植物生态学报 ›› 2016, Vol. 40 ›› Issue (9): 893-901.DOI: 10.17521/cjpe.2016.0163
所属专题: 青藏高原植物生态学:群落生态学
岳楷1, 杨万勤1,2, 彭艳1, 黄春萍1,3, 张川1, 吴福忠1,2,*()
收稿日期:
2016-05-09
接受日期:
2016-07-23
出版日期:
2016-09-10
发布日期:
2016-09-29
通讯作者:
吴福忠
基金资助:
Kai YUE1, Wan-Qin YANG1,2, Yan PENG1, Chun-Ping HUANG1,3, Chuan ZHANG1, Fu-Zhong WU1,2,*()
Received:
2016-05-09
Accepted:
2016-07-23
Online:
2016-09-10
Published:
2016-09-29
Contact:
Fu-Zhong WU
摘要:
溪流广泛分布于高寒森林地表, 凋落于其中的林木凋落物的分解是整个森林生态系统物质循环的重要环节, 水体流动过程中的冲刷和淋洗作用及其他独特的环境条件可能显著影响凋落物中木质素的降解。该研究采用凋落袋法对比研究了岷江上游高寒森林4种典型且初始质量差异显著的凋落叶, 即康定柳(Salix paraplesia)、高山杜鹃(Rhododendron lapponicum)、方枝柏(Sabina saltuaria)和四川红杉(Larix mastersiana), 在不同生境(林下、溪流和河岸带)下分解过程中木质素残留质量和浓度(质量百分率)的动态变化特征。经过两年的分解, 发现溪流显著促进了凋落叶中木质素的降解; 同一物种凋落叶在不同生境下木质素残留质量差异显著(p < 0.05), 整体表现为溪流<河岸带<林下; 在凋落叶分解的初期木质素有明显的降解, 其浓度表现为先降低后升高, 但不同物种之间存在显著(p < 0.05)的差异; 在整个分解过程中, 木质素残留质量总体呈现出了降低的趋势。此外, 生境类型、分解时期和区域性环境因子(温度、pH值和营养元素的有效性)能显著影响木质素的降解率。这些结果表明, 传统上认为木质素在凋落叶分解初期相对稳定的观点可能并不准确, 其浓度很可能是先下降后升高, 这也与有关木质素动态的最新研究结果相一致。另一方面, 在不同分解时期和不同生境下, 凋落叶木质素降解率表现出了显著差异, 表明区域性环境因子在凋落叶分解和木质素降解过程中具有重要的作用。
岳楷, 杨万勤, 彭艳, 黄春萍, 张川, 吴福忠. 高寒森林溪流对凋落叶分解过程中木质素降解的影响. 植物生态学报, 2016, 40(9): 893-901. DOI: 10.17521/cjpe.2016.0163
Kai YUE, Wan-Qin YANG, Yan PENG, Chun-Ping HUANG, Chuan ZHANG, Fu-Zhong WU. Effects of streams on lignin degradation during foliar litter decomposition in an alpine forest. Chinese Journal of Plant Ecology, 2016, 40(9): 893-901. DOI: 10.17521/cjpe.2016.0163
生境 Habitat | AT (℃) | C (g·kg-1) | N (g·kg-1) | P (g·kg-1) | pH | ||
---|---|---|---|---|---|---|---|
林下 Forest floor | 2.0 ± 5.2 | 126 ± 26 | 5.8 ± 1.1 | 1.2 ± 0.2 | 6.6 ± 0.02 | ||
生境 Habitat | AT (°C) | HCO3- (mg·L-1) | NH4+ (mg·L-1) | NO3-(mg·L-1) | PO43- (μg·L-1) | pH | FV (m·s-1) |
溪流 Stream | 5.1 ± 2.6 | 13.9 ± 1.96 | 0.10 ± 0.05 | 0.29 ± 0.07 | 7.85 ± 0.38 | 6.6 ± 0.4 | 0.53 ± 0.15 |
河岸带 Riparian zone | 4.8 ± 3.4 | 19.7 ± 1.33 | 0.04 ± 0.02 | 0.34 ± 0.08 | 7.84 ± 0.41 | 6.9 ± 0.3 | 0.05 ± 0.01 |
表1 不同生境下凋落叶分解过程中环境因子特征(平均值±标准偏差, n = 90)
Table 1 Characteristics of environmental conditions of different habitats during the process of foliar litter decomposition (mean ± SD, n = 90)
生境 Habitat | AT (℃) | C (g·kg-1) | N (g·kg-1) | P (g·kg-1) | pH | ||
---|---|---|---|---|---|---|---|
林下 Forest floor | 2.0 ± 5.2 | 126 ± 26 | 5.8 ± 1.1 | 1.2 ± 0.2 | 6.6 ± 0.02 | ||
生境 Habitat | AT (°C) | HCO3- (mg·L-1) | NH4+ (mg·L-1) | NO3-(mg·L-1) | PO43- (μg·L-1) | pH | FV (m·s-1) |
溪流 Stream | 5.1 ± 2.6 | 13.9 ± 1.96 | 0.10 ± 0.05 | 0.29 ± 0.07 | 7.85 ± 0.38 | 6.6 ± 0.4 | 0.53 ± 0.15 |
河岸带 Riparian zone | 4.8 ± 3.4 | 19.7 ± 1.33 | 0.04 ± 0.02 | 0.34 ± 0.08 | 7.84 ± 0.41 | 6.9 ± 0.3 | 0.05 ± 0.01 |
物种 Species | C (%) | N (%) | P (%) | 木质素 Lignin (%) | C:N | C:P | N:P | Lignin:N |
---|---|---|---|---|---|---|---|---|
康定柳 S. paraplesia | 34.8 ± 0.9c | 2.64 ± 0.15a | 0.17 ± 0.01a | 24.7 ± 1.3d | 13.2 ± 0.8d | 207 ± 19.7c | 15.7 ± 1.7a | 9.38 ± 0.8c |
高山杜鹃 R. lapponicum | 38.6 ± 1.1b | 0.69 ± 0.10d | 0.10 ± 0.02d | 29.8 ± 0.8b | 57.2 ± 10.2a | 375 ± 53.6a | 6.75 ± 1.5c | 44.3 ± 8.3a |
方枝柏 S. saltuaria | 46.9 ± 1.8a | 1.05 ± 0.06c | 0.15 ± 0.01b | 28.1 ± 0.8c | 45.1 ± 3.9b | 304 ± 12.6b | 6.79 ± 0.7c | 26.9 ± 1.8b |
四川红杉 L. mastersiana | 37.5 ± 0.5b | 1.59 ± 0.11b | 0.12 ± 0.01c | 37.8 ± 1.0a | 23.6 ± 1.8c | 320 ± 24.6b | 13.6 ± 0.8b | 30.1 ± 2.1b |
表2 康定柳、高山杜鹃、方枝柏和四川红杉凋落叶初始质量特征(平均数±标准偏差, n = 9)
Table 2 Initial chemical properties of Salix paraplesia, Rhododendron lapponicum, Sabina saltuaria, and Larix mastersiana foliar litters (mean ± SD, n = 9)
物种 Species | C (%) | N (%) | P (%) | 木质素 Lignin (%) | C:N | C:P | N:P | Lignin:N |
---|---|---|---|---|---|---|---|---|
康定柳 S. paraplesia | 34.8 ± 0.9c | 2.64 ± 0.15a | 0.17 ± 0.01a | 24.7 ± 1.3d | 13.2 ± 0.8d | 207 ± 19.7c | 15.7 ± 1.7a | 9.38 ± 0.8c |
高山杜鹃 R. lapponicum | 38.6 ± 1.1b | 0.69 ± 0.10d | 0.10 ± 0.02d | 29.8 ± 0.8b | 57.2 ± 10.2a | 375 ± 53.6a | 6.75 ± 1.5c | 44.3 ± 8.3a |
方枝柏 S. saltuaria | 46.9 ± 1.8a | 1.05 ± 0.06c | 0.15 ± 0.01b | 28.1 ± 0.8c | 45.1 ± 3.9b | 304 ± 12.6b | 6.79 ± 0.7c | 26.9 ± 1.8b |
四川红杉 L. mastersiana | 37.5 ± 0.5b | 1.59 ± 0.11b | 0.12 ± 0.01c | 37.8 ± 1.0a | 23.6 ± 1.8c | 320 ± 24.6b | 13.6 ± 0.8b | 30.1 ± 2.1b |
图1 不同生境下康定柳(A)、高山杜鹃(B)、方枝柏(C)、四川红杉(D)凋落叶分解过程中木质素残留质量(g)的动态特征(平均值±标准偏差, n = 9)。不同小写字母表示同一分解时期同一物种在不同生境下木质素残留质量差异显著(p < 0.05)。
Fig. 1 Dynamics of lignin mass remaining (g) in the decomposing foliar litter of Salix paraplesia (A), Rhododendron lapponicum (B), Sabina saltuaria (C), and Larix mastersiana (D) under different habitat conditions (mean ± SD, n = 9). Different lowercase letters indicate significant (p < 0.05) differences of lignin mass remaining for a given litter species in a specific decomposition period under different habitat conditions.
图2 不同生境下康定柳(A)、高山杜鹃(B)、方枝柏(C)、四川红杉(D)凋落叶分解过程中木质素浓度(质量百分率, %)的动态特征(平均值±标准偏差, n = 9)。不同小写字母表示同一物种凋落叶在同一生境的不同分解时期木质素浓度差异显著(p < 0.05)。FP, 冻结期; GS, 生长季节; IV, 初始值; LGS, 生长季节后期; PP, 冻结初期; TP, 融化期; 1, 第一年; 2, 第二年。
Fig. 2 Dynamics of lignin concentration (percent litter mass, %) during Salix paraplesia (A), Rhododendron lapponicum (B), Sabina saltuaria (C), and Larix mastersiana (D) foliar litter decomposition (p < 0.05) under different habitat conditions (mean ± SD, n = 9). Different lowercase letters indicate significant (p < 0.05) differences of lignin concentration among different decomposition periods for a given litter species incubated in a specific type of habitat. FP, freezing period; GS, growing season; IV, initial value; LGS, late growing season; PP, pre-freezing period; TP, thawing period; 1, first year; 2, second year.
图3 不同生境下康定柳(A)、高山杜鹃(B)、方枝柏(C)、四川红杉(D)凋落叶分解过程中木质素降解率(%/月)的动态特征(平均值±标准偏差, n = 9)。不同小写字母表示同一物种凋落叶在同一生境下不同分解时期木质素降解率差异显著(p < 0.05)。FP, 冻结期; IV, 初始值; GS, 生长季节; LGS, 生长季节后期; PP, 冻结初期; TP, 融化期; 1, 第一年; 2, 第二年。
Fig. 3 Dynamics of lignin degradation rate (%/month) during Salix paraplesia (A), Rhododendron lapponicum (B), Sabina saltuaria (C), and Larix mastersiana (D) foliar litter decomposition (p < 0.05) under different habitat conditions (mean ± SD, n = 9). Different lowercase letters indicate significant (p < 0.05) differences of lignin degradation rate among different decomposition periods for a given litter species incubated in a specific type of habitat. FP, freezing period; GS, growing season; IV, initial value; LGS, late growing season; PP, pre-freezing period; TP, thawing period; 1, first year; 2, second year.
影响因子 Influence factor | 自由度 Degree of freedom | F | p |
---|---|---|---|
物种 Species | 3 | 165.753 | < 0.001 |
生境 Habitat | 2 | 75.197 | < 0.001 |
时期 Period | 9 | 504.141 | < 0.001 |
物种×生境 Species × habitat | 6 | 40.353 | < 0.001 |
物种×时期 Species × period | 27 | 17.003 | < 0.001 |
生境×时期 Habitat × period | 18 | 18.317 | < 0.001 |
物种×生境×时期 Species × habitat × period | 54 | 12.020 | < 0.001 |
表3 凋落叶分解过程中木质素降解率与物种、生境和分解时期的重复测量方差分析
Table 3 Repeated-measure ANOVA analysis on the effects of litter species, habitat type, and decomposition period on lignin degradation rate during litter decomposition process
影响因子 Influence factor | 自由度 Degree of freedom | F | p |
---|---|---|---|
物种 Species | 3 | 165.753 | < 0.001 |
生境 Habitat | 2 | 75.197 | < 0.001 |
时期 Period | 9 | 504.141 | < 0.001 |
物种×生境 Species × habitat | 6 | 40.353 | < 0.001 |
物种×时期 Species × period | 27 | 17.003 | < 0.001 |
生境×时期 Habitat × period | 18 | 18.317 | < 0.001 |
物种×生境×时期 Species × habitat × period | 54 | 12.020 | < 0.001 |
生境 Habitat | 回归式 Regression model | ||||
---|---|---|---|---|---|
a0 | a1X1 | a2X2 | a3X3 | a4X4 | |
林下 Forest floor | ŷ = 0.424 | -0.017 C:N (0.353) | +0.042 C (0.607) | ||
溪流 Stream | ŷ = 1.602 | -0.031 Lignin:N (0.785) | +13.231 P (0.874) | ||
河岸带 Riparian zone | ŷ = -7.311 | +0.032 Lignin (0.783) | +20.108 P (0.888) | +0.164 N:P (0.932) | +0.090 C (0.940) |
表4 凋落叶木质素两年总降解率(%/月)与初始质量的逐步回归分析
Table 4 Stepwise regression analysis between lignin degradation rate (%/month) of the 2 years and foliar litter initial chemical properties
生境 Habitat | 回归式 Regression model | ||||
---|---|---|---|---|---|
a0 | a1X1 | a2X2 | a3X3 | a4X4 | |
林下 Forest floor | ŷ = 0.424 | -0.017 C:N (0.353) | +0.042 C (0.607) | ||
溪流 Stream | ŷ = 1.602 | -0.031 Lignin:N (0.785) | +13.231 P (0.874) | ||
河岸带 Riparian zone | ŷ = -7.311 | +0.032 Lignin (0.783) | +20.108 P (0.888) | +0.164 N:P (0.932) | +0.090 C (0.940) |
林下 Forest floor | AT | C | N | P | pH | ||
---|---|---|---|---|---|---|---|
康定柳 Salix paraplesia | 26.925*** | 35.094*** | 1.987 | 0.340 | 6.194* | ||
高山杜鹃 Rhododendron lapponicum | 16.022*** | 0.064 | 5.700* | 17.816*** | 2.431 | ||
方枝柏 Sabina saltuaria | 10.134** | 1.037 | 23.348*** | 23.681*** | 8.314** | ||
四川红杉 Larix mastersiana | 30.336*** | 7.748** | 32.560*** | 10.076** | 13.489*** | ||
溪流 Stream | AT | HCO3- | NH4+ | NO3- | PO43- | pH | FV |
康定柳 Salix paraplesia | 0.001 | 0.572 | 2.692 | 13.248*** | 0.522 | 6.208* | 0.385 |
高山杜鹃 Rhododendron lapponicum | 1.286 | 1.722 | 6.088* | 8.832** | 1.612 | 1.652 | 1.590 |
方枝柏 Sabinasaltuaria | 1.245 | 4.809* | 7.579** | 8.964** | 0.001 | 6.454* | 0.103 |
四川红杉 Larix mastersiana | 2.815 | 2.179 | 4.681* | 11.866** | 0.063 | 5.594* | 0.053 |
河岸带 Riparian zone | AT | HCO3- | NH4+ | NO3- | PO43- | pH | FV |
康定柳 Salix paraplesia | 35.148*** | 5.748* | 12.267** | 0.256 | 1.305 | 16.431*** | 2.540 |
高山杜鹃 Rhododendron lapponicum | 3.702 | 2.822 | 2.029 | 1.369 | 0.001 | 7.300** | 5.752* |
方枝柏 Sabina saltuaria | 1.564 | 4.024* | 6.775* | 4.609* | 0.115 | 3.545 | 6.232* |
四川红杉 Larix mastersiana | 36.978*** | 15.189*** | 19.985*** | 4.305* | 0.055 | 0.371 | 11.602** |
表5 不同生境下凋落叶分解过程中木质素降解率(%/月)与环境因子回归分析的F值
Table 5 F-value for the regression analysis between lignin degradation rate (%/month) and environmental factors under different habitats during foliar litter decomposition
林下 Forest floor | AT | C | N | P | pH | ||
---|---|---|---|---|---|---|---|
康定柳 Salix paraplesia | 26.925*** | 35.094*** | 1.987 | 0.340 | 6.194* | ||
高山杜鹃 Rhododendron lapponicum | 16.022*** | 0.064 | 5.700* | 17.816*** | 2.431 | ||
方枝柏 Sabina saltuaria | 10.134** | 1.037 | 23.348*** | 23.681*** | 8.314** | ||
四川红杉 Larix mastersiana | 30.336*** | 7.748** | 32.560*** | 10.076** | 13.489*** | ||
溪流 Stream | AT | HCO3- | NH4+ | NO3- | PO43- | pH | FV |
康定柳 Salix paraplesia | 0.001 | 0.572 | 2.692 | 13.248*** | 0.522 | 6.208* | 0.385 |
高山杜鹃 Rhododendron lapponicum | 1.286 | 1.722 | 6.088* | 8.832** | 1.612 | 1.652 | 1.590 |
方枝柏 Sabinasaltuaria | 1.245 | 4.809* | 7.579** | 8.964** | 0.001 | 6.454* | 0.103 |
四川红杉 Larix mastersiana | 2.815 | 2.179 | 4.681* | 11.866** | 0.063 | 5.594* | 0.053 |
河岸带 Riparian zone | AT | HCO3- | NH4+ | NO3- | PO43- | pH | FV |
康定柳 Salix paraplesia | 35.148*** | 5.748* | 12.267** | 0.256 | 1.305 | 16.431*** | 2.540 |
高山杜鹃 Rhododendron lapponicum | 3.702 | 2.822 | 2.029 | 1.369 | 0.001 | 7.300** | 5.752* |
方枝柏 Sabina saltuaria | 1.564 | 4.024* | 6.775* | 4.609* | 0.115 | 3.545 | 6.232* |
四川红杉 Larix mastersiana | 36.978*** | 15.189*** | 19.985*** | 4.305* | 0.055 | 0.371 | 11.602** |
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