常绿和落叶阔叶物种当年生小枝茎长度和茎纤细率对展叶效率的影响

doi:10.17521/cjpe.2016.0376

摘要/Abstract

摘要：

枝和叶是组成树冠的两个最主要的结构单元, 枝条的结构特征及其与叶性状之间的变化关系决定了植物对光照、空间等资源的利用和适应策略。该研究以小枝茎长度和茎纤细率表征茎结构, 以叶面积比(即单位小枝茎干质量的总叶面积)、叶密度(即单位茎长度的叶数量)和叶茎生物量比(即单位小枝茎干质量的总叶干质量)表征展叶效率, 以浙江省清凉峰自然保护区25个常绿阔叶物种和60个落叶阔叶物种为对象, 探讨小枝茎结构变化对展叶效率的影响。结果显示, 展叶效率无论是用叶面积比、叶密度还是用叶茎生物量比表示, 常绿和落叶物种小枝的茎长结构性状均与展叶效率呈显著的负相关关系, 即随着小枝茎长度和茎纤细率的增加, 展叶效率逐渐降低, 可能反映了机械安全和光照限制的作用。同时, 尽管茎长度-展叶效率的斜率在常绿和落叶物种间无显著差异, 并且常绿物种叶密度与小枝茎结构性状之间的关系截距显著大于落叶物种, 即常绿物种具有较高的出叶强度。但由于单叶面积和比叶面积显著小于落叶物种, 在某一给定的小枝茎长结构下, 常绿物种每单位质量的茎生物量投资获得的总叶面积和总叶质量都比落叶物种少, 即常绿物种的展叶效率比落叶物种低。这可能反映了常绿物种高消耗慢收益的保守型策略。这些结果表明, 小枝茎长结构对展叶效率具有显著的影响, 并将随叶习性的不同而改变, 与植物对环境的适应策略密切相关。

关键词: 茎长度, 茎纤细率, 展叶效率, 常绿物种, 落叶物种

Abstract:

Aims Branches and leaves are the two main structural units of tree crown composition. Among the adaptive strategies of plants, the functional traits of branches and the relationships between branch traits and leaf traits determine the capacity of trees to access light and space. In this study, our objective is to test the hypothesis that leaf display efficiency is affected by the stem length to stem slender ratio within current-year twigs.Methods The stem length to stem slender ratios of current-year twigs were used as the proxy of stem structure traits. Leaf area ratio (total leaf area per stem mass), leaf density (leaf number per stem length) and leaf/stem mass ratio (total leaf mass per stem mass) were used as the proxies of leaf display efficiency. The relationship between stem structure traits and leaf display efficiency within current-year twigs were studied for 25 evergreen and 60 deciduous broadleaved woody species in Qingliang Mountain, Zhejiang, China. The standardized major axis estimation method was used to examine the scaling relationship between stem structural traits and leaf display efficiency within current-year twigs.Important findings The proxies of leaf display efficiency, measured by leaf area ratio, leaf density or leaf/stem mass ratio, were all significantly and negative correlated with stem length to stem slender ratio within current-year twigs in both evergreen and deciduous broadleaved woody species. This suggested that leaf display efficiency decreased with stem length to stem slender ratios within current-year twigs, which may reflect the role of mechanical safety and light within twigs. The slope of the relationship between leaf display efficiency and stem long-dimension structure traits in evergreen species was not significantly different from the one in deciduous species. In contrast, the y-intercept of the relationship between leaf density and stem long-dimension structure traits was significantly larger in evergreen species than in deciduous species, i.e. the leafing intensity of evergreen species was higher than that of deciduous species. Individual leaf area and specific leaf area were smaller in evergreen species than in deciduous species, which resulted in deciduous species have a larger leaf area per stem mass and leaf mass per stem mass at a given stem length to stem slender ratio compared to evergreen species. It may reflect the conservative adaptive strategy of high consumption and slow benefit in evergreen species. Our results demonstrated that leaf display efficiency could be affected by stem length, and would change with leaf life-span (deciduous versus evergreen).

Key words: stem length, stem slender ratio, leaf display efficiency, evergreen species, deciduous species

李俊慧, 彭国全, 杨冬梅. 常绿和落叶阔叶物种当年生小枝茎长度和茎纤细率对展叶效率的影响. 植物生态学报, 2017, 41(6): 650-660. DOI: 10.17521/cjpe.2016.0376

Jun-Hui LI, Guo-Quan PENG, Dong-Mei YANG. Effect of stem length to stem slender ratio of current-year twigs on the leaf display efficiency in evergreen and deciduous broadleaved trees. Chinese Journal of Plant Ecology, 2017, 41(6): 650-660. DOI: 10.17521/cjpe.2016.0376

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https://www.plant-ecology.com/CN/Y2017/V41/I6/650

图/表 6

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指标 (y 轴-x 轴) Index (y axis-x axis)	功能组 Functional group	样本量 Sample size	决定系数 Coefficient of determination	斜率(95%置信区间) Slope (95% confidence interval)	截距 Intercept
叶数量-小枝茎长度 Leaf number-stem length	落叶 Deciduous	60	0.192	0.510 (0.404, 0.645)	0.238
叶数量-小枝茎长度 Leaf number-stem length	常绿 Evergreen	25	0.279	0.463 (0.323, 0.662)	0.415
叶面积比-小枝茎纤细率 Leaf area ratio-stem slender ratio	落叶 Deciduous	60	0.613	-0.867 (-1.021, -0.737)	4.525
叶面积比-小枝茎纤细率 Leaf area ratio-stem slender ratio	常绿 Evergreen	25	0.620	-0.791 (-1.028, -0.608)	4.175
叶密度-小枝茎纤细率 Density of leaf number-stem slender ratio	落叶 Deciduous	60	0.617	-0.804 (-0.945, -0.684)	1.115
叶密度-小枝茎纤细率 Density of leaf number-stem slender ratio	常绿 Evergreen	25	0.781	-0.717 (-0.876, -0.587)	1.106
叶茎质量比-小枝茎纤细率 Leaf/stem mass ratio-stem slender ratio	落叶 Deciduous	60	0.629	-0.869 (-1.019, -0.741)	2.231
叶茎质量比-小枝茎纤细率 Leaf/stem mass ratio-stem slender ratio	常绿 Evergreen	25	0.823	-0.613 (-0.734, -0.512)	1.859
叶面积比-小枝茎长度 Leaf area ratio-stem length	落叶 Deciduous	60	0.628	-0.973 (-1.141, -0.829)	3.955
叶面积比-小枝茎长度 Leaf area ratio-stem length	常绿 Evergreen	25	0.683	-0.939 (-1.195, -0.738)	3.679
叶密度-小枝茎长度 Density of leaf number-stem length	落叶 Deciduous	60	0.741	-0.902 (-1.030, -0.789)	0.587
叶密度-小枝茎长度 Density of leaf number-stem length	常绿 Evergreen	25	0.787	-0.852 (-1.038, -0.699)	0.657
叶茎质量比-小枝茎长度 Leaf/stem mass ratio-stem length	落叶 Deciduous	60	0.614	-0.974 (-1.146, -0.828)	1.660
叶茎质量比-小枝茎长度 Leaf/stem mass ratio-stem length	常绿 Evergreen	25	0.852	-0.729 (-0.859, -0.618)	1.474
总叶面积-茎干质量 Total leaf area-stem dry mass	落叶 Deciduous	60	0.437	0.815 (0.670, 0.991)	0.504
总叶面积-茎干质量 Total leaf area-stem dry mass	常绿 Evergreen	25	0.439	0.722 (0.526, 0.992)	0.502

指标 (y 轴-x 轴) Index (y axis-x axis)	功能组 Functional group	样本量 Sample size	决定系数 Coefficient of determination	斜率(95%置信区间) Slope (95% confidence interval)	截距 Intercept
叶数量-小枝茎长度 Leaf number-stem length	落叶 Deciduous	60	0.192	0.510 (0.404, 0.645)	0.238
叶数量-小枝茎长度 Leaf number-stem length	常绿 Evergreen	25	0.279	0.463 (0.323, 0.662)	0.415
叶面积比-小枝茎纤细率 Leaf area ratio-stem slender ratio	落叶 Deciduous	60	0.613	-0.867 (-1.021, -0.737)	4.525
叶面积比-小枝茎纤细率 Leaf area ratio-stem slender ratio	常绿 Evergreen	25	0.620	-0.791 (-1.028, -0.608)	4.175
叶密度-小枝茎纤细率 Density of leaf number-stem slender ratio	落叶 Deciduous	60	0.617	-0.804 (-0.945, -0.684)	1.115
叶密度-小枝茎纤细率 Density of leaf number-stem slender ratio	常绿 Evergreen	25	0.781	-0.717 (-0.876, -0.587)	1.106
叶茎质量比-小枝茎纤细率 Leaf/stem mass ratio-stem slender ratio	落叶 Deciduous	60	0.629	-0.869 (-1.019, -0.741)	2.231
叶茎质量比-小枝茎纤细率 Leaf/stem mass ratio-stem slender ratio	常绿 Evergreen	25	0.823	-0.613 (-0.734, -0.512)	1.859
叶面积比-小枝茎长度 Leaf area ratio-stem length	落叶 Deciduous	60	0.628	-0.973 (-1.141, -0.829)	3.955
叶面积比-小枝茎长度 Leaf area ratio-stem length	常绿 Evergreen	25	0.683	-0.939 (-1.195, -0.738)	3.679
叶密度-小枝茎长度 Density of leaf number-stem length	落叶 Deciduous	60	0.741	-0.902 (-1.030, -0.789)	0.587
叶密度-小枝茎长度 Density of leaf number-stem length	常绿 Evergreen	25	0.787	-0.852 (-1.038, -0.699)	0.657
叶茎质量比-小枝茎长度 Leaf/stem mass ratio-stem length	落叶 Deciduous	60	0.614	-0.974 (-1.146, -0.828)	1.660
叶茎质量比-小枝茎长度 Leaf/stem mass ratio-stem length	常绿 Evergreen	25	0.852	-0.729 (-0.859, -0.618)	1.474
总叶面积-茎干质量 Total leaf area-stem dry mass	落叶 Deciduous	60	0.437	0.815 (0.670, 0.991)	0.504
总叶面积-茎干质量 Total leaf area-stem dry mass	常绿 Evergreen	25	0.439	0.722 (0.526, 0.992)	0.502

性状 Trait	常绿 Evergreen	落叶 Deciduous	显著水平 Significant value, p
茎纤细率 Stem slender ratio (cm·cm^-1)	41.80 ± 3.98	46.87 ± 2.80	0.318
茎直径 Stem diameter (cm)	0.18 ± 0.01	0.18 ± 0.01	0.887
茎长度 Stem length (cm)	6.77 ± 0.62	8.01 ± 0.50	0.158
茎干质量 Stem mass (mg)	112.03 ± 11.72	132.50 ± 13.22	0.352
总叶干质量 Total leaf mass (mg)	853.32 ± 74.69	840.87 ± 62.55	0.909
叶密度指数 Leaf density index (No.·cm^-1)	1.20 ± 0.20	0.76 ± 0.06	0.007
叶面积比 Leaf area ratio (cm²·g^-1)	1β081.71 ± 146.01	1β573.41 ± 130.10	0.030
单叶面积 Individual leaf area (cm²)	16.21 ± 1.97	34.66 ± 2.34	<0.001
总叶面积 Total leaf area (cm²)	95.19 ± 10.34	163.68 ± 11.69	0.001
比叶面积 Specific leaf area (cm²·g^-1)	114.31 ± 6.89	200.76 ± 4.42	<0.001
叶茎质量比 Leaf/stem mass ratio (mg·mg^-1)	9.14 ± 0.99	7.96 ± 0.66	0.335

性状 Trait	常绿 Evergreen	落叶 Deciduous	显著水平 Significant value, p
茎纤细率 Stem slender ratio (cm·cm^-1)	41.80 ± 3.98	46.87 ± 2.80	0.318
茎直径 Stem diameter (cm)	0.18 ± 0.01	0.18 ± 0.01	0.887
茎长度 Stem length (cm)	6.77 ± 0.62	8.01 ± 0.50	0.158
茎干质量 Stem mass (mg)	112.03 ± 11.72	132.50 ± 13.22	0.352
总叶干质量 Total leaf mass (mg)	853.32 ± 74.69	840.87 ± 62.55	0.909
叶密度指数 Leaf density index (No.·cm^-1)	1.20 ± 0.20	0.76 ± 0.06	0.007
叶面积比 Leaf area ratio (cm²·g^-1)	1β081.71 ± 146.01	1β573.41 ± 130.10	0.030
单叶面积 Individual leaf area (cm²)	16.21 ± 1.97	34.66 ± 2.34	<0.001
总叶面积 Total leaf area (cm²)	95.19 ± 10.34	163.68 ± 11.69	0.001
比叶面积 Specific leaf area (cm²·g^-1)	114.31 ± 6.89	200.76 ± 4.42	<0.001
叶茎质量比 Leaf/stem mass ratio (mg·mg^-1)	9.14 ± 0.99	7.96 ± 0.66	0.335