0.01) and, chlorophyll content (P < 0.05) of the community, for which only the biomass proportions of only Gramineae and sedge increased significantly (both P < 0.001), and coverage of only Gramineae increased significantly (P < 0.01). Plant height of Gramineae was always higher than that of the other functional groups whether nitrogen was added or not (P < 0.001, respectively). Therefore, a high LMF and HMR of Gramineae enhances their competition potential for aboveground resources, and enables them to respond quickly and effectively to nitrogen addition.

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参考文献

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图 1 草本植物植株示意图

Figure 1. Model of herbaceous plants

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表 1 各功能群植物的地上生物量、盖度、平均高度以及植株数(n = 10)

Table 1 Aboveground biomass, coverage, average height and number from functional groups (n = 10)

功能群 Functional group	地上生物 Aboveground biomass/(g·m^–2)		盖度 Coverage/%		高度 Height/cm		株数 Plant number/(plant·m^–2)
功能群 Functional group	对照 CK	氮添加 + N	对照 CK	氮添加 + N	对照 CK	氮添加 + N	对照 CK	氮添加 + N
杂类草 Forb	99.73 ± 17.49	123.59 ± 19.08^***	54.67 ± 3.53	55.12 ± 5.23	8.10 ± 2.49	11.61 ± 1.75	501.60 ± 5.00	392.40 ± 3.48^***
禾本科 Gramineae	131.76 ± 18.52	235.44 ± 27.41^***	26.86 ± 2.15	36.10 ± 4.59^***	46.81 ± 4.63	53.83 ± 5.42	71.20 ± 9.28	87.20 ± 10.84
莎草科 Sedge	10.48 ± 1.47	32.75 ± 3.81^***	2.32 ± 1.32	1.68 ± 0.92	8.47 ± 5.58	12.41 ± 6.90	19.60 ± 11.16	12.40 ± 13.80
豆科 Legume	0.66 ± 0.48	0.83 ± 0.77	1.35 ± 0.69	0.57 ± 0.47	3.94 ± 1.32	6.50 ± 2.12	16.80 ± 2.64	2.04 ± 4.24^***
蕨类 Fern	3.81 ± 0.83	2.98 ± 0.83	2.29 ± 1.49	1.05 ± 1.26	11.33 ± 3.85	12.25 ± 4.60	5.20 ± 7.72	2.40 ± 9.20
合计 Total	246.44 ± 25.66	395.59 ± 41.76^***	87.49 ± 1.31	94.52 ± 1.53^**	13.38 ± 3.84	18.02 ± 1.66^**	614.40 ± 7.68	496.44 ± 3.32^***
，，*分别表示对照与处理间差异显著(P < 0.05，P < 0.01，P < 0.001)。　，，* indicate significant differences between control and treatment at 0.05, 0.01, and 0.001 levels, respectively.

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表 2 代表物种叶重比、株高质量比、株高叶重比和叶绿素含量

Table 2 Leaf mass fraction, height/mass ratio, height/leaf mass ratio and chlorophyll of selected species

类群 Functional group	叶重比^* Leaf mass fraction/%	株高质量比^* Height to mass ratio/(cm·g^–1)	株高叶重比^* Height to leaf mass ratio/(cm·g^–1)	叶绿素 Chlorophyll/SPAD
类群 Functional group	叶重比^* Leaf mass fraction/%	株高质量比^* Height to mass ratio/(cm·g^–1)	株高叶重比^* Height to leaf mass ratio/(cm·g^–1)	对照 CK	氮添加 + N
杂类草 Forb (n = 8)	66.12 ± 3.08	7.20 ± 1.41	10.80 ± 2.67	36.34 ± 2.33	46.83 ± 2.40
禾本科 Gramineae (n = 4)	77.86 ± 1.32	34.32 ± 2.79	39.66 ± 10.25	29.25 ± 0.38	48.03 ± 1.83
平均 Total (n = 12)	70.39 ± 2.64	17.07 ± 5.34	21.29 ± 5.78	35.09 ± 8.36	47.04 ± 8.19
代表该指标在功能群间差异显著(P < 0.05) 　 represents significan differences among functional groups at the 0.05 level.

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表 3 代表物种叶重比、株高质量比、株高叶重比与地上生物量、高度、盖度的相关系数

Table 3 Correlation coefficients between leaf mass fraction, height/mass ratio, height/leaf mass ratio and aboveground biomass, plant height and coverage for selected species

类别 Item	指标 Parameter	地上生物量 Aboveground biomass		高度 Height		盖度 Coverage
类别 Item	指标 Parameter	对照 CK	氮添加 + N	对照 CK	氮添加 + N	对照 CK	氮添加 + N
杂类草 Forb	叶重比 Leaf mass fraction	0.30	0.55	0.17	–0.06	0.32	0.65
	株高质量比 Height to mass ratio	0.00	–0.10	–0.31	–0.13	0.27	0.20
	株高叶重比 Height to leaf mass ratio	0.01	–0.10	–0.28	–0.09	0.25	0.08
禾本科 Gramineae	叶重比 Leaf mass fraction	–0.20	–0.26	–0.28	–0.33	–0.05	–0.11
	株高质量比 Height to mass ratio	–0.44	–0.42	–0.22	–0.28	–0.35	–0.39
	株高叶重比 Height to leaf mass ratio	–0.45	–0.42	–0.21	–0.27	–0.37	–0.41
所有代表种 Total	叶比例 Leaf mass fraction	0.42	0.73^*	0.63^*	0.59^*	0.41	0.72^*
	比重高 Height to mass ratio	0.01	0.20	0.69^*	0.64^*	0.03	0.23
	比叶重高 Height to leaf mass ratio	0.12	0.18	0.67^*	0.63^*	0.10	0.18
代表相关分析显著相关(P < 0.05)。　 represents significant correlation (P < 0.05).

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参考文献(31)

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[10]	BAI Y, WU J, CLARK C, NAEEM S, PAN Q, HUANG J H, ZHANG L, HAN X. Tradeoffs and thresholds in the effects of nitrogen addition on biodiversity and ecosystem functioning: Evidence from Inner Mongolia grasslands. Global Change Biology, 2010, 16(1): 358-372. doi: 10.1111/j.1365-2486.2009.01950.x
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[13]	DICKSON T L, MITTELBACH G G, REYNOLDS H L, GROSS K L. Height and clonality traits determine plant community responses to fertilization. Ecology, 2014, 95(9): 2443-2452. doi: 10.1890/13-1875.1
[14]	辛小娟, 王刚, 杨莹博, 任正炜. 氮、磷添加对亚高山草甸地上/地下生物量分配的影响. 生态科学, 2014, 33(3): 452-458. XIN X J, WANG G, YANG Y B, REN Z W. Effects of N, P addition on above/below ground biomass allocation in a subalpine meadow. Ecological Science, 2014, 33(3): 452-458.
[15]	SPEHN E M, JOSHI J, SCHMID B, DIEMER M, KÖRNER C. Above-ground resource use increases with plant species richness in experimental grassland ecosystems. Functional Ecology, 2000, 14(3): 326-337. doi: 10.1046/j.1365-2435.2000.00437.x
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叶重比及株高质量比解释亚高寒草甸禾本科对氮素添加的积极响应

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Using leaf mass fractions and height/mass ratios to explain the positive response of Gramineae to nitrogen addition on a sub-alpine meadow

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