亚热带不同树种凋落叶分解对氮添加的响应

doi:10.17521/cjpe.2019.0299

植物生态学报 ›› 2020, Vol. 44 ›› Issue (3): 214-227.DOI: 10.17521/cjpe.2019.0299

所属专题：全球变化与生态系统；凋落物

亚热带不同树种凋落叶分解对氮添加的响应

陈思路^1,²,蔡劲松⁴,林成芳^1,^2,^3,^*(),宋豪威^1,²,杨玉盛^1,^2,³

¹福建师范大学地理科学学院, 福州 350007
²湿润亚热带山地生态国家重点实验室培育基地, 福州 350007
³福建三明森林生态系统与全球变化研究站, 福建三明 365000
⁴福建省三明市郊国有林场, 福建三明 365000

收稿日期:2019-11-04 接受日期:2020-02-06 出版日期:2020-03-20 发布日期:2020-04-30
通讯作者: 林成芳
基金资助:
国家自然科学基金(31770663);福建省自然科学基金(2018J01718)

Response of leaf litter decomposition of different tree species to nitrogen addition in a subtropical forest

CHEN Si-Lu^1,²,CAI Jin-Song⁴,LIN Cheng-Fang^1,^2,^3,^*(),SONG Hao-Wei^1,²,YANG Yu-Sheng^1,^2,³

¹School of Geographical Sciences, Fujian Normal University, Fuzhou 350007, China
²State Key Laboratory of Humid Subtropical Mountain Ecology, Fuzhou, 350007, China
³Sanming Research Station of Forest Ecosystem and Global Change, Sanming, Fujian 365000, China
⁴State-owned Forest Farm in the Suburbs of Sanming, Fujian Province, Sanming, Fujian 365000, China

Received:2019-11-04 Accepted:2020-02-06 Online:2020-03-20 Published:2020-04-30
Contact: Cheng-Fang LIN
Supported by:
National Natural Science Foundation of China(31770663);Natural Science Foundation of Fujian(2018J01718)

摘要/Abstract

摘要：

为探究不同质量凋落物对氮(N)沉降的响应, 该研究采用尼龙网袋分解法, 在亚热带福建三明格氏栲(Castanopsis kawakamii)自然保护区的米槠(Castanopsis carlesii)天然林, 选取4种本区常见的具有不同初始化学性质的树种凋落叶进行模拟N沉降(N添加)分解实验(施N水平为对照0和50 kg·hm ^-2·a ^-1)。研究结果表明: 在2年的分解期内, 对照处理的各树种凋落叶的分解速率依次为观光木(Michelia odora, 0.557 a ^-1)、米槠(0.440 a ^-1)、台湾相思(Acacia confusa, 0.357 a ^-1)、杉木(Cunninghamia lanceolata, 0.354 a ^-1); N添加处理凋落叶分解速率依次为观光木(0.447 a ^-1)、米槠(0.354 a ^-1)、杉木(0.291 a ^-1)、台湾相思(0.230 a ^-1), 除杉木凋落叶外, N添加显著降低了其他3种凋落叶分解速率。N添加不仅使4种树木凋落叶分解过程中的N释放减慢, 同时还抑制凋落叶化学组成中木质素和纤维素的降解; N添加在凋落叶分解过程中总体上提高β-葡萄糖苷酶(βG)和酸性磷酸酶活性, 对纤维素水解酶的活性影响不一致, 而降低β-N-乙酰氨基葡萄糖苷酶活性和酚氧化酶活性。凋落叶分解速率与凋落叶中的碳获取酶(βG)活性以及其化学组分中的可萃取物含量极显著正相关, 与初始碳浓度、纤维素和木质素含量极显著负相关, 与初始N含量没有显著相关性。凋落物类型和N添加的交互作用虽未影响干质量损失速率, 但对木质素和纤维素的降解具有显著效应。综上所述, 化学组分比初始N含量能更好地预测凋落叶分解速率, 而N添加主要通过抑制分解木质素的氧化酶(如PHO)来降低凋落叶分解速率。

关键词: 凋落叶分解, 氮沉降, 酶活性, 亚热带森林

Abstract:

Aims Litter decomposition plays a vital role in nutrient recycling of forest ecosystems. The decomposition rate of leaves can vary among tree species with different substrate quality and environmental conditions, such as the supply of exogenous nitrogen (N). However, the effects of exogenous N on leaf litter decomposition of different tree species in subtropical forests with high nitrogen deposition background remain poorly understood. Thus this study was designed to address the effect of N addition on litter decomposition of different tree species in a subtropical forest ecosystem.
Methods Leaf litters of four common tree species with contrasting substrate quality were collected and decomposed in fertilized (50 kg N·hm ^-2·a ^-1) and control (0 kg N·hm ^-2·a ^-1) plots for up to two years by using the nylon bag method, in Sanming Castanopsis kawakamii nature reserve of Fujian Province.
Important findings The litter decomposition rate in control plots ranked as follows: Michelia odora (0.557 a ^-1), Castanopsis carlesii (0.440 a ^-1), Acacia confusa (0.357 a ^-1), Cunninghamia lanceolata (0.354 a ^-1), while the decomposition rate in N addition plots ranked as follows: Michelia odora (0.447 a ^-1), Castanopsis carlesii (0.354 a ^-1), Cunninghamia lanceolata (0.291 a ^-1), Acacia confusa (0.230 a ^-1). Overall, N addition significantly increased the litter mass remaining of Michelia odora, Acacia confusa and Castanopsis carlesii, but not Cunninghamia lanceolata. N addition not only slowed down the release of N, but also retarded the degradation of lignin and cellulose in the decomposition process. Moreover, N addition increased the activities of β-glucosidase (βG) and acid phosphatase, had species-specific effects on the activity of cellulose hydrolase, and decreased the activity of β-N-acetylglucosaminidase and phenoloxidase (PHO). The litter mass loss rate was positively correlated with the activities of carbon acquiring enzyme (βG) in litter layer and the extractives, negatively correlated with carbon concentration, cellulose and lignin, but did not correlate significantly with the initial N concentration. Further analysis found an interactive effect of litter type and N addition on the degradation of cellulose and lignin, but not on dry mass loss. Overall, our results demonstrated that litter chemical components may be better parameters to predict the decomposition rate of leaf litters than the initial nutrient concentrations, and N addition could decrease leaf litter decomposition by inhibiting oxidase (e.g. PHO). We call for further experiments to involve more species and longer time for revealing the response of leaf litter decomposition and its extracellular enzyme activity to N addition.

Key words: leaf litter decomposition, nitrogen deposition, enzyme activities, subtropical forest

陈思路, 蔡劲松, 林成芳, 宋豪威, 杨玉盛. 亚热带不同树种凋落叶分解对氮添加的响应. 植物生态学报, 2020, 44(3): 214-227. DOI: 10.17521/cjpe.2019.0299

CHEN Si-Lu, CAI Jin-Song, LIN Cheng-Fang, SONG Hao-Wei, YANG Yu-Sheng. Response of leaf litter decomposition of different tree species to nitrogen addition in a subtropical forest. Chinese Journal of Plant Ecology, 2020, 44(3): 214-227. DOI: 10.17521/cjpe.2019.0299

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2019.0299

https://www.plant-ecology.com/CN/Y2020/V44/I3/214

图/表 11

表1 亚热带米槠天然林群落特征与土壤表层(0-10 cm)理化性质

Table 1 Community characteristics and topsoil (0-10 cm) physiochemical properties of a subtropical Castanopsis carlesii natural forest

林分特征 Forest characteristics				土壤理化性质 Soil physiochemical properties
林分密度 Stand density (株·hm^-2)	郁闭度 Canopy density	平均树高 Mean tree height (m)	平均胸径 Mean DBH (cm)	容重 Bulk density (g·cm^-3)	全C Total carbon (%)	全N Total nitrogen (%)	全P Total phosphorus (%)
1 955	0.89	11.9	20	1.12	2.45	0.186	0.021 7

表2 亚热带森林不同凋落叶初始化学性质(平均值±标准偏差, n = 3)

Table 2 Initial chemical properties of leaf litter in a subtropical forest (mean ± SD, n = 3)

树种 Species	C (g·kg^-1)	N ( g·kg^-1)	P ( g·kg^-1)	可萃取物 Extractive (%)	纤维素 Cellulose (%)	木质素 Lignin (%)	C:N	C:P	木质素:N Lignin:N	木质素:P Lignin:P
米槠 Castanopsis carlesii	486.44 ± 2.15	14.22 ± 0.19	0.34 ± 0.03	55.50 ± 1.58	24.50 ± 1.07	20.00 ± 1.02	34.21 ± 2.13	1 158.20 ± 0.78	1.41 ± 0.05	47.62± 1.56
杉木 Cunninghamia lanceolata	525.05 ± 2.39	16.03 ± 0.16	0.79 ± 0.10	51.40 ± 0.95	24.30 ± 1.02	24.30 ± 0.85	32.75 ± 0.95	664.62 ± 0.50	1.52 ± 0.05	30.76 ± 5.46
观光木 Michelia odora	390.24 ± 1.68	24 .26 ± 0.12	0.92 ± 0.05	55.20 ± 0.97	25.50 ± 0.95	19.30 ± 1.16	16.10 ± 0.26	424.45 ± 0.36	0.80 ± 0.28	20.98 ± 2.15
台湾相思 Acacia confusa	501.37 ± 2.81	27.61 ± 0.36	0.56 ± 0.02	51.10 ± 1.87	26.60 ± 0.45	22.30 ± 0.81	18.14 ± 0.09	894.69 ± 0.52	0.81 ± 0.35	39.82 ± 0.97

图1 亚热带森林不同处理凋落叶分解过程中质量残留率变化(平均值±标准偏差, n = 3)。A, 米槠凋落叶。B, 杉木凋落叶。C, 观光木凋落叶。D, 台湾相思凋落叶。CT, 对照; +N, 氮添加。*, p < 0.05; **, p < 0.01。

Fig. 1 Variations of litter mass remaining of different tree species during decomposition under different treatments in a subtropical forest (mean ± SD, n = 3). A, Castanopsis carlesii litter. B, Cunninghamia lanceolata litter. C, Michelia odora litter. D, Acacia confusa litter. CT, control; +N, nitrogen addition.

表3 亚热带森林不同处理凋落物分解质量残留率随时间的指数回归方程

Table 3 Exponential regression equations of mass remaining rate of leaf litter with time under different treatments in a subtropical forest

树种 Species	处理 Treatment	Olson负指数方程 Olson negative exponential equation	R²	分解常数 Decay constant k (a^-1)	年实际干质量损失率 Annual observed mass loss rate (%)	年预期干质量损失率 Annual predicted mass loss rate (%)	半分解时间 T_50% (a)	95%分解时间 T_95% (a)
米槠 Castanopsis carlesii	CT	Y = 91.26e^-0.440^x	0.949	0.440^Ba	47.794	42.194	1.292	5.941
米槠 Castanopsis carlesii	+N	Y = 96.35e^-0.354^x	0.976	0.354^Bb	34.257	32.435	1.836	8.232
杉木 Cunninghamia lanceolata	CT	Y = 95.79e^-0.354^x	0.969	0.354^Ca	36.242	30.661	1.862	8.366
杉木 Cunninghamia lanceolata	+N	Y = 94.91e^-0.291^x	0.968	0.291^BCa	29.510	28.215	2.245	10.167
观光木 Michelia odora	CT	Y = 81.65e^-0.557^x	0.899	0.557^Aa	49.333	38.934	0.993	5.124
观光木 Michelia odora	+N	Y = 88.03e^-0.447^x	0.934	0.447^Ab	39.768	36.776	1.334	6.481
台湾相思 Acacia confusa	CT	Y = 91.26e^-0.357^x	0.964	0.357^Ca	35.240	31.303	1.798	8.248
台湾相思 Acacia confusa	+N	Y = 96.35e^-0.230^x	0.908	0.230^Cb	24.887	26.328	2.685	13.479

表4 亚热带森林凋落叶分解质量损失、木质素、纤维素和氮释放的三因素重复测量方差分析

Table 4 Analysis (indicated by F values from ANOVA with three factors repeated measurements) of decomposition time, litter type, nitrogen addition and their interactions on litter mass and lignin, cellulose and N release in a subtropical forest

差异来源 Different source	质量损失率 Mass loss rate	木质素损失率 Lignin loss rate	纤维素损失率 Cellulose loss rate	氮释放率 N release rate
T	356.428^***	18 920.622^***	8 803.023^***	1 266.179^***
L	60.641^*	1 798.360^**	4 305.721^***	70.640^**
N	287.734^**	2 916.781^***	5 176.453^***	421.188^**
T ′ L	4.114	1 605.655^***	1 016.727^***	14.430^*
T ′ N	2.681	124.624^**	72.162^*	15.670^*
L ′ N	8.713	582.266^**	667.323^***	7.278
T ′ L ′ N	1.038	271.636^**	407.487^***	2.044

图2 亚热带森林不同处理凋落叶分解过程中木质素残留率变化(平均值±标准偏差, n = 3)。A, 米槠凋落叶。B, 杉木凋落叶。C, 观光木凋落叶。D, 台湾相思凋落叶。CT, 对照; +N, 氮添加。

Fig. 2 Dynamics of litter lignin remaining rates under different treatments during decomposition in a subtropical forest (mean ± SD, n = 3). A, Castanopsis carlesii litter. B, Cunninghamia lanceolata litter. C, Michelia odora litter. D, Acacia confusa litter. CT, control; +N, nitrogen addition.

图3 亚热带森林不同处理凋落叶分解过程中纤维素残留率变化(平均值±标准偏差, n = 3)。A, 米槠凋落叶。B, 杉木凋落叶。C, 观光木凋落叶。D, 台湾相思凋落叶。CT, 对照; +N, 氮添加。

Fig. 3 Dynamics of litter cellulose remaining rates under different treatments during decomposition in a subtropical forest (mean ± SD, n = 3). A, Castanopsis carlesii litter. B, Cunninghamia lanceolata litter. C, Michelia odora litter. D, Acacia confusa litter. CT, control; +N, nitrogen addition.

图4 亚热带森林不同处理凋落叶分解过程中氮残留率变化(平均值±标准偏差, n = 3)。A, 米槠凋落叶。B, 杉木凋落叶。C, 观光木凋落叶。D, 台湾相思凋落叶。CT, 对照; +N, 氮添加。

Fig. 4 Dynamics of litter nitrogen remaining rates under different treatments during decomposition in a subtropical forest (mean ± SD, n = 3). A, Castanopsis carlesii litter. B, Cunninghamia lanceolata litter. C, Michelia odora litter. D, Acacia confusa litter. CT, control; +N, nitrogen addition.

图5 亚热带森林不同处理凋落叶分解过程中平均酶活性(平均值±标准偏差, n = 3)。A, 米槠凋落叶; B, 杉木凋落叶; C, 观光木凋落叶; D, 台湾相思凋落叶。CT, 对照; +N, 氮添加。AP, 酸性磷酸酶; βG, β-葡萄糖苷酶; CBH, 纤维素水解酶; NAG, β-N-乙酰氨基葡萄糖苷酶; PEO, 过氧化物酶; PHO, 酚氧化酶。不同小写字母表示同一种酶活性在对照和N添加处理之间差异显著(p < 0.05)。

Fig. 5 Litter enzyme activities under different treatments during decomposition in a subtropical forest (mean ± SD, n = 3). A, Castanopsis carlesii litter; B, Cunninghamia lanceolata litter; C, Michelia odora litter; D, Acacia confusa litter. CT, control; +N, nitrogen addition. AP, acid phosphatase; βG, β-1,4-glucosidase; CBH, cellobiohydrolase; NAG, β-1,4-N-acetylglucosaminidase; PEO, peroxidase; PHO, phenol oxidase. Different lowercase letters indicate significant differences among control and nitrogen addition treatments in the same enzyme (p < 0.05).

表5 亚热带森林凋落物酶活性的双因素方差分析

Table 5 Analysis (indicated by F values from ANOVA with two factors) of litter type, nitrogen addition and their interactions on litter enzyme activity in a subtropical forest

差异来源 Different source	βG	CBH	NAG	AP	PHO	PEO
L	23.755^***	35.348^***	80.072^***	82.374^***	32.250^***	97.990^***
N	27.609^***	0.040	60.288^***	7.380^*	4.684^*	256.223^***
L × N	21.797^***	29.083^***	149.790^***	5.044^*	19.714^***	15.464^***

表6 亚热带森林对照样地凋落物质量损失速率与累积酶活性和初始化学性质的Pearson相关分析(n = 12, 使用平均值)

Table 6 Pearson correlation coefficients (r) for litter mass loss rate versus cumulative enzyme activity and initial chemistry in the control plots after 2-year decomposition in a subtropical forest (n = 12, mean value used)

酶活性 Enzyme activity								初始性质 Initial chemistry
AP	βG	CBH	NAG	PEO	PHO	C	N	可萃取物 Extractive (%)	纤维素 Cellulose (%)	木质素 Lignin (%)
-1.98	0.661^*	0.511	0.372	0.297	0.533	-0.817^**	-0.530	0.871^**	-0.724^**	-0.848^**

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亚热带不同树种凋落叶分解对氮添加的响应

Response of leaf litter decomposition of different tree species to nitrogen addition in a subtropical forest

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