不同生境条件下蕨类和被子植物的气孔形态特征及其对光强变化的响应

doi:10.3724/SP.J.1258.2014.00081

植物生态学报 ›› 2014, Vol. 38 ›› Issue (8): 868-877.DOI: 10.3724/SP.J.1258.2014.00081

不同生境条件下蕨类和被子植物的气孔形态特征及其对光强变化的响应

熊慧¹, 马承恩², 李乐³, 曾辉¹^,², 郭大立³^,^*()

¹北京大学深圳研究生院城市人居环境科学与技术重点实验室, 深圳 518055
²北京大学城市与环境学院生态学系, 北京 100871
³中国科学院地理科学与资源研究所, 生态系统网络观测与模拟重点实验室, 千烟洲生态试验站, 北京 100101

收稿日期:2014-03-11 接受日期:2014-05-25 出版日期:2014-03-11 发布日期:2014-08-18
通讯作者: 郭大立
作者简介:* E-mail:guodl@igsnrr.ac.cn
基金资助:
中国科学院百人计划项目(KZZD-EW-TZ-11);国家自然科学基金项目(31325006);国家自然科学基金项目(31021001)

Stomatal characteristics of ferns and angiosperms and their responses to changing light intensity at different habitats

XIONG Hui¹, MA Cheng-En², LI Le³, ZENG Hui¹^,², GUO Da-Li³^,^*()

¹Key Laboratory for Urban Habitat Environmental Science and Technology, Peking University Shenzhen Graduate School, Shenzhen 518055, China
²Department of Ecology, College of Urban and Environmental Sciences, Peking University, Beijing 100871, China
³Qianyanzhou Ecological Station, Key Laboratory of Ecosystem Network Observation and Modeling, Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of Sciences, Beijing 100101, China

Received:2014-03-11 Accepted:2014-05-25 Online:2014-03-11 Published:2014-08-18
Contact: GUO Da-Li

摘要/Abstract

摘要：

气孔是植物与大气环境进行气体交换的重要通道, 在调控植物碳水平衡方面发挥着重要作用。为探讨生境和植物类型对气孔形态特征的影响以及气孔对光强变化的响应格局在不同植物间和不同生境条件下的变异, 选取开阔生境和林下生境的5种蕨类植物和4种被子植物, 测定了它们的气孔形态特征和气孔导度对光强变化的响应。此外, 还收集了8篇文献中开阔和林下生境的45种蕨类植物和70种被子植物的气孔密度和气孔长度数据, 以增大样本量从而更好地探讨不同生境条件下蕨类和被子植物气孔密度及长度的变异格局, 并通过分析生境和植物类型对气孔形态特征的影响来推测生境和植物类型对气孔响应行为的可能影响。实验结果表明, 与林下植物相比, 开阔环境下的植物气孔密度更大, 气孔长度更小, 气孔对光强降低的响应更敏感; 但植物类型对气孔形态特征的影响以及对气孔响应光强的敏感程度的影响均不显著。对文献数据的分析表明, 生境和植物类型对气孔形态特征均有显著影响。考虑到气孔响应快慢与气孔形态特征密切相关, 与蕨类植物相比, 被子植物小而密的气孔可能为其更快地响应环境变化提供了基础。研究表明生境和植物类型对气孔响应行为均有显著影响。

关键词: 蕨类植物, 生境, 光照强度, 气孔密度, 气孔长度, 气孔导度

Abstract:

Aims Stomata are critical in controlling the exchange of water vapour and carbon dioxide and maintaining the balance between plant water and carbon relations. Here, we investigated the effects of habitat (open and understory) and plant type (ferns and angiosperms) on stomatal morphology and stomatal responses to changing light intensity.
Methods We measured stomatal morphology and stomatal conductance in response to transitions in light intensity in five ferns and four angiosperms from different habitats. To increase the sample size, we also collected data on stomatal characteristics for 45 ferns and 70 angiosperms from published studies.
Important findings For all the nine species, the plants in open-habitat had significantly greater stomatal density, shorter stomatal length and greater sensitivity to decreasing light intensity than those in the understory, but the effect of plant type was not significant. Combined analysis with published data indicated that the effects of both habitat and plant type on stomatal morphology were significant. As stomatal sensitivity was closely linked to stomatal morphology, more and smaller stomata might enable angiosperms to respond more quickly to environmental perturbations than ferns. We conclude that both habitat and plant type affect the stomatal response to light.

Key words: fern, habitat, light intensity, stomatal density, stomatal length, stomatal conductance

熊慧, 马承恩, 李乐, 曾辉, 郭大立. 不同生境条件下蕨类和被子植物的气孔形态特征及其对光强变化的响应. 植物生态学报, 2014, 38(8): 868-877. DOI: 10.3724/SP.J.1258.2014.00081

XIONG Hui, MA Cheng-En, LI Le, ZENG Hui, GUO Da-Li. Stomatal characteristics of ferns and angiosperms and their responses to changing light intensity at different habitats. Chinese Journal of Plant Ecology, 2014, 38(8): 868-877. DOI: 10.3724/SP.J.1258.2014.00081

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链接本文: https://www.plant-ecology.com/CN/10.3724/SP.J.1258.2014.00081

https://www.plant-ecology.com/CN/Y2014/V38/I8/868

图/表 9

图1 开阔和林下生境不同土壤深度的土壤含水量(平均值±标准误差, n = 6)。***, p < 0.001。

Fig. 1 Soil water content at different soil depth in open and understory habitats (means ± SE, n = 6). ***, p < 0.001.

表1 9种植物的植物类型、生境、缩写和植株高度(平均值±标准误差)

Table 1 Plant type, habitat, species abbreviations and plant height of the nine species (mean ± SE)

物种 Species	植物类型 Plant type	生境 Habitat	缩写 Abbreviations	高度 Height (m)
芒萁 Dicranopteris pedata	蕨类植物 Fern	开阔 Open	Dp	0.78 ± 0.09
菜蕨 Callipteris esculenta	蕨类植物 Fern	开阔 Open	Ce	1.13 ± 0.09
长叶铁角蕨 Asplenium prolongatum	蕨类植物 Fern	林下 Understory	Ap	0.34 ± 0.01
福建观音座莲 Angiopteris fokiensis	蕨类植物 Fern	林下 Understory	Af	1.85 ± 0.09
狗脊 Woodwardia japonica	蕨类植物 Fern	林下 Understory	Wj	0.94 ± 0.09
少花柏拉木 Blastus pauciflorus	被子植物 Angiosperm	开阔 Open	Bp	1.02 ± 0.09
三叶地锦 Parthenocissus semicordata	被子植物 Angiosperm	林下 Understory	Ps	藤本 Alina
对叶楼梯草 Elatostema sinense	被子植物 Angiosperm	林下 Understory	Es	0.55 ± 0.03
心叶毛蕊茶 Camellia cordifolia	被子植物 Angiosperm	林下 Understory	Cc	3.43 ± 0.54

图2 开阔和林下生境的4种被子植物和5种蕨类植物的气孔密度和气孔长度(平均值±标准误差)。物种缩写同表1; A, F分别代表被子植物(n = 4)和蕨类植物(n = 5); U, O分别代表林下生境(n = 6)和开阔生境(n = 3); **, p < 0.01; *, p < 0.05。

Fig. 2 Stomatal density (SD) and stomatal length (SL) for 4 angiosperms and 5 ferns in open and understory habitats (mean ± SE). Abbreviations of species name see Table 1; A and F represent angiosperms (n = 4) and ferns (n = 5), respectively; U and O represent plants in understory (n = 6) and open habitats (n = 3), respectively; **, p < 0.01; *, p < 0.05.

图3 开阔和林下生境的蕨类和被子植物的气孔密度和气孔长度的关系。数据包括本研究实测数据以及文献中的数据。

Fig. 3 Relationship between stomatal density and stomatal length in ferns and angiosperms in open and understory habitats. Data include measurements in this study and from literature.

图4 生境和类型对植物气孔密度和气孔长度的影响(平均值±标准误差)。数据包括本研究实测数据以及文献中的数据; ***, p < 0.001。

Fig. 4 Effects of habitat and plant type on stomatal density and stomatal length (mean ± SE). Data include measurements in this study and from literature; ***, p < 0.001.

图5 开阔生境被子植物(OA)、开阔生境蕨类植物(OF)、林下生境被子植物(UA)和林下生境蕨类植物(UF)的气孔密度和气孔长度比较(平均值±标准误差)。不同的字母表示差异显著(p < 0.05); 数据包括本研究实测数据以及文献中的数据。

Fig. 5 Comparison of stomatal density and stomatal length among four categories of plants: angiosperm in open habitat (OA), fern in open habitat (OF), angiosperm in understory (UA), and fern in understory (UF) (mean ± SE). Different letters indicate significant difference (p < 0.05); Data include measurements in this study and from literature.

表2 生境和植物类型对9种植物气孔响应光强变化过程中的各个指标的影响。

Table 2 Effects of habitat and plant type on each index in response to changing light intensity of nine species.

变异来源 Source of variable	植物生境 Habitat	植物类型 Plant type	类型×生境 Habitat × Plant type
光合速率最大值(A, μmol·m^-2·s^-1)	0.017	0.165	0.172
气孔导度最大值(G_s, mol·m^-2·s^-1)	0.024	0.169	0.126
气孔关闭绝对速率(G_closure, G_s·s^-1)	0.000	0.116	0.260
气孔关闭相对速率(P_closure, %G_s·s^-1)	0.032	0.241	0.934
气孔张开绝对速率(G_open, G_s·s^-1)	0.493	0.814	0.751
气孔张开相对速率(P_open, %G_s·s^-1)	0.601	0.179	0.985

图6 开阔和林下生境的两种蕨类植物(A-D)和两种被子植物(E-H)的气孔导度和光合速率随光强变化的响应曲线。虚线表示光照强度; A, E的黑色箭头可以看出光强降低时(1000到100 μmol·m-2·s-1)开阔生境的蕨类和被子植物气孔导度迅速降低, 气孔快速响应; C, G的黑色箭头可以看出光强降低时(1000到100 μmol·m-2·s-1)林下生境的蕨类和被子植物气孔导度变化很小, 气孔响应缓慢。

Fig. 6 Responses of stomatal conductance (Gs) and CO2 assimilation rate (A) in two ferns (A-D) and two angiosperms (E-H) in open and understory habitats for 30 minutes after each of four transitions in light intensity (PPFD) (dashed line). The two species in open habitats showed rapid stomatal closure following transition from high to low light (1000 to 100 μmol·m-2·s-1) (A and E, black arrows); whereas the two species in understory showed slow stomatal closure following transition from high to low light (1000 to 100 μmol·m-2·s-1) (C and G, black arrows).

Table 3 Correlations between the eight morphological and physiological traits of nine species

性状指标 Traits	气孔密度 SD	气孔长度 SL	光合速率最大值 A	气孔导度最大值 G_s	气孔关闭绝对速率 G_closure	气孔关闭相对速率 P_closure	气孔张开绝对速率 G_open
气孔长度 SL	-0.588⁺
光合速率最大值 A	0.773^*	-0.363
气孔导度最大值 G_s	0.755^*	-0.371	0.984^**
气孔关闭绝对速率 G_closure	0.911^**	-0.555	0.901^**	0.906^**
气孔关闭相对速率 P_closure	0.661⁺	-0.639⁺	0.325	0.306	0.658⁺
气孔张开绝对速率 G_open	0.090	-0.135	0.648	0.673^*	0.440	-0.063
气孔张开相对速率 P_open	-0.448	-0.102	-0.166	-0.217	-0.291	-0.122	0.414

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不同生境条件下蕨类和被子植物的气孔形态特征及其对光强变化的响应

Stomatal characteristics of ferns and angiosperms and their responses to changing light intensity at different habitats

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