九种维管植物水力性状的演化趋势

doi:10.17521/cjpe.2017.0258

摘要/Abstract

摘要：

维管植物从蕨类植物、裸子植物到被子植物的演化过程中, 生理结构和功能都被认为更趋向于适应干旱的环境。但是关于3类植物是否提升了水分传导和水分利用效率的研究目前多基于叶脉和气孔的形态结构推断, 而对水分传导速率的直接测量则集中于被子植物内部。因此, 同时测定3类植物叶片和枝条尺度的水力性状, 可以研究维管植物水力性状的演化趋势和协同演化关系。该研究测定了中国科学院华南植物园内的蕨类植物(芒萁(Dicranopteris pedata)、华南毛蕨(Cyclosorus parasiticus)、乌毛蕨(Blechnum orientale)), 裸子植物(罗汉松(Podocarpus macrophyllus)、竹柏(Podocarpus nagi)、落羽杉(Taxodium distichum))和被子植物(海南木莲(Manglietia fordiana var. hainanensis)、东京油楠(Sindora tonkinensis)、羊蹄甲(Bauhinia purpurea))共9种植物的水力结构与功能的相关性状。发现9种植物从原始到更演化物种的枝条比导水率(K_S)、叶片比导水率(K_L)和叶片水力导度(K_leaf)依次升高; 叶片蒸腾速率(E)和潜在水分利用效率(WUE_i)升高趋势不显著; 气孔面积指数(SPI)和木材密度(WD)在3个类群间无明显变化趋势。水力功能性状在枝条(K_S)和叶片(K_L、K_leaf和E)尺度显著相关, 但和两个结构性状并无显著相关性。对显著相关的性状进行系统发育独立差分析发现相关性依然存在, 表明枝条导水和叶片导水、叶片失水的协同性独立于系统发育而存在。该文以9种维管植物为例, 发现演化过程中水分传导相关的功能性状有显著提升, 且导水和失水的性状存在着协同演化关系。上述维管植物水力性状演化趋势的结论需要对更多物种、更精细的水力结构性状的进行测定验证。

关键词: 功能性状, 适应性, 水分传导, 系统发育独立差, 协同演化

Abstract:

Aims As vascular plants evolve from ferns to gymnosperms and angiosperms, their physiological structures and functions are assumed more adaptable to arid environment. Whether the three plant groups from early to late evolved lineages have improved their water transport and use efficiency has been studied on the basis of the morphological structure of leaf veins and stomata.Moreover, the water transportation rate was directly measured in the angiosperms. Therefore, we measured structural and functional traits related to water relations in all three plant groups simultaneously, to test the hypothesis on the evolutionary process of plant hydraulics.

Methods We selected three species in each group grown in South China Botanical Garden, Guangzhou, China, including ferns (Dicranopteris pedata, Cyclosorus parasiticus and Blechnum orientale), gymnosperms (Podocarpus macrophyllus, Podocarpus nagi and Taxodium distichum) and angiosperms (Manglietia fordiana var. hainanensis, Sindora tonkinensis and Bauhinia purpurea).

Important findings Sapwood and leaf specific hydraulic conductivities (K_S and K_L, respectively), and leaf conductance (K_leaf) significantly increased from ferns, gymnosperms to angiosperms. However, no significant trends were found in transpiration rate (E) and intrinsic water use efficiency. Meanwhile, neither the size and density of stomata nor wood density showed significant difference among three plant groups. The hydraulic functional traits (K_S, K_Land K_leaf) had significantly positive correlations with each other, but had no relationships with the two measured structural traits. Phylogenetic independent contrasts analyses showed that the coordination between K_S and K_leaf, and between K_S and E were independent of the phylogeny. Based on the nine vascular species, this study demonstrated that water transport related traits are improved as vascular plants evolved, and the co-evolution between water transport and transpiration traits were identified. For further study, it is necessary to consolidate our data with investigations of more detailed water-transport structures in more species from different evolutionary lineages.

Key words: functional traits, adaptation, water transport, phylogenetic independent contrast, correlated evolution

赵乐文, 陈梓熠, 邹滢, 付子钊, 吴桂林, 刘小容, 罗琦, 林忆雪, 李雄炬, 刘智通, 刘慧. 九种维管植物水力性状的演化趋势. 植物生态学报, 2018, 42(2): 220-228. DOI: 10.17521/cjpe.2017.0258

ZHAOLe-Wen, CHEN Zi-Yi, ZOU Ying, FU Zi-Zhao, WU Gui-Lin, LIU Xiao-Rong, LUO Qi, LIN Yi-Xue, LI Xiong-Ju, LIU Zhi-Tong, LIU Hui. Changes in hydraulic traits of nine vascular plants from different evolutionary lineages. Chinese Journal of Plant Ecology, 2018, 42(2): 220-228. DOI: 10.17521/cjpe.2017.0258

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2017.0258

https://www.plant-ecology.com/CN/Y2018/V42/I2/220

图/表 5

表1 本试验中不同演化类群9种维管植物概况

Table 1 Summary of the nine species from three evolutionary lineages in this study

	物种 Species	缩略词 Abbreviation	科 Family
蕨类植物 Ferns	芒萁 Dicranopteris pedata	Dp	里白科 Gleicheniaceae
	华南毛蕨 Cyclosorus parasiticus	Cp	金星蕨科 Thelypteridaceae
	乌毛蕨 Blechnum orientale	Bo	乌毛蕨科 Blechnaceae
裸子植物 Gymnosperms	罗汉松 Podocarpus macrophyllus	Pm	罗汉松科 Podocarpaceae
	竹柏 Podocarpus nagi	Pn	罗汉松科 Podocarpaceae
	落羽杉 Taxodium distichum	Td	杉科 Taxodiaceae
被子植物 Angiosperms	海南木莲 Manglietia fordiana var. hainanensis	Mh	木兰科 Magnoliaceae
	东京油楠 Sindora tonkinensis	St	豆科 Fabaceae
	羊蹄甲 Bauhinia purpurea	Bp	豆科 Fabaceae

图1 不同演化类群9种植物的枝条比导水率(KS)、叶片比导水率(KL)和叶片水力导度(Kleaf)(平均值±标准误差, n = 3-5)。最右图柱上不同字母是HSD多重比较的结果, 9个物种的系统发育树在柱状图最下方, 物种缩写见表1。Fern, 蕨类植物; Gym, 裸子植物; Ang, 被子植物。

Fig. 1 Comparison of sapwood-specific hydraulic conductivity (KS), leaf-specific hydraulic conductivity (KL) and leaf hydraulic conductance (Kleaf) among the nine species from three evolutionary lineages (mean ± SE, n = 3-5). Letters on top of each bar in the right column are HSD multiple comparison results, the phylogenetic tree of the nine species is drawn at the bottom. See species abbrivations in Table 1. Fern, ferns; Gym, gynosperms; Ang, angiosperms.

图2 不同演化类群9种植物的蒸腾速率(E)和潜在水分利用效率(WUEi)(平均值±标准误差, n = 3-5)。缩写注释同图1。

Fig. 2 Comparison of leaf transpiration rate (E) and intrinsic water use efficiency (WUEi) among the nine species from three evolutionary lineages (mean ± SE, n = 3-5). Notes are the same with Fig. 1.

图3 不同演化类群9种植物的气孔面积指数(SPI)和木材密度(WD)(平均值±标准误差, n = 3-5)。缩写注释同图1。

Fig. 3 Comparison of stomatal area index (SPI) and wood density (WD) among the nine species from three evolutionary lineages (mean ± SE, n = 3-5). Notes are the same with Fig. 1.

图4 不同演化类群9种植物的枝条比导水率(KS)与叶片水力导度(Kleaf)(A, B)和蒸腾速率(E)(C, D)的相关性。A、C是这两对性状的直接相关, B、D是这两对性状的系统发育独立差分析。A, C内数值为平均值±标准误差(n = 3-5), A, B, C, D均报告Pearson相关系数(r)和显著性(p)。

Fig. 4 Correlations between sapwood-specific hydraulic conductivity (KS) and leaf hydraulic conductance (Kleaf)(A, B), and leaf transpiration rate (E)(C, D) among the nine species from three evolutionary lineages. A, C, traditional cross-species correlations; B, D, correlations among phylogenetically independent contrasts. In A and C, data are mean ± SE (n = 3-5), Pearson correlation coefficients (r) and p values are reported.

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