氮添加对山西太岳山天然油松林主要植物叶片性状的影响

doi:10.17521/cjpe.2015.1043

摘要/Abstract

摘要：

为探讨植物性状对大气氮沉降的响应与适应机制, 该文以中国特有的、在北方温性针叶林中广泛分布的天然油松(Pinus tabuliformis)林为研究对象, 在2009-2013年开展了氮添加对植物叶片性状影响的野外控制试验, 4个氮添加浓度分别为0 kg·hm^-2·a^-1 (CK)、50 kg·hm^-2·a^-1 (低氮)、100 kg·hm^-2·a^-1 (中氮)和150 kg·hm^-2·a^-1(高氮)。试验过程中分别测定了油松、蒙古栎(Quercus mongolica)、茶条槭(Acer ginnala)、毛榛(Corylus mandshurica)、沙梾(Cornus bretschneideri)、绣线菊(Spiraea salicifolia)、金银忍冬(Lonicera maackii)、羊须草(Carex callitrichos)、龙常草(Diarrhena mandshurica)、大火草(Anemone tomentosa)和玉竹(Polygonatum odoratum)等11种主要植物的9种叶片性状, 包括叶厚度(LT)、比叶面积(SLA)、干物质含量(LDMC)、叶氮含量(LNC)、叶磷含量(LPC)等。结果表明: 1)在氮添加影响下, 玉竹等个别物种的LT和SLA、绣线菊等部分物种的叶面积(LA)和LDMC差异显著, 上述所有物种的LNC与大多数物种的叶绿素含量(CC)、LPC显著增加, 油松等9种植物叶片N:P发生显著变化, 不同年龄、不同类型的植物叶片对氮添加的响应不同。2)叶性状之间普遍存在显著相关性, 如SLA与LNC和LPC极显著正相关, LT与LNC和LPC极显著负相关, 且相关性随氮添加强度变化。3) 11种植物的叶片特征空间分布规律与叶经济谱的描述一致, 氮添加使植物在特征空间中的位置向叶片薄、生长快、叶寿命短的“快速投资-收益型”一端发生移动; 在垂直方向上, 阔叶乔木、灌木及草本的位置与针叶乔木的移动方向相反。当环境改变时, 植物会改变生存策略, 调整资源分配, 从而保证物种间相对位置和群落整体结构的稳定性。叶经济谱的形成不依赖于环境的变化, 而是植物一种固有的属性。

关键词: 大气氮沉降, 叶经济谱, 叶片功能性状, 天然油松林, 氮添加

Abstract:

Aims Our objectives were to explore the effects of simulated nitrogen deposition on leaf traits and ecological stoichiometry characteristics of common species in natural Pinus tabuliformis forests.
Methods We conducted the experiment of nitrogen (N) addition from 2009 to 2013 in the natural Pinus tabuliformis forests in Taiyue Mountain, Shanxi, China. The levels of N addition were 0 (control), 50 (low-N), 100 (medium-N) and 150 (high-N) kg·hm^-2·a^-1, respectively. Eleven common plant species in 12 20 m × 20 m plots were selected, including Pinus tabuliformis, Quercus mongolica, Acer ginnala, Corylus mandshurica, Cornus bretschneideri, Spiraea salicifolia, Lonicera maackii, Carex callitrichos, Diarrhena mandshurica, Anemone tomentosa, and Polygonatum odoratum. Nine leaf traits were measured, including leaf thickness (LT), specific leaf area (SLA), leaf dry matter content (LDMC), leaf nitrogen content (LNC), leaf phosphorus content (LPC), and other four.
Important findings We found that: 1) LT and SLA of Polygonatum odoratum significantly differed among four levels of N addition. Leaf area (LA) and LDMC of several species, such as Spiraea salicifolia, had significant difference among the N addition concentration. LNC of all species, chlorophyll content (CC) and LPC of most species increased significantly with the addition of N. Leaf N:P of 9 species varied significantly, and leaves with different types and ages showed different responses to N addition. 2) Leaf traits were significantly correlated with each other. For instance, SLA was significantly positively correlated with LNC and LPC. In contrast LT was negatively connected with LNC and LPC. In addition, the degree of correlation changed with the level of N addition. 3) The pattern of species distribution in leaf trait space was consistent with the prediction from the theory of Leaf Economic Spectrum (LES). N addition drove species moving along axis 1 in the trait space, and propelled them towards different directions along axis 2, which indicated that these species tended to take the “fast investment-return” strategy. These results suggested that with the change of environmental conditions, plants changed their survival strategy and adjusted resource allocation to maintain the stability of communities. This is the inherent characteristic of plants, thus the formation of LES did not depend on the environment change.

Key words: atmospheric nitrogen deposition, leaf economic spectrum, leaf functional traits, natural Pinus tabuliformis forest, nitrogen addition

肖迪, 王晓洁, 张凯, 何念鹏, 侯继华. 氮添加对山西太岳山天然油松林主要植物叶片性状的影响. 植物生态学报, 2016, 40(7): 686-701. DOI: 10.17521/cjpe.2015.1043

Di XIAO, Xiao-Jie WANG, Kai ZHANG, Nian-Peng HE, Ji-Hua HOU. Effects of nitrogen addition on leaf traits of common species in natural Pinus tabuliformis forests in Taiyue Mountain, Shanxi Province, China. Chinese Journal of Plant Ecology, 2016, 40(7): 686-701. DOI: 10.17521/cjpe.2015.1043

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2015.1043

https://www.plant-ecology.com/CN/Y2016/V40/I7/686

图/表 9

表1 天然油松林各处理样地的林分特征和土壤理化性质

Table 1 Stand characteristics and soil physical-chemical properties of the nitrogen-loaded plots in the natural forest of Pinus tabuliformis

处理 Treatment	林分特征 Stand characteristics						土壤理化性质 Physical-chemical properties of soil
	林龄 Stand age (a)	密度 Density (plant·hm^-2)	平均胸径 Mean diameter at breast height (cm)	平均株高 Mean plant height (m)	坡度 Slope (°)		土壤容重 Soil volume weight (g·cm^-3)	土壤pH Soil pH	土壤有机碳含量 Soil organic carbon content (mg·g^-1)	土壤氮含量 Soil nitrogen content (mg·g^-1)	土壤磷含量 Soil phosphorus content (mg·g^-1)
	对照 Control	75	1 267	23.9	17.7	24		0.94	7.12	45.30	1.70	0.43
低氮 Low-N	75	1 567	20.6	17.8	21		0.97	7.12	61.02	2.14	0.38
中氮 Medium-N	75	1 208	23.5	17.4	25		1.07	7.19	42.14	1.91	0.38
高氮 High-N	75	1 225	23.4	19	23		1.06	7.28	36.08	2.25	0.44

图1 氮添加对叶厚度(LT, A)、叶面积(LA, B)、比叶面积(SLA, C)、叶绿素含量(CC, D)和叶干物质含量(LDMC, E)的影响(平均值±标准误差, n = 3)。Ag, 茶条槭; At, 大火草; Cb, 沙梾; Cc, 羊须草; Cm, 毛榛; Dm, 龙常草; Lm, 金银忍冬; Po, 玉竹; Pt-cy, 油松(当年生叶); Pt-fy, 油松(往年生叶); Qm, 蒙古栎; Ss, 绣线菊。CK, 氮添加浓度为0 kg·hm-2·a-1; LN, 氮添加浓度为50 kg·hm-2·a-1; MN, 氮添加浓度为100 kg·hm-2·a-1; HN, 氮添加浓度为150 kg·hm-2·a-1。不同小写字母表示氮添加处理间在p < 0.05水平上差异显著。

Fig. 1 Effects of nitrogen addition on leaf thickness (LT, A)、leaf area (LA, B)、specific leaf area (SLA, C)、chlorophyll content (CC, D) and leaf dry matter content (LDMC, E)(mean ± SE, n = 3). Ag, Acer ginnala; At, Anemone tomentosa; Cb, Cornus bretchneideri; Cc, Carex callitrichos; Cm, Corylus mandshurica; Dm, Diarrhena mandshurica; Lm, Lonicera maackii; Po, Polygonatum odoratum; Pt-cy, Pinus tabuliformis (current-year leaves); Pt-fy, P. tabuliformis (former-year leaves); Qm, Quercus mongolica; Ss, Spiraea salicifolia. CK, 0 kg·hm-2·a-1 nitrogen; LN, 50 kg·hm-2·a-1 nitrogen; MN, 100 kg·hm-2·a-1 nitrogen; HN, 150 kg·hm-2·a-1 nitrogen. Different lowercase letters indicate the significant difference at 5% level between nitrogen treatments.

图2 氮添加对叶碳含量(LCC, A)、叶氮含量(LNC, B)、叶磷含量(LPC, C)及氮磷比(N:P, D)的影响(平均值±标准误差, n = 3)。Ag, 茶条槭; At, 大火草; Cb, 沙梾; Cc, 羊须草; Cm, 毛榛; Dm, 龙常草; Lm, 金银忍冬; Po, 玉竹; Pt-cy, 油松(当年生叶); Pt-fy, 油松(往年生叶); Qm, 蒙古栎; Ss, 绣线菊。CK, 氮添加浓度为0 kg·hm-2·a-1; LN, 氮添加浓度为50 kg·hm-2·a-1; MN, 氮添加浓度为100 kg·hm-2·a-1; HN, 氮添加浓度为150 kg·hm-2·a-1。不同小写字母表示氮添加处理间在p < 0.05水平上差异显著。

Fig. 2 Effects of nitrogen addition on leaf carbon content (LCC, A), leaf nitrogen content (LNC, B), leaf phosphorus content (LPC, C), and N-P ratio (N:P, D)(mean ± SE, n = 3). Ag, Acer ginnala; At, Anemone tomentosa; Cb, Cornus bretchneideri; Cc, Carex callitrichos; Cm, Corylus mandshurica; Dm, Diarrhena mandshurica; Lm, Lonicera maackii; Po, Polygonatum odoratum; Pt-cy, Pinus tabuliformis (current-year leaves); Pt-fy, P. tabuliformis (former-year leaves); Qm, Quercus mongolica; Ss, Spiraea salicifolia. CK, 0 kg·hm-2·a-1 nitrogen; LN, 50 kg·hm-2·a-1 nitrogen; MN, 100 kg·hm-2·a-1 nitrogen; HN, 150 kg·hm-2·a-1 nitrogen. Different lowercase letters indicate the significant difference at 5% level between nitrogen treatments.

表2 不同生活型植物叶片性状对氮添加的响应(平均值±标准误差)

Table 2 Variation of leaf traits in different life forms under different levels of nitrogen addition (mean ± SE)

叶性状 Leaf trait	处理 Treatment	生活型 Life form
叶性状 Leaf trait	处理 Treatment	针叶乔木 Coniferous tree	阔叶乔木 Broadleaf tree	灌木 Shrub	草本 Herb
叶厚度 Leaf thickness (μm)	CK LN MN HN	716.23 ± 15.16^a 675.67 ± 27.71^a 689.33 ± 27.24^a 661.00 ± 17.85^a	160.45 ± 10.44^a 136.88 ± 10.45^a 147.48 ± 5.23^a 154.85 ± 9.76^a	167.45 ± 18.16^a 166.78 ± 13.89^a 152.16 ± 7.41^a 167.69 ± 16.35^a	199.43 ± 28.20^a 198.07 ± 23.30^a 168.19 ± 20.76^a 161.35 ± 17.11^a
叶面积 Leaf area (cm²)	CK LN MN HN	6.59 ± 0.14^ab 7.13 ± 0.33^a 7.18 ± 0.01^a 6.45 ± 0.12^b	22.91 ± 2.70^a 23.92 ± 4.38^a 29.33 ± 4.91^a 26.49 ± 3.74^a	25.59 ± 5.56^a 23.79 ± 4.83^a 25.97 ± 5.02^a 30.02 ± 5.96^a	39.75 ± 12.70^a 39.36 ± 12.85^a 44.70 ± 13.22^a 51.44 ± 20.18^a
比叶面积 Specific leaf area (m²·kg^-1)	CK LN MN HN	12.42 ± 0.53^a 12.62 ± 0.33^a 11.54 ± 0.40^a 12.67 ± 0.79^a	24.32 ± 1.36^ab 25.95 ± 0.91^a 22.19 ± 1.43^ab 21.73 ± 1.19^b	36.06 ± 0.86^a 35.15 ± 1.27^a 32.66 ± 1.69^a 32.00 ± 1.56^a	34.22 ± 1.12^a 33.30 ± 0.92^a 32.98 ± 1.16^a 31.60 ± 1.09^a
叶绿素含量 Chlorophyll content (SPAD)	CK LN MN HN	45.73 ± 1.36^a 40.37 ± 1.34^a 44.03 ± 1.95^a 42.23 ± 2.87^a	43.78 ± 1.91^b 46.87 ± 2.17^ab 48.75 ± 1.82^ab 49.47 ± 1.21^a	35.20± 1.46^a 37.03 ± 1.60^a 39.65 ± 1.83^a 39.71 ± 1.83^a	30.78 ± 1.50^a 31.81 ± 1.74^a 34.86 ± 1.47^a 33.62 ± 1.43^a
叶干物质含量 Leaf dry matter content (g·kg^-1)	CK LN MN HN	391.87 ± 5.43^b 421.10 ± 7.77^a 420.03 ± 3.91^a 416.17 ± 7.31^a	381.12 ± 15.37^a 377.22 ± 3.14^a 395.85 ± 11.58^a 384.52 ± 5.94^a	284.66 ± 12.41^a 287.51 ± 11.45^a 290.32 ± 15.77^a 295.34 ± 11.52^a	236.01 ± 16.81^a 241.20 ± 16.62^a 213.13 ± 23.40^a 249.49 ± 11.31^a
叶有机碳含量 Leaf carbon content (mg·g^-1)	CK LN MN HN	587.59 ± 10.05^a 541.67 ± 27.75^a 553.05 ± 1.01^a 548.83 ± 4.88^a	473.87 ± 15.47^a 486.62 ± 17.09^a 481.64 ± 13.02^a 487.60 ± 13.23^a	484.68 ± 17.84^a 450.92 ± 7.49^a 454.37 ± 6.12^a 468.28 ± 11.67^a	452.51 ± 7.97^a 441.45 ± 8.30^a 449.41 ± 9.61^a 450.11 ± 6.43^a
叶氮含量 Leaf nitrogen content (mg·g^-1)	CK LN MN HN	12.09 ± 0.39^b 13.48 ± 0.47^a 11.91 ± 0.50^b 12.50 ± 0.38^ab	25.25 ± 0.79^Cc 26.60 ± 0.32^BCb 27.56 ± 0.47^Bb 29.78 ± 0.37^Aa	22.47 ± 0.61^Bc 25.63 ± 0.56^Ab 25.81 ± 0.47^Ab 26.93 ± 0.45^Aa	22.96 ± 0.75^Cc 28.04 ± 0.80^Bb 28.20 ± 0.62^Bb 30.47 ± 0.74^Aa
叶磷含量 Leaf phosphorus content (mg·g^-1)	CK LN MN HN	1.08 ± 0.12^a 1.02 ± 0.10^a 1.06 ± 0.11^a 0.98 ± 0.07^a	1.60 ± 0.11^ABb 1.48 ± 0.05^Bb 1.70 ± 0.07^ABab 1.92 ± 0.17^Aa	1.17 ± 0.05^Cc 1.41 ± 0.03^Bb 1.71 ± 0.07^Aa 1.75 ± 0.05^Aa	1.26 ± 0.04^Bc 1.68 ± 0.08^Ab 1.91 ± 0.08^Aa 1.86 ± 0.08^Aab
叶氮磷比 leaf N:P	CK LN MN HN	12.54 ± 1.11^a 14.75 ± 1.14^a 12.44 ± 0.89^a 14.40 ± 0.80^a	16.39 ± 0.96^a 18.16 ± 0.46^a 16.44 ± 0.52^a 16.70 ± 1.08^a	19.86 ± 0.89^Aa 18.47 ± 0.61^ABa 15.80 ± 0.72^Bb 15.77 ± 0.59^Bb	18.50 ± 0.50^Aa 17.26 ± 0.66^ABab 15.16 ± 0.44^Bc 16.95 ± 0.59^ABb

表2 不同生活型植物叶片性状对氮添加的响应(平均值±标准误差)

Table 2 Variation of leaf traits in different life forms under different levels of nitrogen addition (mean ± SE)

叶性状 Leaf trait	处理 Treatment	生活型 Life form
叶性状 Leaf trait	处理 Treatment	针叶乔木 Coniferous tree	阔叶乔木 Broadleaf tree	灌木 Shrub	草本 Herb
叶厚度 Leaf thickness (μm)	CK LN MN HN	716.23 ± 15.16^a 675.67 ± 27.71^a 689.33 ± 27.24^a 661.00 ± 17.85^a	160.45 ± 10.44^a 136.88 ± 10.45^a 147.48 ± 5.23^a 154.85 ± 9.76^a	167.45 ± 18.16^a 166.78 ± 13.89^a 152.16 ± 7.41^a 167.69 ± 16.35^a	199.43 ± 28.20^a 198.07 ± 23.30^a 168.19 ± 20.76^a 161.35 ± 17.11^a
叶面积 Leaf area (cm²)	CK LN MN HN	6.59 ± 0.14^ab 7.13 ± 0.33^a 7.18 ± 0.01^a 6.45 ± 0.12^b	22.91 ± 2.70^a 23.92 ± 4.38^a 29.33 ± 4.91^a 26.49 ± 3.74^a	25.59 ± 5.56^a 23.79 ± 4.83^a 25.97 ± 5.02^a 30.02 ± 5.96^a	39.75 ± 12.70^a 39.36 ± 12.85^a 44.70 ± 13.22^a 51.44 ± 20.18^a
比叶面积 Specific leaf area (m²·kg^-1)	CK LN MN HN	12.42 ± 0.53^a 12.62 ± 0.33^a 11.54 ± 0.40^a 12.67 ± 0.79^a	24.32 ± 1.36^ab 25.95 ± 0.91^a 22.19 ± 1.43^ab 21.73 ± 1.19^b	36.06 ± 0.86^a 35.15 ± 1.27^a 32.66 ± 1.69^a 32.00 ± 1.56^a	34.22 ± 1.12^a 33.30 ± 0.92^a 32.98 ± 1.16^a 31.60 ± 1.09^a
叶绿素含量 Chlorophyll content (SPAD)	CK LN MN HN	45.73 ± 1.36^a 40.37 ± 1.34^a 44.03 ± 1.95^a 42.23 ± 2.87^a	43.78 ± 1.91^b 46.87 ± 2.17^ab 48.75 ± 1.82^ab 49.47 ± 1.21^a	35.20± 1.46^a 37.03 ± 1.60^a 39.65 ± 1.83^a 39.71 ± 1.83^a	30.78 ± 1.50^a 31.81 ± 1.74^a 34.86 ± 1.47^a 33.62 ± 1.43^a
叶干物质含量 Leaf dry matter content (g·kg^-1)	CK LN MN HN	391.87 ± 5.43^b 421.10 ± 7.77^a 420.03 ± 3.91^a 416.17 ± 7.31^a	381.12 ± 15.37^a 377.22 ± 3.14^a 395.85 ± 11.58^a 384.52 ± 5.94^a	284.66 ± 12.41^a 287.51 ± 11.45^a 290.32 ± 15.77^a 295.34 ± 11.52^a	236.01 ± 16.81^a 241.20 ± 16.62^a 213.13 ± 23.40^a 249.49 ± 11.31^a
叶有机碳含量 Leaf carbon content (mg·g^-1)	CK LN MN HN	587.59 ± 10.05^a 541.67 ± 27.75^a 553.05 ± 1.01^a 548.83 ± 4.88^a	473.87 ± 15.47^a 486.62 ± 17.09^a 481.64 ± 13.02^a 487.60 ± 13.23^a	484.68 ± 17.84^a 450.92 ± 7.49^a 454.37 ± 6.12^a 468.28 ± 11.67^a	452.51 ± 7.97^a 441.45 ± 8.30^a 449.41 ± 9.61^a 450.11 ± 6.43^a
叶氮含量 Leaf nitrogen content (mg·g^-1)	CK LN MN HN	12.09 ± 0.39^b 13.48 ± 0.47^a 11.91 ± 0.50^b 12.50 ± 0.38^ab	25.25 ± 0.79^Cc 26.60 ± 0.32^BCb 27.56 ± 0.47^Bb 29.78 ± 0.37^Aa	22.47 ± 0.61^Bc 25.63 ± 0.56^Ab 25.81 ± 0.47^Ab 26.93 ± 0.45^Aa	22.96 ± 0.75^Cc 28.04 ± 0.80^Bb 28.20 ± 0.62^Bb 30.47 ± 0.74^Aa
叶磷含量 Leaf phosphorus content (mg·g^-1)	CK LN MN HN	1.08 ± 0.12^a 1.02 ± 0.10^a 1.06 ± 0.11^a 0.98 ± 0.07^a	1.60 ± 0.11^ABb 1.48 ± 0.05^Bb 1.70 ± 0.07^ABab 1.92 ± 0.17^Aa	1.17 ± 0.05^Cc 1.41 ± 0.03^Bb 1.71 ± 0.07^Aa 1.75 ± 0.05^Aa	1.26 ± 0.04^Bc 1.68 ± 0.08^Ab 1.91 ± 0.08^Aa 1.86 ± 0.08^Aab
叶氮磷比 leaf N:P	CK LN MN HN	12.54 ± 1.11^a 14.75 ± 1.14^a 12.44 ± 0.89^a 14.40 ± 0.80^a	16.39 ± 0.96^a 18.16 ± 0.46^a 16.44 ± 0.52^a 16.70 ± 1.08^a	19.86 ± 0.89^Aa 18.47 ± 0.61^ABa 15.80 ± 0.72^Bb 15.77 ± 0.59^Bb	18.50 ± 0.50^Aa 17.26 ± 0.66^ABab 15.16 ± 0.44^Bc 16.95 ± 0.59^ABb

表3 叶性状间Pearson相关系数及氮添加处理和物种的影响效应分析

Table 3 Pearson correlation coefficients among leaf traits and the effects of N-treatment and species

	LT	LA	SLA	CC	LDMC	LCC	LNC	LPC	N:P	N-Tr.	Sp.	Sig. (N-Tr.× Sp.)
LT	1									-0.06	-0.50^**	0.00
LA	-0.17^**	1								0.06	0.37^**	0.00
SLA	-0.73^**	0.42^**	1							-0.12^*	0.71^**	0.01
CC	0.10	0.25^**	-0.34^**	1						0.18^**	-0.30^**	0.00
LDMC	0.11	-0.43^**	-0.56^**	0.35^**	1					0.04	-0.72^**	0.00
LCC	0.44^**	-0.55^**	-0.64^**	0.22^**	0.52^**	1				-0.04	-0.58^**	0.00
LNC	-0.73^**	0.56^**	0.71^**	0.13^*	-0.37^**	-0.60^**	1			0.25^**	0.62^**	0.00
LPC	-0.46^**	0.42^**	0.50^**	0.03	-0.40^**	-0.54^**	0.71^**	1		0.34^**	0.39^**	0.00
N:P	-0.36^**	0.17^**	0.29^**	0.14^*	0.08	-0.06	0.29^**	-0.41^**	1	-0.17^**	0.26^**	0.00

图3 氮添加对叶氮含量(LNC)与比叶面积(SLA) (A)、LNC与叶厚度(LT) (B)、叶磷含量(LPC)与SLA (C)和LPC与LT (D)关系的影响。CK, 氮添加浓度为0 kg·hm-2·a-1; LN, 氮添加浓度为50 kg·hm-2·a-1; MN, 氮添加浓度为100 kg·hm-2·a-1; HN, 氮添加浓度为150 kg·hm-2·a-1。

Fig. 3 Effects of nitrogen addition on leaf nitrogen content (LNC) vs. specific leaf area (SLA) (A), LNC vs. leaf thickness (LT) (B), leaf phosphorus content (LPC) vs. SLA (C) and LPC vs. LT (D). CK, 0 kg·hm-2·a-1 nitrogen; LN, 50 kg·hm-2·a-1 nitrogen; MN, 100 kg·hm-2·a-1 nitrogen; HN, 150 kg·hm-2·a-1 nitrogen.

表4 氮添加处理对叶氮含量、叶磷含量与比叶面积、叶厚度比值影响的单因素方差分析(平均值±标准误差, n = 36)

Table 4 Results of one-way ANOVA for the effect of nitrogen addition on the ratio of leaf nitrogen content (LNC) over specific leaf area (SLA) and leaf thickness (LT), and leaf phosphorus content (LPC) over specific leaf area (SLA) and leaf thickness (LT) (mean ± SE, n = 36)

处理 Treatment	叶氮含量/比叶面积 LNC:SLA	叶磷含量/比叶面积 LPC:SLA	叶氮含量/叶厚度 LNC:LT	叶磷含量/叶厚度 LPC:LT
CK	0.74 ± 0.022^d	0.05 ± 0.002^b	0.12 ± 0.007^c	0.01 ± 0.000^b
LN	0.85 ± 0.018^c	0.05 ± 0.002^b	0.15 ± 0.009^bc	0.01 ± 0.001^b
MN	0.92 ± 0.027^b	0.06 ± 0.002^a	0.16 ± 0.009^ab	0.01 ± 0.001^a
HN	1.05 ± 0.033^a	0.07 ± 0.003^a	0.18 ± 0.012^a	0.01 ± 0.001^a

表5 叶性状在主成分分析中的载荷

Table 5 Loadings of leaf traits in principal components analyses

载荷 Loading	叶厚度 Leaf thickness	叶面积 Leaf area	比叶面积 Specific leaf area	叶绿素含量 Chlorophyll content	叶干物质含量 Leaf dry matter content	叶碳含量 Leaf carbon content	叶氮含量 Leaf nitrogen content	叶磷含量 Leaf phosphorus content	氮磷比 N-P ratio	氮添加处理 N addition treatment
主成分1 Principal component 1	0.70	-0.64	-0.83	0.09	0.58	0.80	-0.91	-0.80	-0.11	-0.20
主成分2 Principal component 2	-0.40	0.08	0.16	0.32	0.33	0.10	0.24	-0.43	0.94	-0.57

图4 天然油松林主要植物的叶性状主成分分析结果: 天然油松林主要植物的叶性状载荷(A)、因子得分(B)及对氮添加的响应(C)。分析前对数据进行了log10转化。CC, 叶绿素含量; LA, 叶面积; LCC, 叶有机碳含量; LDMC, 叶干物质含量; LNC, 叶氮含量; LPC, 叶磷含量; LT, 叶厚度; N:P, 氮磷比; SLA, 比叶面积。Ag, 茶条槭; At, 大火草; Cb, 沙梾; Cc, 羊须草; Cm, 毛榛; Dm, 龙常草; Lm, 金银忍冬; Po, 玉竹; Pt-cy, 油松(当年生叶); Pt-fy, 油松(往年生叶); Qm, 蒙古栎; Ss, 绣线菊。C, 针叶乔木; B, 阔叶乔木; H, 草本; S, 灌木。CK, 氮添加浓度为0 kg·hm-2·a-1; LN, 氮添加浓度为50 kg·hm-2·a-1; MN, 氮添加浓度为100 kg·hm-2·a-1; HN, 氮添加浓度为150 kg·hm-2·a-1。

Fig. 4 Results of principle component analyses (PCA) of leaf traits across common species in the natural Pinus tabuliformis forest. Loadings of leaf traits across primary species in the natural forest of P. tabuliformis (A), factor scores of different species (B) and responses to nitrogen addition (C). Data were log10-transformed before analysis. CC, chlorophyll content; LA, leaf area; LCC, leaf carbon content; LDMC, leaf dry matter content; LNC, leaf nitrogen content; LPC, leaf phosphorus content; LT, leaf thickness; N:P, N-P ratio; SLA, specific leaf area. Ag, Acer ginnala; At, Anemone tomentosa; Cb, Cornus bretchneideri; Cc, Carex callitrichos; Cm, Corylus mandshurica; Dm, Diarrhena mandshurica; Lm, Lonicera maackii; Po, Polygonatum odoratum; Pt-cy, Pinus tabuliformis (current-year leaves); Pt-fy, P. tabuliformis (former-year leaves); Qm, Quercus mongolica; Ss, Spiraea salicifolia. C, coniferous tree; B, broadleaf tree; H, herb; S, shrub. CK, 0 kg·hm-2·a-1 nitrogen; LN, 50 kg·hm-2·a-1 nitrogen; MN, 100 kg·hm-2·a-1 nitrogen; HN, 150 kg·hm-2·a-1 nitrogen.

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