大兴安岭泥炭地植物叶片碳氮磷含量及其化学计量学特征

doi:10.17521/cjpe.2018.0214

植物生态学报 ›› 2018, Vol. 42 ›› Issue (12): 1154-1167.DOI: 10.17521/cjpe.2018.0214

所属专题：生态化学计量

大兴安岭泥炭地植物叶片碳氮磷含量及其化学计量学特征

李瑞¹,胡朝臣¹,许士麒¹,吴迪¹,董玉平¹,孙新超¹,毛瑢^2,³,王宪伟^2,^*(),刘学炎^1,^*()

1 天津大学表层地球系统科学研究院, 天津 300072
2 中国科学院东北地理与农业生态研究所, 长春 130102
3 江西农业大学林学院, 南昌 330045

收稿日期:2018-08-27 修回日期:2018-12-03 出版日期:2018-12-20 发布日期:2019-04-04
通讯作者: 王宪伟,刘学炎 ORCID: 0000-0003-1097-151,李瑞 ORCID: 0000-0002-1793-7997
基金资助:
国家自然科学基金(41471056);国家自然科学基金(41522301);国家自然科学基金(41730855);国家重点研发计划(2016YFA0600802)

Leaf C, N, and P concentrations and their stoichiometry in peatland plants of Da Hinggan Ling, China

LI Rui¹,HU Chao-Chen¹,XU Shi-Qi¹,WU Di¹,DONG Yu-Ping¹,SUN Xin-Chao¹,MAO Rong^2,³,WANG Xian-Wei^2,^*(),LIU Xue-Yan^1,^*()

1 Institute of Surface-Earth System Science, Tianjin University, Tianjin 300072, China
2 Northeast Institute of Geography and Agroecology, Chinese Academy of Sciences, Changchun 130102, China
3 College of Forestry, Jiangxi Agricultural University, Nanchang 330045, China

Received:2018-08-27 Revised:2018-12-03 Online:2018-12-20 Published:2019-04-04
Contact: WANG Xian-Wei,LIU Xue-Yan ORCID: 0000-0003-1097-151, LI Rui ORCID: 0000-0002-1793-7997
Supported by:
Supported by the National Natural Science Foundation of China(41471056);Supported by the National Natural Science Foundation of China(41522301);Supported by the National Natural Science Foundation of China(41730855);the National Key R & D Program of China(2016YFA0600802)

摘要/Abstract

摘要：

叶片碳(C)、氮(N)、磷(P)含量及其化学计量特征为植物养分状况和元素限制性提供依据。为了解不同生活型植物叶片C、N、P化学计量特征的变化,该研究测定、分析了大兴安岭地区18个泥炭地常见的3种草本植物——白毛羊胡子草(Eriophorum vaginatum)、玉簪薹草(Carex globularis)、小叶章(Deyeuxia angustifolia), 5种落叶灌木——柴桦(Betula fruticosa)、越桔柳(Salix myrtilloides)、细叶沼柳(Salix rosmarinifolia)、笃斯越桔(Vaccinium uliginosum)、越桔(Vaccinium vitis-idaea)和3种常绿灌木——杜香(Ledum palustre)、地桂(Chamaedaphne calyculata)、头花杜鹃(Rhododendron capitatum)的叶片C、N、P含量。结果表明: (1)落叶和常绿灌木叶片C、N、P含量总体高于草本植物而C:N、C:P、N:P低于草本植物, 说明不同生活型植物具有不同的养分利用策略,灌木叶片C、N、P储存高于草本植物而N、P利用效率低于草本植物; (2)小叶章和头花杜鹃叶片N:P小于10, 同时其N含量小于全球植物叶片平均N含量, 相比其他植物来说更易受N限制; (3)采样地点解释了叶片C、N、P指标变异的12.8%-40.8%, 植物种类对叶片C、N、P指标变异的解释量占9.3%-25.5%; (4)草本植物C、N、P指标的地点间变异系数高于落叶和常绿灌木, 草本植物C、N、P指标对地点因素变化的响应较灌木敏感; (5)草本植物N含量种间变异系数高于落叶和常绿灌木, 落叶灌木P含量种间变异系数高于草本植物和常绿灌木, 草本植物和落叶灌木N、P吸收的种间生理分化较常绿灌木高。

关键词: 大兴安岭, 北方泥炭地, 植物营养, 化学计量学

Abstract:

Aims Leaf carbon (C), nitrogen (N), and phosphorus (P) concentrations and their stoichiometry can provide a basis for plant nutrient status and element limitation. Our objective was to explore variations of leaf C:N:P stoichiometry in plants of different growth forms.

Methods We analyzed leaf C, N, and P concentrations in three graminoids (Eriophorum vaginatum, Carex globularis, Deyeuxia angustifolia), five deciduous shrubs (Betula fruticosa, Salix myrtilloides, Salix rosmarinifolia, Vaccinium vitis-idaea, Vaccinium uliginosum), and three evergreen shrubs (Ledum palustre, Chamaedaphne calyculata, Rhododendron capitatum) across 18 peatland sites in the Da Hinggan Ling, northeastern China.

Important findings (1) Leaf C, N, and P concentrations were higher, and the leaf C:N, C:P, and N:P values were lower, in deciduous and evergreen shrubs than in graminoids, indicating that plants of different growth forms had different nutrient utilization strategies. Shrubs had higher C, N and P storage and lower N and P use efficiency than graminoids. (2) Leaf N:P values in Deyeuxia angustifolia and R. capitatum were less than 10, and their leaf N concentrations were lower than the global mean leaf N concentration, indicating that those species were limited by N more than other plants. (3) The sampling sites explained 12.8%-40.8% of the variations in leaf C, N, and P stoichiometry, and plant species explained 9.3%-25.5%. (4) Graminoids had greater inter-site coefficient of variance (CV) values in leaf C, N, and P variables than deciduous and evergreen shrubs, indicating greater sensitive to site factors. (4) The inter-species CV values in leaf N were greater in graminoids than in deciduous and evergreen shrubs, and the inter-species CV values in leaf P were greater in deciduous shrubs than in graminoids and evergreen shrubs, indicating greater physiological differentiation in N and P use strategies in graminoids and deciduous shrubs than in evergreen shrubs.

Key words: Da Hinggan Ling, northern peatland, plant nutrition, stoichiometry

李瑞, 胡朝臣, 许士麒, 吴迪, 董玉平, 孙新超, 毛瑢, 王宪伟, 刘学炎. 大兴安岭泥炭地植物叶片碳氮磷含量及其化学计量学特征. 植物生态学报, 2018, 42(12): 1154-1167. DOI: 10.17521/cjpe.2018.0214

LI Rui, HU Chao-Chen, XU Shi-Qi, WU Di, DONG Yu-Ping, SUN Xin-Chao, MAO Rong, WANG Xian-Wei, LIU Xue-Yan. Leaf C, N, and P concentrations and their stoichiometry in peatland plants of Da Hinggan Ling, China. Chinese Journal of Plant Ecology, 2018, 42(12): 1154-1167. DOI: 10.17521/cjpe.2018.0214

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2018.0214

https://www.plant-ecology.com/CN/Y2018/V42/I12/1154

图/表 10

图1 大兴安岭泥炭地研究点地理位置示意图。

Fig. 1 Locations of the peatlands investigated in the Da Hinggan Ling.

表1 大兴安岭地区泥炭地研究点的气候和采集植物种信息

Table 1 Information on climate and plant species studied in the peatland sites in the Da Hinggan Ling

地点 Site	年平均气温MAT (℃)	年降水量 MAP (mm)	海拔 Elevation (m)	调查植物种(重复数) Plant species studied (replicates)
阿里河 Alihe	-5.57	481	490	A(2), C(3), D(3), E(3)
阿木尔 Amuer	-4.48	459	533	A(3), C(2), D(3), E(2), G(3), H(1), I(3)
大林河 Dalinhe	-4.20	443	466	A(3), D(3), E(3), G(3), I(3), J(3)
富克山 Fukeshan	-4.21	444	468	A(3), D(3), E(2), G(1), H(1), I(3), K(1)
根河 Genhe	-5.22	464	839	A(3), C(3), D(3), E(4), I(2), K(3)
呼源 Huyuan	-4.48	501	665	A(3), B(1), D(3), E(1), G(2), I(3), K(2)
呼中 Huzhong	-4.06	479	534	A(3), D(3), G(3), I(3), J(1)
金河 Jinhe	-5.41	477	862	A(3), C(2), D(3), G(1)
林海 Linhai	-5.51	520	571	A(3), C(3), D(3), F(2), G(3), I(1), K(3)
满归 Mangui	-4.78	464	620	A(3), D(3), E(2), G(3), J(3), K(3)
南瓮河 Nanwenghe	-2.71	498	485	A(3), B(3), C(3), D(3), E(3), F(3), G(3)
盘古 Pangu	-3.62	477	406	A(3), D(3), E(3), G(1), H(2), I(3), J(3)
十二站 Shi’erzhan	-2.40	495	394	A(3), C(2), D(3), E(3), G(2), J(1)
塔河 Tahe	-3.56	487	440	A(3), C(2), D(3), E(3), F(1)
图强 Tuqiang	-4.29	452	477	A(3), D(3), E(1), F(2), G(1), I(3), J(2), K(2)
新林 Xinlin	-3.63	498	526	A(3), D(3), E(3), G(3), I(3), J(3), K(2)
伊图里河 Ih Tol Gol	-5.13	475	822	A(3), C(2), D(3), G(2)
壮林 Zhuanglin	-4.48	459	528	A(3), D(3), G(3), I(3), J(2), K(1)

表2 大兴安岭泥炭地植物叶片C、N、P指标之间的相关性

Table 2 The correlations among leaf C, N, and P variables in peatland plants of Da Hinggan Ling

	C (%)	N (%)	P (%)	C:N	C:P
C (%)	1
N (%)	0.294^**	1
P (%)	0.069	0.287^**	1
C:N	-0.002	-0.886^**	-0.258^**	1
C:P	0.005	-0.292^**	-0.854^**	0.318^**	1
N:P	-0.007	0.238^**	-0.750^**	-0.242^**	0.812^**

图2 大兴安岭泥炭地植物叶片C、N、P含量(A、B、C)及其计量值(D、E、F)。黑色的短横线分别为每个植物种C、N、P指标的平均值。深灰、灰色和浅灰的箱子分别为草本、落叶灌木和常绿灌木植物, 箱子高度为25%-75%数值分布, 箱子中的横线为平均值, 箱须为标准偏差, 箱须上方不同的大写字母和下方不同的小写字母分别表示生活型之间和生活型内部物种之间的差异显著(p < 0.05)。Bf, 柴桦; Cc, 地桂; Cg, 玉簪薹草; Da, 小叶章; Ev, 白毛羊胡子草; Lp, 杜香; Rc, 头花杜鹃; Sm, 越桔柳; Sr, 细叶沼柳; Vu, 笃斯越桔; Vv, 越桔。

Fig. 2 Leaf C, N, and P concentrations (A, B, C) and their stoichiometric ratios (D, E, F) in peatland plants of the Da Hinggan Ling. The black and short horizontal lines are the average values for leaf C, N, and P variables for each species. Boxes in dark grey, grey, and light grey mark graminoids, deciduous shrub, and evergreen shrub plants, respectively. The box encompasses the 25th to 75th percentiles; lines in boxes mark the mean values, and whiskers are the standard deviation value. Different capital letters above boxes and lowercase letters below boxes mark significant differences among growth forms and species (p < 0.05). Bf, Betula fruticosa; Cc, Chamaedaphne calyculata; Cg, Carex globularis; Da, Deyeuxia angustifolia; Ev, Eriophorum vaginatum; Lp, Ledum palustre; Rc, Rhododendron capitatum; Sm, Salix myrtilloides; Sr, Salix rosmarinifolia; Vu, Vaccinium uliginosum; Vv, Vaccinium vitis-idaea.

表3 采样地点(D)和植物种类(S)对大兴安岭泥炭地植物叶片C、N、P含量和计量值影响的一般线性模型(GLM)结果

Table 3 Summary of General Linear Model (GLM) statistics, showing the effects of sampling site (D) and plant species (S) on leaf C, N, and P concentrations and stoichiometry in peatland plants of Da Hinggan Ling

变异来源 Source	C			N			P			C:N			C:P			N:P
变异来源 Source	MS	SS (%)	F	MS	SS (%)	F	MS	SS (%)	F	MS	SS (%)	F	MS	SS (%)	F	MS	SS (%)	F
D	84.3	38.7	24.016^*	1.7	40.8	29.415^*	0.04	25.8	13.031^*	378.2	38.3	23.732^*	74 004.0	23.9	9.548^*	67.2	12.8	5.883^*
S	17.9	14.0	5.094^*	0.2	9.7	4.131^*	0.01	16.9	5.022^*	53.4	9.3	3.353^*	29 120.8	16.0	3.757^*	78.5	25.5	6.872^*
D × S	5.5	20.6	1.580^*	0.1	26.4	2.349^*	0.01	26.2	1.699^*	31.3	25.7	1.967^*	8 315.6	21.0	1.073	18.4	27.5	1.614^*

图3 大兴安岭泥炭地植物叶片C、N、P含量(A、B、C)及其计量值(D、E、F)的地点间变异系数(KCV)。黑、灰、浅灰色直线分别为草本、落叶灌木和常绿灌木植物KCV的平均值, 植物名称缩写下面的数字为出现的地点数。Bf, 柴桦; Cc, 地桂; Cg, 玉簪薹草; Da, 小叶章; Ev, 白毛羊胡子草; Lp, 杜香; Rc, 头花杜鹃; Sm, 越桔柳; Sr, 细叶沼柳; Vu, 笃斯越桔; Vv, 越桔。

Fig. 3 Inter-site coefficient of variation (KCV) in leaf C, N, and P concentrations (A, B, C) and their stoichiometric ratios (D, E, F) in peatland plants of Da Hinggan Ling. Lines in black, grey, and light grey mark the mean KCV values of graminoids, deciduous shrubs and evergreen shrubs, respectively. The number below each species indicates the number of occurring locations. Bf, Betula fruticosa; Cc, Chamaedaphne calyculata; Cg, Carex globularis; Da, Deyeuxia angustifolia; Ev, Eriophorum vaginatum; Lp, Ledum palustre; Rc, Rhododendron capitatum; Sm, Salix myrtilloides; Sr, Salix rosmarinifolia; Vu, Vaccinium uliginosum; Vv, Vaccinium vitis-idaea.

图4 大兴安岭泥炭地植物叶片C、N、P含量(A、B、C)及其计量值(D、E、F)的种间变异系数(ZCV)。黑、灰、浅灰色直线分别为草本、落叶灌木和常绿灌木植物ZCV的平均值, 地点名称后的数字分别为该地点草本、落叶灌木和常绿灌木植物种数。

Fig. 4 Inter-species coefficient of variation (ZCV) in leaf C, N, and P concentrations (A, B, C) and their stoichiometric ratios (D, E, F) in peatland plants of Da Hinggan Ling. Lines in black, grey, and light grey mark mean ZCV values for graminoids, decidous shrubs and evergreen shrubs, respectively. The numbers after site names indicate the number of species for graminoids, decidous and evergreen shrubs, respectively.

表4 大兴安岭泥炭地植物叶片C、N、P含量及其计量值与其他研究结果的比较

Table 4 Comparisons of leaf C, N, and P concentrations and their stoichiometry between plants of the Da Hinggan Ling peatlands and those in other studies

研究区域 Study region	C (%)			N (%)			P (%)			C:N			C:P			N:P			数据来源
研究区域 Study region	Mean	SD	n	Mean	SD	n	Mean	SD	n	Mean	SD	n	Mean	SD	n	Mean	SD	n	Data source
泥炭地 Peatlands	42.2	2.9	275	1.9	0.4	276	0.18	0.07	263	23.6	6.2	275	269	110	262	11.6	4.6	263	本研究 This Study
森林 Forests	48.0	5.3	102	1.8	0.5	102	0.20	0.12	102	29.1	9.5	102	314	152	102	11.5	5.0	102	Ren et al., 2012
荒漠 Deserts				1.1	0.8	276	0.10	0.08	276							11.5	5.1	276	Zhang et al., 2014
草原 Grasslands	43.8	3.0	213	2.8	0.9	213	0.19	0.84	525	17.9	5.7	213				15.3	5.2	525	He et al., 2006, 2008
中国 China				2.0	0.8	554	0.15	0.10	745							16.3	9.3	547	Han et al., 2005
全球 Globe	46.2	7.2	76	2.0	0.9	1 251	0.18	0.11	923	23.8	17.3	62	301	237	43	13.8	9.5	894	Elser et al., 2000a; Reich & Oleksyn, 2004

表5 大兴安岭泥炭地植物菌根类型、叶形和植物高度概况

Table 5 Plant mycorrhizal types, leaf shapes and plant height in the Da Hinggan Ling peatlands

生活型 Growth form	植物种 Species	菌根类型 Mycorrhizal type	植株高度 Plant height (cm)	叶形 Leaf shape	参考文献 Reference
草本植物 Graminoids	白毛羊胡子草 Eriophorum vaginatum	无菌根 Non-mycorrhizae	40-80	线形 Linear leaf	Hobbie & Hobbie, 2006; Wu & Hong, 2010
	玉簪薹草 Carex globularis	无菌根 Non-mycorrhizae	30-60	线形 Linear leaf
	小叶章 Deyeuxia angustifolia	丛枝菌根 Arbuscular mycorrhizae	30-100	线形 Linear leaf
落叶灌木 Deciduous shrubs	柴桦 Betula fruticosa	外生菌根 Ecto-mycorrhizae	50-150	卵形 Ovate leaf
	越桔柳 Salix myrtilloides	外生菌根 Ecto-mycorrhizae	30-80	椭圆形 Oblong leaf
	细叶沼柳 Salix rosmarinifolia	外生菌根 Ecto-mycorrhizae	50-100	披针形 Lanceolate leaf
	笃斯越桔 Vaccinium uliginosum	杜鹃花科菌根 Ericoid mycorrhizae	50-80	倒卵形 Obovate leaf
	越桔 Vaccinium vitis-idaea	杜鹃花科菌根 Ericoid mycorrhizae	10-30	倒卵形 Obovate leaf
常绿灌木 Evergreen shrubs	杜香 Ledum palustre	杜鹃花科菌根 Ericoid mycorrhizae	40-50	线形 Linear leaf
	地桂 Chamaedaphne calyculata	杜鹃花科菌根 Ericoid mycorrhizae	30-150	椭圆形 Oblong leaf
	头花杜鹃 Rhododendron capitatum	杜鹃花科菌根 Ericoid mycorrhizae	40-100	椭圆形 Oblong leaf

图5 大兴安岭泥炭地植物叶片N、P含量与N:P阈值。虚线分别表示N:P阈值为10, 14, 16和20。星星表示全球植物叶片N含量平均值(1.9%)和P含量平均值(0.12%)(Yan et al., 2017)。

Fig. 5 Leaf N and P concentrations and N:P threshold in peatland plants of Da Hinggan Ling. Dash lines represent the N:P ratios of 10, 14, 16 and 20, respectively. Star-shaped symbol represents the global mean leaf N concentration (1.9%) and P concentration (0.12%) (Yan et al., 2017). Bf, Betula fruticosa; Cc, Chamaedaphne calyculata; Cg, Carex globularis; Da, Deyeuxia angustifolia; Ev, Eriophorum vaginatum; Lp, Ledum palustre; Rc, Rhododendron capitatum; Sm, Salix myrtilloides; Sr, Salix rosmarinifolia; Vu, Vaccinium uliginosum; Vv, Vaccinium vitis-idaea.

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大兴安岭泥炭地植物叶片碳氮磷含量及其化学计量学特征

Leaf C, N, and P concentrations and their stoichiometry in peatland plants of Da Hinggan Ling, China

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