浙江天童常绿阔叶林不同演替阶段木本植物的水力结构特征

doi:10.17521/cjpe.2015.0258

摘要/Abstract

摘要：

水力结构是植物应对环境形成的与水分运输相关的形态策略.探索不同演替阶段和群落不同高度层植物的水力结构特征, 有助于理解植物的水分运输和利用策略.该研究以浙江天童常绿阔叶林演替前中后期群落的上层木(占据林冠层的树种)和下层木(灌木层物种)为对象, 测定了演替共有种(至少存在于两个演替阶段的物种)和更替种(仅存在于某一演替阶段的物种)的枝边材比导率,叶比导率和胡伯尔值, 以及边材疏导面积,末端枝总叶面积和枝条水势, 分析植物水力结构在群落上层木和下层木间以及在演替阶段间的差异, 及其与枝叶性状的相关关系.结果显示: (1)上层木植物边材比导率和叶比导率显著高于下层木植物(p < 0.05); (2)上层木和下层木的边材比导率与叶比导率在演替阶段间均无显著差异(p > 0.05); 上层木的胡伯尔值在演替阶段间无显著差异, 下层木的胡伯尔值随演替显著下降(p < 0.05); (3)上层木共有种仅边材比导率随演替进行显著降低(p < 0.05), 更替种的3个水力结构参数在演替阶段间无显著差异; 下层木共有种水力结构参数在演替阶段间无明显差异, 更替种仅胡伯尔值随演替减小(p < 0.05); (4)植物边材比导率与枝疏导面积和末端枝所支撑的总叶面积显著正相关(p < 0.01), 胡伯尔值与枝条水势及末端枝总叶面积显著负相关(p < 0.01).以上结果表明: 天童常绿阔叶林演替各阶段上层木比下层木具有更大的输水能力和效率; 随着演替进行, 上层木与下层木的共有种和更替种边材比导率的相反变化表明上层木水力结构的变化可能由微生境变化引起, 而下层木水力结构特征的变化可能由物种更替造成.

关键词: 水力结构, 共有种, 上层木, 更替种, 下层木

Abstract:

Aims Hydraulic architecture is a morphological strategy in plants to transport water in coping with environmental conditions. Change of hydraulic architecture for plants occupying different canopy layers within community and for the same plant at different successional stages reflect existence and adaptation in plant's water transportation strategies. The objective of this study was to examine how hydraulic architecture varies with canopy layers within a community and with forest succession.Methods The study site is located in Tiantong National Forest Park, Zhejiang Province, China. Hydraulic architectural traits studied include sapwood-specific hydraulic conductivity, leaf-specific hydraulic conductivity, Huber value, sapwood channel area of twigs, total leaf area per terminal twig, and water potential of twigs. We measured those traits for species that occur in multiple successional stages (we called it "overlapping species") and for species that occur only in one successional stage (we called it "turnover species") along a successional series of evergreen broadleaved forests. For a given species, we sampled both overstory and understory trees. Hydraulic architectural traits between overstory and understory trees in the same community and at successional stages were compared. Pearson correlation was used to exam the relationship between hydraulic architectural traits and the twig/leaf traits.Important findings Sapwood-specific hydraulic conductivities and leaf-specific hydraulic conductivities were significantly higher in overstory trees than those in understory trees, but did not significantly differ from successional stages. Huber value decreased significantly for understory trees, but did not change for overstory trees through forest successional stages. For overstory trees, a trend of decreasing sapwood-specific hydraulic conductivity was observed for overlapping species but not for turnover species with successional stages. In contrast, for understory trees, a trend of decreasing Huber values was observed for turner species but not for overlapping species with successional stages. Across tree species, sapwood-specific hydraulic conductivity was positively correlated with sapwood channel area and total leaf area per terminal twig size. Huber value was negatively correlated to water potential of twigs and total leaf area per terminal twig size. These results suggest that water transportation capacity and efficiency are higher in overstory trees than in understory trees across successional stages in evergreen broadleaved forests in Tiantong region. The contrasting trends of sapwood-specific hydraulic conductivity between overlapping species and turnover species indicate that shift of microenvironment conditions might lead to changes of hydraulic architecture in overstory trees, whereas species replacement might result in changes of hydraulic architecture in understory trees.

Key words: hydraulic architecture, overlapping species, overstory trees, turnover species, understory trees

赵延涛, 许洺山, 张志浩, 周刘丽, 张晴晴, 宋彦君, 阎恩荣. 浙江天童常绿阔叶林不同演替阶段木本植物的水力结构特征. 植物生态学报, 2016, 40(2): 116-126. DOI: 10.17521/cjpe.2015.0258

Yan-Tao ZHAO, Ming-Shan XU, Zhi-Hao ZHANG, Liu-Li ZHOU, Qing-Qing ZHANG, ARSHAD Ali, Yan-Jun SONG, En-Rong YAN. Hydraulic architecture of evergreen broad-leaved woody plants at different successional stages in Tiantong National Forest Park, Zhejiang Province, China. Chinese Journal of Plant Ecology, 2016, 40(2): 116-126. DOI: 10.17521/cjpe.2015.0258

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2015.0258

https://www.plant-ecology.com/CN/Y2016/V40/I2/116

图/表 5

表1 浙江天童常绿阔叶林次生林演替系列各阶段样地特征和所选物种

Table 1 Characteristics of study plots in a secondary successional series of evergreen broad-leaved forests in Tiantong, Zhejiang Province

演替阶段 Successional stage	群落类型 Community type	年龄 Age (a)	优势种 Dominant species	上层木 Overstory tree			下层木 Understory tree
演替阶段 Successional stage	群落类型 Community type	年龄 Age (a)	优势种 Dominant species	层高 Height (m)	共有种Overlapping species	更替种 Turnover species	层高Height (m)	共有种Overlapping species	更替种 Turnover species
S1	檵木灌丛 Loropetalum chinese shrub	20	檵木 Loropetalum chinese 赤楠 Syzygium buxifolium 白栎 Quercus fabri	8	木荷 Schima superba	白栎 Quercus fabri	2.5	山矾 Symplocos sumuntia 窄基红褐柃 Eurya Rubiginosa	赤楠 Syzygium buxifolium
S2	木荷群落 Schima superba community	60	木荷 Schima superba 石栎 Lithocarpus glabra	17	木荷 Schima superba	石栎 Lithocarpus glabra	4.0		马银花 Rhododendron ovatum
S3	栲群落 Castanopsis fargesii community	120	栲 Castanopsis fargesii	22		栲 Castanopsis fargesii	5.5		交让木 Daphniphyllum macropodum

图1 浙江天童常绿阔叶林演替系列上层木和下层木的水力结构(平均值±标准误差).All表示3个演替阶段合并.S1,S2和S3表示演替阶段.不同大写字母表示同一演替阶段不同高度层之间差异显著(p < 0.05); 不同小写字母表示同一高度层不同演替阶段之间差异显著(p < 0.05).

Fig. 1 Hydraulic architecture of trees between overstory and understory species along a successional series in evergreen broad-leaved forest in Tiantong, Zhejiang (mean ± SE). All means were combined for three successional stages. S1, S2 and S3 represent successional stages. Different capital letters indicate significant difference between canopy layers at the same successional stage (p < 0.05). Different lowercase letters indicate significant difference between successional stages for the same canopy layer (p < 0.05).

表2 高度层与演替阶段及其交互作用对植物水力结构的双因素方差分析结果

Table 2 Results of two-way ANOVA for testing the main effects of canopy layers, succession stages and their interactions on hydraulic architecture of tree species

变量 Variable	高度层 Vertical layer			演替阶段 Succession stage			高度层×演替阶段 Vertical layer × succession stage
变量 Variable	df	F	p	df	F	p	df	F	p
边材比导率 Sapwood-specific hydraulic conductivity	1	43.62	<0.001	2	2.44	0.10	2	1.36	0.27
叶比导率 Leaf-specific hydraulic conductivity	1	5.18	<0.050	2	2.86	0.07	2	0.32	0.73
胡伯尔值 Huber value	1	4.65	<0.050	2	2.92	0.07	2	1.14	0.33

图2 浙江天童常绿阔叶林演替系列上共有种和更替种的水力结构差异(平均值±标准误差).A, 上层木共有种.B, 下层木共有种.C, 上层木更替种.D, 下层木更替种.S1,S2和S3表示演替阶段.不同演替阶段不同小写字母表示差异显著(p < 0.05), F和p表示单因素方差分析结果.

Fig. 2 Differences of hydraulic architecture for over lapping and turnover species in both overstory and understory layers along a successional series in evergreen broad-leaved forest in Tiantong, Zhejiang (mean ± SE). A, Overstory over lapping species. B, Understory over lapping species. C, Overstory turnover species. D, Understory turnover species. S1, S2 and S3 represent successional stages. Different letters among successional stages indicate significant difference (p < 0.05). F and p are results of one-way ANOVA.

表3 浙江天童常绿阔叶林植物水力结构与枝叶性状的Pearson相关系数

Table 3 Coefficients of Pearson correlation between plant hydraulic architecture and branch and leaf traits in evergreen broad-leaved forests in Tiantong, Zhejiang

	边材比导率 Sapwood-specific hydraulic conductivity	叶比导率 Leaf-specific hydraulic conductivity	胡伯尔值 Huber value	枝条输导面积 Branches channel area	末端枝条总叶面积 Terminal leaf area
叶比导率 Leaf-specific hydraulic conductivity	0.50^***
胡伯尔值 Huber value	-0.12	0.55^***
枝条输导面积 Branches channel area	0.38^**	0.02	-0.17
末端枝条总叶面积 Terminal leaf area	0.33^**	-0.09	-0.36^**	0.75^***
枝条水势 Branches water potential	0.13	-0.10	-0.37^**	0.56^***	0.50^***

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