青藏高原高寒草地常见植物叶属性对氮、磷添加的响应

doi:10.13209/j.0479-8023.2017.057

摘要/Abstract

摘要：

基于海北站氮磷添加实验平台, 研究青藏高原高寒草地常见植物叶片干物质含量(LDMC)、比叶面积(SLA)和叶片碳、氮、磷含量(LC, LN, LP)属性对N添加、P添加和NP同时添加的响应。从10个物种总体来看, N添加使LN显著增加9.4%。P添加使LP显著增加81.8%。N添加和P添加对LDMC和SLA有显著的交互作用。在无N添加条件下, P添加使LDMC减小2.3%, SLA增加3.5%; 在N添加条件下, P添加使LDMC减小10.1%, SLA增加15.3%。N添加和P添加对基于叶经济谱的物种排序无显著影响, 而NP同时添加显著改变了基于叶经济谱的物种排序。结果表明, 通过提高SLA和叶片NP含量, 使高寒草地植物对光、土壤可利用性N和P的获取与利用能力增强。NP同时添加可以显著改变叶属性的种间变异, 进而导致青藏高原高寒草地常见植物在叶经济谱上形成新的分布格局。因此, 在利用叶属性预测群落结构和生态系统功能对NP同时添加的响应时, 应该考虑物种的特异性响应。

关键词: 叶经济谱, N添加, P添加, 属性可塑性, 资源竞争策略

Abstract:

To investigate species’ resource competition strategies with traits of the leaf economics spectrum across contrasting environments and to examine the effects of nutrient additions on the ranking of species based on their leaf economics spectrum in an alpine grassland on the Qinghai-Tibetan Plateau, five leaf traits (LDMC: leaf dry matter content, SLA: specific leaf area, LC: leaf carbon concentration, LN: leaf nitrogen concentration and LP: leaf phosphorus concentration) were measured for 10 plant common species in all plots. The results showed that N addition significantly increased LN by 9.4% and P addition significantly increased LP by 81.8%. There were significant interactions between N addition and P addition on SLA and LDMC, which increased SLA by 15.3% and decreased LDMC by 10.1%. In addition, there were species-specific responses of leaf traits across multivariate trait space to nutrient additions. The variation in species responses to NP addition significantly changed the species ranking based on the leaf economics spectrum. These results showed that co-occurring species followed a conservative strategy in the infertile environment and an exploitative strategy in the fertilized ones by increasing SLA and leaf nitrogen and phosphorus concentrations. Different species responses to NP addition caused a new species distribution based on the leaf economics spectrum. These results suggest that, before using leaf traits to predict responses of community structure and ecosystem functioning to nitrogen and phosphorus additions, it is necessary to take the species-specific responses into consideration.

Key words: leaf economics spectrum, nitrogen addition, phosphorus addition, trait plasticity, resource competition

李颖, 林笠, 朱文琰, 张振华, 贺金生. 青藏高原高寒草地常见植物叶属性对氮、磷添加的响应[J]. 北京大学学报自然科学版, 2017, 53(3): 535-544.

Ying LI, Li LIN, Wenyan ZHU, Zhenhua ZHANG, Jinsheng HE. Responses of Leaf Traits to Nitrogen and Phosphorus Additions across Common Species in an Alpine Grassland on the Qinghai-Tibetan Plateau[J]. Acta Scientiarum Naturalium Universitatis Pekinensis, 2017, 53(3): 535-544.

导出引用管理器 EndNote|Ris|BibTeX

链接本文: https://xbna.pku.edu.cn/CN/10.13209/j.0479-8023.2017.057

https://xbna.pku.edu.cn/CN/Y2017/V53/I3/535

图/表 8

表1 海北高寒草地10种常见植物基本特征

Table 1 Common species and their functional groups in the alpine meadow of Haibei, Qinghai, China

物种	科	功能群
异针茅Stipa aliena	禾本科	多年生禾草类
垂穗披碱草Elymus nutans	禾本科	多年生禾草类
矮嵩草Kobresia humilis	莎草科	多年生禾草类
麻花艽Gentiana straminea	龙胆科	多年生杂类草
美丽风毛菊Saussurea pulchra	菊科	多年生杂类草
高山豆Tibetia himalaica	豆科	多年生杂类草
青海苜蓿Medicago archiducis-nicolai	豆科	多年生杂类草
圆萼刺参Morina chinensis	川续断科	多年生杂类草
钉柱委陵菜Potentilla saundersiana	蔷薇科	多年生杂类草
重冠紫菀Aster diplostephioides	菊科	多年生杂类草

表2 以N添加、P添加和物种为固定因子对叶干物质含量、比叶面积和叶片C, N, P含量的三因素方差分析

Table 2 Three-way ANOVA test of effects of N addition, P addition, species and their interactions on leaf dry matter content, specific leaf area, leaf C, N and P concentrations

变异来源	df	LDMC	SLA	LC	LN	LP
N添加 (N)	1	28.24^***	12.80^**	12.00^**	37.30^***	3.61
P添加 (P)	1	60.61^***	31.70^***	33.00^***	3.49	1081.42^***
Species (S)	9	656.03^***	144.82^***	58.00^***	174.55^***	50.37^***
N×P	1	18.85^**	11.08^**	3.00	4.13	0.04
N×S	9	1.25	1.33	4.00^***	12.46^***	0.87
P×S	9	0.98	1.34	1.00	2.17^*	16.66^***
N×P×S	9	0.63	0.66	1.00	2.13^*	1.31

图1 海北高寒草地常见种的LDMC, SLA, LC, LN和LP对N添加、P添加和NP添加的响应(平均值±标准误差)

Fig. 1 Responses of LDMC, SLA, LC, LN and LP to nitrogen and phosphorus additions for common species (mean±SE)

表3 海北高寒草地常见植物的LDMC, SLA, LC, LN和LP对N添加、P添加和NP添加的响应(平均值±标准误差)

Table 3 Responses of LDMC, SLA, LC, LN and LP to nutrient additions for common species (mean±SE)

处理	LDMC/(g · kg^-1)	SLA/ (cm²· g^-1)	LC/(g · kg^-1)	LN/(g · kg^-1)	LP/(g · kg^-1)
对照	254.66±13.94	168.94±9.30	441.77±1.78	24.53±1.18	1.65±0.05
N添加	250.21±13.54	171.29±8.52	443.03±1.46	26.83±1.09	1.60±0.05
P添加	248.83±13.99	174.41±8.44	436.08±1.65	24.48±1.28	3.08±0.13
NP添加	224.83±11.84	197.46±9.26	440.35±1.71	28.98±1.10	2.88±0.10

图2 不同N、P添加条件下海北高寒草地常见种在多元叶属性空间内的排序（Sa: 异针茅; En: 垂穗披碱草; Kh: 矮嵩草; Ma: 青海苜蓿; Th: 高山豆; Gs: 麻花艽; Sp: 美丽风毛菊; Mc: 圆萼刺参; Ad: 重冠紫菀; Ps: 钉柱委陵菜）

Fig. 2 Common species ranking based on traits of the leaf economic spectrum under different nitrogen and phosphorus conditions in the alpine grassland in Haibei, Qinghai, China

表4 海北高寒草地禾草类植物和杂类草植物的LDMC, SLA, LC, LN和LP对N添加、P添加和NP添加的响应(平均值±标准误差)

Table 4 Responses of LDMC, SLA, LC, LN and LP to nutrient additions for graminoids and forbs (mean ±SE)

功能群	处理	LDMC/(g·kg^-1)	SLA/ (cm²·g^-1)	LC/(g·kg^-1)	LN/(g·kg^-1)	LP/(g·kg^-1)
禾草类	对照	360.12±16.95	127.73±10.92	446.90±1.24^a	20.03±0.62^b	1.35±0.08^b
	N添加	359.18±14.86	133.81±11.47	448.51±1.21^a	22.54±0.61^ab	1.29±0.06^b
	P添加	358.96±17.93	137.50±12.10	440.48±1.58^b	19.83±0.84^b	2.74±0.17^a
	NP添加	318.01±11.50	158.31±16.29	447.10±1.60^a	25.42±1.04^a	2.54±0.12^a
杂类草	对照	207.97±8.42	187.97±10.58	439.57±2.35	26.46±1.50	1.78±0.04^b
	N添加	203.51±7.81	187.35±9.55	440.69±1.83	28.67±1.39	1.73±0.05^b
	P添加	201.63±7.92	190.23±9.33	434.19±2.14	26.48±1.63	3.11±0.14^a
	NP添加	184.90±7.88	214.24±9.57	437.45±2.10	30.50±1.40	3.02±0.12^a

表5 海北高寒草地10种常见植物在4种养分添加处理下基于主成分分析主轴1的排序

Table 5 Rankings of ten common species based on the PCA axis 1 in the four nutrient additions treatments in the alpine meadow of Haibei, Qinghai

处理	物种排序
处理	1	2	3	4	5	6	7	8	9	10
对照	Sa	Kh	En	Gs	Mc	Ps	Sp	Ad	Ma	Th
N添加	Sa	Kh	En	Gs	Mc	Ps	Sp	Ad	Ma	Th
P添加	Sa	Mc	Gs	Kh	En	Ps	Sp	Ad	Ma	Th
NP添加	Sa	Kh	Gs	Mc	En	Ps	Sp	Ma	Ad	Th

图3 海北高寒草地常见种在多元叶属性空间内对 N 添加、P添加和NP添加的响应（灰色矢量代表 N, P 添加后各物种在属性空间中的位置变化, 黑色矢量代表N, P添加后物种的平均位置变化。物种缩写同图2）

Fig. 3 Shifts of species leaf traits across multivariate trait space in response to N addition, P addition and NP addition in the alpine grassland in Haibei, Qinghai

参考文献 53

[1]	Violle C, Navas M L, Vile D, et al.Let the concept of trait be functional!. Oikos, 2007, 116: 882-892
[2]	Arnold S J.Morphology, performance and fitness. American Zoologist, 1983, 23: 347-361
[3]	孟婷婷, 倪健, 王国宏. 植物功能性状与环境和生态系统功能. 植物生态学报, 2010, 31(1): 150-165
[4]	张林, 罗天祥. 植物叶寿命及其相关叶性状的生态学研究进展. 植物生态学报, 2004, 28(6): 844-852
[5]	Wright I J, Reich P B, Westoby M, et al.The worldwide leaf economics spectrum. Nature, 2004, 428: 821-827
[6]	He J S, Wang Z, Wang X, et al.A test of the generality of leaf trait relationships on the Tibetan Plateau. New Phytologist, 2006, 170(4): 835-848
[7]	Grime J P, Thompson K, Hunt R et al. Integrated screening validates primary axes of specialisation in plants. Oikos, 1997, 79(2): 259-281
[8]	Reich P B, Ellsworth D S, Walters M B, et al.Generality of leaf trait relationships: a test across six biomes. Ecology, 1999, 80(6): 1955-1969
[9]	Díaz S, Hodgson J G, Thompson K, et al.The plant traits that drive ecosystems: evidence from three continents. Journal of Vegetation Science, 2004, 15(3): 295-304
[10]	Reich P B.The world-wide ‘fast-slow’ plant econo-mics spectrum: a traits manifesto. Journal of Ecology, 2014, 102(2): 275-301
[11]	He J S, Wang X, Flynn D F, et al.Taxonomic, phylogenetic, and environmental trade-offs between leaf productivity and persistence. Ecology, 2009, 90(10): 2779-2791
[12]	Cunningham S A, Summerhayes B, Westoby M.Evolutionary divergences in leaf structure and che-mistry, comparing rainfall and soilnutrient gradients. Ecological Monographs, 1999, 69: 569-588
[13]	Dormann C F, Woodin S J.Climate change in the Arctic: using plant functional types in a meta-analysis of field experiments. Functional Ecology, 2002, 16(1): 4-17
[14]	万宏伟, 杨阳, 白世勤, 等. 羊草草原群落 6 种植物叶片功能特性对氮素添加的响应. 植物生态学报, 2008, 32(3): 611-621
[15]	Lienin P, Kleyer M.Plant leaf economics and reproductive investment are responsive to gradients of land use intensity. Agriculture, Ecosystems & Envi-ronment, 2011, 145(1): 67-76
[16]	Garnier E, Laurent G, Bellmann A, et al.Consistency of species ranking based on functional leaf traits. New Phytologist, 2001, 152(1): 69-83
[17]	Al Haj Khaled R, Duru M, Theau J P, et al. Variation in leaf traits through seasons and N-availability levels and its consequences for ranking grassland species. Journal of Vegetation Science, 2005, 16(4): 391-398
[18]	Ordoñez J C, van Bodegom P M, Witte J P M, et al. A global study of relationships between leaf traits, climate and soil measures of nutrient fertility. Global Ecology and Biogeography, 2009, 18: 137-149
[19]	Heberling J M, Fridley J D.Biogeographic constraints on the world-wide leaf economics spectrum. Global Ecology and Biogeography, 2012, 21(12): 1137-1146
[20]	Wright J P, Sutton-Grier A.Does the leaf economic spectrum hold within local species pools across varying environmental conditions?. Functional Ecology, 2012, 26: 1390-1398
[21]	Maire V, Gross N, Hill D, et al.Disentangling coordination among functional traits using an indivi-dual-centred model: impact on plant performance at intra-and inter-specific levels. PLoS ONE, 2013, 8: e77372
[22]	Walters M B, Gerlach J P.Intraspecific growth and functional leaf trait responses to natural soil resource gradients for conifer species with contrasting leaf habit. Tree Physiology, 2013, 33: 297-310
[23]	周兴民. 中国嵩草草甸. 北京: 科学出版社, 2001
[24]	杨晓霞, 任飞, 周华坤, 等. 青藏高原高寒草甸植物群落生物量对氮、磷添加的响应. 植物生态学报, 2014, 38(2): 159-166
[25]	马建静, 吉成均, 韩梅, 等. 青藏高原高寒草地和内蒙古高原温带草地主要双子叶植物叶片解剖特征的比较研究. 中国科学: 生命科学, 2012, 42: 158-172
[26]	Geng Y, Wang L, Jin D, et al.Alpine climate alters the relationships between leaf and root morphological traits but not chemical traits. Oecologia, 2014, 175(2): 445-455
[27]	Hong J, Wang X, Wu J.Stoichiometry of root and leaf nitrogen and phosphorus in a dry alpine steppe on the northern Tibetan Plateau. PLoS ONE, 2014: e109052
[28]	李英年, 赵新全, 曹广民, 等. 海北高寒草甸生态系统定位站气候、植被生产力背景的分析. 高原气象, 2004, 23(4): 558-567
[29]	赵新全, 周兴民. 青藏高原高寒草甸生态系统管理的生态学基础: 海北高寒草甸生态系统研究站. 人类环境杂志, 1999, 28(8): 642-647
[30]	Kuo S.Methods of soil analysis, Part 3, Chemical methods. Madison: Soil Science Society of America, 1996: 869-919
[31]	Adamidis G C, Kazakou E, Fyllas N M, et al.Species adaptive strategies and leaf economic relationships across serpentine and non-serpentine habitats on Lesbos, eastern Mediterranean. PLoS ONE, 2014, 9(5): e96034
[32]	Padgett P E, Allen E B.Differential responses to nitrogen fertilization in native shrubs and exotic annuals common to Mediterranean coastal sage scrub of California. Plant Ecology, 1999, 144(1): 93-101
[33]	Lawrence D.Nitrogen and phosphorus enhance growth and luxury consumption of four secondary forest tree species in Borneo. Journal of Tropical Ecology, 2001, 17(6): 859-869
[34]	Xia J, Wan S.Global response patterns of terrestrial plant species to nitrogen addition. New Phytologist, 2008, 179(2): 428-439
[35]	胡文祥, 李伟, 杜国祯. 基于物种性状的两种高寒草甸优势禾草对施肥的响应. 兰州大学学报(自然科学版), 2011, 47(6): 68-74
[36]	宾振钧, 王静静, 张文鹏, 等. 氮肥添加对青藏高原高寒草甸6个群落优势种生态化学计量学特征的影响. 植物生态学报, 2014, 38(3): 231-237
[37]	宾振钧, 张仁懿, 张文鹏, 等. 氮磷硅添加对青藏高原高寒草甸垂穗披碱草叶片碳氮磷的影响. 生态学报, 2015, 35(14): 1-10
[38]	赵新风, 徐海量, 张鹏, 等. 养分与水分添加对荒漠草地植物钠猪毛菜功能性状的影响. 植物生态学报, 2014, 38(2): 134-146
[39]	Garnier E, Vancaeyzeele S.Carbon and nitrogen content of congeneric annual and perennial grass species: relationships with growth. Plant, Cell & Environment, 1994, 17(4): 399-407
[40]	Wright I J, Cannon K.Relationships between leaf lifespan and structural defences in a low-nutrient, sclerophyll flora. Functional Ecology, 2001, 15(3): 351-359
[41]	Wilson P J, Thompson K E N, Hodgson J G. Specific leaf area and leaf dry matter content as alternative predictors of plant strategies. New Phytologist, 1999, 143(1): 155-162
[42]	Li Y, Johnson D A, Su Y, et al.Specific leaf area and leaf dry matter content of plants growing in sand dunes. Botanical Bulletin of Academia Sinica, 2005, 46: 127-134
[43]	Aerts R, Chapin III F S. The mineral nutrition of wild plants revisited: a re-evaluation of processes and patterns. Advances in Ecological Research, 2000, 30: 1-67
[44]	Chapin III F S. The mineral nutrition of wild plants. Annual Review of Ecology and Systematics, 1980, 11: 233-260
[45]	Van Wijk M T, Williams M, Gough L, et al. Luxury consumption of soil nutrients: a possible competitive strategy in above-ground and below-ground biomass allocation and root morphology for slow-growing arctic vegetation?. Journal of Ecology, 2003, 91(4): 664-676
[46]	Funk J L, Jones C G, Lerdau M T.Leaf-and shoot-level plasticity in response to different nutrient and water availabilities. Tree Physiology, 2007, 27(12): 1731-1739
[47]	Pontes D S L, Louault F, Carrère P, et al. The role of plant traits and their plasticity in the response of pasture grasses to nutrients and cutting frequency. Annals of Botany, 2010: mcq066
[48]	Fonseca C R, Overton J M, Collins B, et al.Shifts in trait-combinations along rainfall and phosphorus gradients. Journal of Ecology, 2000, 88(6): 964-977
[49]	Pensa M, Karu H, Luud A, et al.Within-species correlations in leaf traits of three boreal plant species along a latitudinal gradient. Plant Ecology, 2010, 208(1): 155-166
[50]	Wright I J, Reich P B, Cornelissen J H, et al.Modulation of leaf economic traits and trait rela-tionships by climate. Global Ecology and Biogeo-graphy, 2005, 14(5): 411-421
[51]	Cordell S, Goldstein G, Meinzer F C, et al.Mor-phological and physiological adjustment to N and P fertilization in nutrient-limited Metrosideros polymor-pha canopy trees in Hawaii. Tree Physiology, 2001, 21: 43-50
[52]	Bubier J L, Smith R, Juutinen S, et al.EVects of nutrient addition on leaf chemistry, morphology, and photosynthetic capacity of three bog shrubs. Oeco-logia, 2011, 167: 355-368
[53]	Van de Weg M J, Shaver G R, Salmon V G. Contrasting effects of long term versus short-term nitrogen addition on photosynthesis and respiration in the Arctic. Plant Ecology, 2013, 214: 1273-1286