亚热带森林附生地衣压力-体积曲线分析及其适用性

doi:10.17521/cjpe.2020.0344

植物生态学报 ›› 2021, Vol. 45 ›› Issue (3): 274-285.DOI: 10.17521/cjpe.2020.0344

亚热带森林附生地衣压力-体积曲线分析及其适用性

董琳琳¹, 普晓妍², 张璐璐¹^,^*(), 宋亮², 鲁志云²^,³, 李苏²^,^*()

¹山东师范大学环境与生态研究院, 济南 250358
²中国科学院西双版纳热带植物园热带森林生态学重点实验室, 昆明 650223
³中国科学院哀牢山亚热带森林生态系统研究站, 云南景东 676209

收稿日期:2020-10-20 接受日期:2021-01-02 出版日期:2021-03-20 发布日期:2021-05-17
通讯作者: 张璐璐,李苏
作者简介:Zhang LL: 675359138@qq.com)
* 李苏: ORCID:0000-0001-5295-6846 (Li S: lis@xtbg.ac.cn;
基金资助:
国家自然科学基金(31770494);国家自然科学基金(3175001);中国科学院青年创新促进会项目(2017441);中国科学院“西部之光”项目和中国科学院135项目(2017XTBG-T01)

Pressure-volume curve analysis of epiphytic lichens and its applicability in subtropical forests

DONG Lin-Lin¹, PU Xiao-Yan², ZHANG Lu-Lu¹^,^*(), SONG Liang², LU Zhi-Yun²^,³, LI Su²^,^*()

¹Institute of Environment and Ecology, Shandong Normal University, Jinan 250358, China
²CAS Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Kunming 650223, China
³Ailaoshan Station for Subtropical Forest Ecosystem Studies, Chinese Academy of Sciences, Jingdong, Yunnan 676209, China

Received:2020-10-20 Accepted:2021-01-02 Online:2021-03-20 Published:2021-05-17
Contact: ZHANG Lu-Lu,LI Su
Supported by:
National Natural Science Foundation of China(31770494);National Natural Science Foundation of China(3175001);Youth Innovation Promotion Association of the Chinese Academy of Sciences (CAS)(2017441);CAS “Light of West China” Program, and the CAS 135 Program(2017XTBG-T01)

摘要/Abstract

摘要：

附生地衣是森林生态系统中的重要组成部分, 在维护森林物种多样性以及水分和养分循环等方面发挥着重要作用。地衣类群的水势特征以及压力-体积(PV)曲线及相关参数是否适用于探索地衣等变水性生物类群应对水分胁迫机制, 仍有待阐明。该研究以中国西南地区云南哀牢山亚热带山地森林中5个类群15种常见附生地衣为对象, 探讨了其PV曲线及水势参数的功能群和物种水平的变化。结果显示, 蓝藻型地衣的内部含水量(WC_internal)和共质体水含量(Rs)显著高于绿藻型地衣, 是其1倍以上, 但其他参数表现出较高的一致性。功能群水平上, 地衣仅WC_internal、Rs和膨压损失点的相对含水量(RWC_TLP)具有显著性差异; 但物种间差异均非常显著。结合主成分分析进一步发现, PV曲线及相关参数在评估地衣整体应对水分胁迫以及揭示生境水分条件选择策略时具有较大的限制性, 受到共生藻和生长型的显著影响; 但内部最大持水力可以解释蓝藻型地衣的生境适应策略, 而部分参数如饱和渗透势(Ψsat)和RWC_TLP可以分别用于解释狭叶地衣、阔叶地衣和枝状地衣对生境水分条件的适应。研究表明PV曲线及水势参数不适用于地衣类群整体的抗旱性评价, 和其他生物类群进行抗旱性比较时更要慎用。

关键词: 地衣, 水势参数, 内部含水量, 压力-体积曲线, 主成分分析, 变水植物

Abstract:

Aims Epiphytic lichens are a widespread and significant component of forest ecosystems, and play an important role in biodiversity maintaining and water and nutrient cycling. This study aims to illustrate the water potential characteristics of the epiphytic lichen group and to assess whether the pressure-volume (PV) curve and relative parameters are suitable for exploring the response of lichens to drought stress.
Methods The water potential parameters and PV curves of 15 epiphytic lichen species from five functional groups were measured in the subtropical forests in the Ailao Mountains, Yunnan Province.
Important findings The internal water content (WC_internal) and symplast water content (Rs) of cyanolichens were significantly higher than those of chlorolichens, while other parameters showed no significant differences. We only observed significant differences in WC_internal, Rs and relative water content at turgor loss point (RWC_TLP) among different functional groups of lichens. All parameters differed significantly among species. Results of principal component analysis (PCA) further indicated that there are extensive limitations for PV curves and water potential parameters in evaluating the overall response of lichen species to water stress and in revealing the selection strategy of water-related habitats, largely due to the influence of the photobiont type and growth form. However, the adaptation strategies of cyanolichens to habitat could be explained by the maximum internal water holding capacity, while the adaptation to water conditions could be explained by the saturated water osmotic potential (Ψsat) in narrowly lobed foliose lichens, and by the RWC_TLP in broadly lobed foliose and fruticose lichens. Our study suggests that PV curves and water potential parameters are not suitable for the general evaluation of the drought resistance of lichen communities, and should be used carefully in comparing the drought resistances between lichens and other plant groups.

Key words: epiphytic lichen, water potential, internal water content, pressure-volume curve, principal component analysis, poikilohydric plant

董琳琳, 普晓妍, 张璐璐, 宋亮, 鲁志云, 李苏. 亚热带森林附生地衣压力-体积曲线分析及其适用性. 植物生态学报, 2021, 45(3): 274-285. DOI: 10.17521/cjpe.2020.0344

DONG Lin-Lin, PU Xiao-Yan, ZHANG Lu-Lu, SONG Liang, LU Zhi-Yun, LI Su. Pressure-volume curve analysis of epiphytic lichens and its applicability in subtropical forests. Chinese Journal of Plant Ecology, 2021, 45(3): 274-285. DOI: 10.17521/cjpe.2020.0344

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https://www.plant-ecology.com/CN/Y2021/V45/I3/274

图/表 6

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物种 Species	共生藻型 Photobiont type	生长型 Growth form	功能群划分 Functional group	主要生境 Main habitat
网肺衣 Lobaria retigera (Lobret)	蓝藻 Cyanobacteria	叶状 Foliose	蓝藻型地衣 Cyanolichen (CYL)	阴湿至高光照湿润生境 Shaded to high-light and humid habitat
猫耳衣 Leptogium menziesii(Lepmen)	蓝藻 Cyanobacteria	叶状 Foliose	CYL	潮湿环境 Humid habitat
黑芽牛皮叶 Sticta fuliginosa(Stiful)	蓝藻 Cyanobacteria	叶状 Foliose	CYL	潮湿高光照生境 Humid and high-light habitat
针芽肺衣 Lobaria isidiophora(Lobisi)	绿藻 Green alga	阔叶型 Broadly lobed foliose	阔叶绿藻型地衣 Broadly lobed foliose chlorolichens (BFL)	中光至高光照生境 Medium to high-light habitat
网大叶梅 Parmotrema reticulatum (Parret)	绿藻 Green alga	阔叶型 Broadly lobed foliose	BFL	中光至高光照、潮湿生境 Medium to high-light, and humid habitat
平滑牛皮叶 Sticta nylanderiana (Stinyl)	绿藻 Green alga	阔叶型 Broadly lobed foliose	BFL	中光至高光照生境 Medium to high-light habitat
黑腹绵腹衣 Anzia hypomelaena (Anzhyp)	绿藻 Green alga	狭叶型 Narrowly lobed foliose	狭叶绿藻型地衣 Narrowly lobed foliose chlorolichens (NFL)	高光照生境 High-light habitat
短根条衣 Everniastrum nepalense (Evenep)	绿藻 Green alga	狭叶型 Narrowly lobed foliose	NFL	高光照生境 High-light habitat
卷梢哑铃孢 Heterodermia boryi (Hetbor)	绿藻 Green alga	狭叶型 Narrowly lobed foliose	NFL	高光照生境 High-light habitat
云南袋衣 Hypogymnia yunnanensis (Hypyun)	绿藻 Green alga	狭叶型 Narrowly lobed foliose	NFL	高光照生境 High-light habitat
裂髓树花 Ramalina conduplicans (Ramcon)	绿藻 Green alga	直立枝状 Shrubby fruticose	直立枝状绿藻型地衣 Shrubby fruticose chlorolichens (SFL)	高光照生境 High-light habitat
槽枝衣 Sulcaria sulcata (Sulsul)	绿藻 Green alga	直立枝状 Shrubby fruticose	SFL	高光照生境 High-light habitat
多花松萝 Usnea florida (Usnflo)	绿藻 Green alga	直立枝状 Shrubby fruticose	SFL	高光照生境 High-light habitat
巢松萝 Usnea dasopoga (Usndas)	绿藻 Green alga	悬垂枝状 Pendent fruticose	悬垂枝状绿藻型地衣 Pendent fruticose chlorolichens (PFL)	高光照且干燥生境 High-light and dry habitat
长松萝 Usnea longissima (Usnlon)	绿藻 Green alga	悬垂枝状 Pendent fruticose	PFL	高光照潮湿生境 High-light and humid habitat

物种 Species	共生藻型 Photobiont type	生长型 Growth form	功能群划分 Functional group	主要生境 Main habitat
网肺衣 Lobaria retigera (Lobret)	蓝藻 Cyanobacteria	叶状 Foliose	蓝藻型地衣 Cyanolichen (CYL)	阴湿至高光照湿润生境 Shaded to high-light and humid habitat
猫耳衣 Leptogium menziesii(Lepmen)	蓝藻 Cyanobacteria	叶状 Foliose	CYL	潮湿环境 Humid habitat
黑芽牛皮叶 Sticta fuliginosa(Stiful)	蓝藻 Cyanobacteria	叶状 Foliose	CYL	潮湿高光照生境 Humid and high-light habitat
针芽肺衣 Lobaria isidiophora(Lobisi)	绿藻 Green alga	阔叶型 Broadly lobed foliose	阔叶绿藻型地衣 Broadly lobed foliose chlorolichens (BFL)	中光至高光照生境 Medium to high-light habitat
网大叶梅 Parmotrema reticulatum (Parret)	绿藻 Green alga	阔叶型 Broadly lobed foliose	BFL	中光至高光照、潮湿生境 Medium to high-light, and humid habitat
平滑牛皮叶 Sticta nylanderiana (Stinyl)	绿藻 Green alga	阔叶型 Broadly lobed foliose	BFL	中光至高光照生境 Medium to high-light habitat
黑腹绵腹衣 Anzia hypomelaena (Anzhyp)	绿藻 Green alga	狭叶型 Narrowly lobed foliose	狭叶绿藻型地衣 Narrowly lobed foliose chlorolichens (NFL)	高光照生境 High-light habitat
短根条衣 Everniastrum nepalense (Evenep)	绿藻 Green alga	狭叶型 Narrowly lobed foliose	NFL	高光照生境 High-light habitat
卷梢哑铃孢 Heterodermia boryi (Hetbor)	绿藻 Green alga	狭叶型 Narrowly lobed foliose	NFL	高光照生境 High-light habitat
云南袋衣 Hypogymnia yunnanensis (Hypyun)	绿藻 Green alga	狭叶型 Narrowly lobed foliose	NFL	高光照生境 High-light habitat
裂髓树花 Ramalina conduplicans (Ramcon)	绿藻 Green alga	直立枝状 Shrubby fruticose	直立枝状绿藻型地衣 Shrubby fruticose chlorolichens (SFL)	高光照生境 High-light habitat
槽枝衣 Sulcaria sulcata (Sulsul)	绿藻 Green alga	直立枝状 Shrubby fruticose	SFL	高光照生境 High-light habitat
多花松萝 Usnea florida (Usnflo)	绿藻 Green alga	直立枝状 Shrubby fruticose	SFL	高光照生境 High-light habitat
巢松萝 Usnea dasopoga (Usndas)	绿藻 Green alga	悬垂枝状 Pendent fruticose	悬垂枝状绿藻型地衣 Pendent fruticose chlorolichens (PFL)	高光照且干燥生境 High-light and dry habitat
长松萝 Usnea longissima (Usnlon)	绿藻 Green alga	悬垂枝状 Pendent fruticose	PFL	高光照潮湿生境 High-light and humid habitat

共生藻型 Photobi-ont type	生长型 Growth form	物种 Species	内部含水量 WC_internal (%, Dry mass)	共质体水 Rs (%, Dry mass)	质外体水 Ra (%, Dry mass)	膨压损失点的相对含水量 RWC_TLP(%)	膨压损失点的渗透势Ψ_TLP (MPa)	饱和渗透势 Ψsat (MPa)
蓝藻 Cyanob-acteria			328.66 ± 36.84 ^Ac	322.08 ± 36.79 ^Ac	6.58 ± 1.19 ^Aab	40.96 ± 3.97 ^Abc	-3.44 ± 0.33 ^Aa	-1.29 ± 0.13 ^Aab
	叶状 Foliose	网肺衣 Lobaria retigera	214.86 ± 11.54	208.02 ± 11.14	6.85 ± 2.64	55.34 ± 2.95	-2.84 ± 0.35	-1.55 ± 0.24
		猫耳衣 Leptogium menziesii	497.80 ± 43.93	492.40 ± 43.56	5.40 ± 1.44	22.68 ± 1.49	-3.86 ± 0.53	-0.85 ± 0.14
		黑芽牛皮叶 Sticta fuliginosa	273.31 ± 32.08	265.81 ± 29.86	7.49 ± 2.30	44.86 ± 3.91	-3.63 ± 0.77	-1.46 ± 0.20
绿藻 Green alga			155.28 ± 8.95 ^B	149.71 ± 8.73 ^B	5.57 ± 0.45 ^A	46.93 ± 1.54 ^A	-3.80 ± 0.22 ^A	-1.71 ± 0.11 ^A
	阔叶型 Broadly lobed foliose		137.05 ± 7.67 ^ab	130.05 ± 7.18 ^ab	7.00 ± 0.80 ^a	56.89 ± 2.73 ^a	-3.45 ± 0.30 ^a	-1.89 ± 0.18 ^c
		针芽肺衣 Lobaria isidiophora	133.11 ± 10.40	128.03 ± 9.95	5.08 ± 1.20	65.68 ± 3.14	-3.85 ± 0.37	-2.47 ± 0.24
		网大叶梅 Parmotrema reticulatum	119.18 ± 12.49	112.58 ± 11.77	6.60 ± 0.79	45.66 ± 3.44	-3.41 ± 0.78	-1.44 ± 0.32
		平滑牛皮叶 Sticta nylanderiana	158.87 ± 12.26	149.55 ± 11.44	9.32 ± 1.57	59.34 ± 2.00	-3.08 ± 0.34	-1.75 ± 0.18
	狭叶型 Narrowly lobed foliose		185.42 ± 23.36 ^ab	179.71 ± 22.70 ^ab	5.71 ± 0.93 ^ab	45.75 ± 2.23 ^bc	-3.74 ± 0.43 ^ab	-1.72 ± 0.23 ^abc
		黑腹绵腹衣 Anzia hypomelaena	349.87 ± 25.89	339.44 ± 25.63	10.43 ± 1.08	33.88 ± 1.99	-1.58 ± 0.19	-0.50 ± 0.07
		短根条衣 Everniastrum nepalense	115.29 ± 6.35	111.44 ± 6.40	3.86 ± 0.40	48.52 ± 2.02	-4.74 ± 0.75	-2.28 ± 0.44
		卷梢哑铃孢 Heterodermia boryi	109.69 ± 9.46	107.74 ± 9.72	1.95 ± 0.46	51.66 ± 1.29	-4.98 ± 0.81	-2.52 ± 0.42
		云南袋衣 Hypogymnia yunnanensis	166.81 ± 7.57	160.20 ± 7.77	6.61 ± 2.20	48.94 ± 6.04	-3.64 ± 0.72	-1.56 ± 0.25
	直立枝状 Shrubby fruticose		153.06 ± 10.08 ^a	149.03 ± 9.88 ^a	4.03 ± 0.77 ^b	39.15 ± 2.46 ^c	-4.82 ± 0.45 ^b	-1.85 ± 0.23 ^bc
		裂髓树花 Ramalina conduplicans	148.43 ± 14.35	142.97 ± 13.80	5.46 ± 1.10	48.02 ± 2.01	-5.82 ± 0.89	-2.70 ± 0.42
		槽枝衣 Sulcaria sulcata	128.74 ± 8.78	125.69 ± 8.03	3.05 ± 1.80	40.92 ± 2.38	-3.97 ± 0.54	-1.56 ± 0.16
		多花松萝 Usnea florida	182.00 ± 20.55	178.44 ± 20.20	3.57 ± 0.96	28.52 ± 2.35	-4.68 ± 0.79	-1.29 ± 0.26
	悬垂枝状 Pendent fruticose		125.67 ± 10.85 ^b	120.23 ± 10.56 ^b	5.44 ± 0.78 ^ab	46.03 ± 3.63 ^b	-2.92 ± 0.28 ^a	-1.19 ± 0.06 ^a
		巢松萝 Usnea dasopoga	153.47 ± 10.12	148.05 ± 9.08	5.43 ± 1.20	40.86 ± 3.71	-3.05 ± 0.33	-1.14 ± 0.10
		长松萝 Usnea longissima	97.86 ± 6.33	92.41 ± 5.72	5.45 ± 1.15	51.20 ± 5.65	-2.79 ± 0.48	-1.24 ± 0.08

共生藻型 Photobi-ont type	生长型 Growth form	物种 Species	内部含水量 WC_internal (%, Dry mass)	共质体水 Rs (%, Dry mass)	质外体水 Ra (%, Dry mass)	膨压损失点的相对含水量 RWC_TLP(%)	膨压损失点的渗透势Ψ_TLP (MPa)	饱和渗透势 Ψsat (MPa)
蓝藻 Cyanob-acteria			328.66 ± 36.84 ^Ac	322.08 ± 36.79 ^Ac	6.58 ± 1.19 ^Aab	40.96 ± 3.97 ^Abc	-3.44 ± 0.33 ^Aa	-1.29 ± 0.13 ^Aab
	叶状 Foliose	网肺衣 Lobaria retigera	214.86 ± 11.54	208.02 ± 11.14	6.85 ± 2.64	55.34 ± 2.95	-2.84 ± 0.35	-1.55 ± 0.24
		猫耳衣 Leptogium menziesii	497.80 ± 43.93	492.40 ± 43.56	5.40 ± 1.44	22.68 ± 1.49	-3.86 ± 0.53	-0.85 ± 0.14
		黑芽牛皮叶 Sticta fuliginosa	273.31 ± 32.08	265.81 ± 29.86	7.49 ± 2.30	44.86 ± 3.91	-3.63 ± 0.77	-1.46 ± 0.20
绿藻 Green alga			155.28 ± 8.95 ^B	149.71 ± 8.73 ^B	5.57 ± 0.45 ^A	46.93 ± 1.54 ^A	-3.80 ± 0.22 ^A	-1.71 ± 0.11 ^A
	阔叶型 Broadly lobed foliose		137.05 ± 7.67 ^ab	130.05 ± 7.18 ^ab	7.00 ± 0.80 ^a	56.89 ± 2.73 ^a	-3.45 ± 0.30 ^a	-1.89 ± 0.18 ^c
		针芽肺衣 Lobaria isidiophora	133.11 ± 10.40	128.03 ± 9.95	5.08 ± 1.20	65.68 ± 3.14	-3.85 ± 0.37	-2.47 ± 0.24
		网大叶梅 Parmotrema reticulatum	119.18 ± 12.49	112.58 ± 11.77	6.60 ± 0.79	45.66 ± 3.44	-3.41 ± 0.78	-1.44 ± 0.32
		平滑牛皮叶 Sticta nylanderiana	158.87 ± 12.26	149.55 ± 11.44	9.32 ± 1.57	59.34 ± 2.00	-3.08 ± 0.34	-1.75 ± 0.18
	狭叶型 Narrowly lobed foliose		185.42 ± 23.36 ^ab	179.71 ± 22.70 ^ab	5.71 ± 0.93 ^ab	45.75 ± 2.23 ^bc	-3.74 ± 0.43 ^ab	-1.72 ± 0.23 ^abc
		黑腹绵腹衣 Anzia hypomelaena	349.87 ± 25.89	339.44 ± 25.63	10.43 ± 1.08	33.88 ± 1.99	-1.58 ± 0.19	-0.50 ± 0.07
		短根条衣 Everniastrum nepalense	115.29 ± 6.35	111.44 ± 6.40	3.86 ± 0.40	48.52 ± 2.02	-4.74 ± 0.75	-2.28 ± 0.44
		卷梢哑铃孢 Heterodermia boryi	109.69 ± 9.46	107.74 ± 9.72	1.95 ± 0.46	51.66 ± 1.29	-4.98 ± 0.81	-2.52 ± 0.42
		云南袋衣 Hypogymnia yunnanensis	166.81 ± 7.57	160.20 ± 7.77	6.61 ± 2.20	48.94 ± 6.04	-3.64 ± 0.72	-1.56 ± 0.25
	直立枝状 Shrubby fruticose		153.06 ± 10.08 ^a	149.03 ± 9.88 ^a	4.03 ± 0.77 ^b	39.15 ± 2.46 ^c	-4.82 ± 0.45 ^b	-1.85 ± 0.23 ^bc
		裂髓树花 Ramalina conduplicans	148.43 ± 14.35	142.97 ± 13.80	5.46 ± 1.10	48.02 ± 2.01	-5.82 ± 0.89	-2.70 ± 0.42
		槽枝衣 Sulcaria sulcata	128.74 ± 8.78	125.69 ± 8.03	3.05 ± 1.80	40.92 ± 2.38	-3.97 ± 0.54	-1.56 ± 0.16
		多花松萝 Usnea florida	182.00 ± 20.55	178.44 ± 20.20	3.57 ± 0.96	28.52 ± 2.35	-4.68 ± 0.79	-1.29 ± 0.26
	悬垂枝状 Pendent fruticose		125.67 ± 10.85 ^b	120.23 ± 10.56 ^b	5.44 ± 0.78 ^ab	46.03 ± 3.63 ^b	-2.92 ± 0.28 ^a	-1.19 ± 0.06 ^a
		巢松萝 Usnea dasopoga	153.47 ± 10.12	148.05 ± 9.08	5.43 ± 1.20	40.86 ± 3.71	-3.05 ± 0.33	-1.14 ± 0.10
		长松萝 Usnea longissima	97.86 ± 6.33	92.41 ± 5.72	5.45 ± 1.15	51.20 ± 5.65	-2.79 ± 0.48	-1.24 ± 0.08

水势参数 Water potential parameter	PC1	PC2
共质体水 Rs	0.584	-
质外体水 Ra	0.307	-0.818
质壁分离时相对含水量 RWC_TLP	-0.494	-0.565
饱和渗透势 Ψsat	0.566	-
标准差 Standard deviation	1.451	1.038
方差贡献率 Proportion of variance	52.66%	26.94%
方差累计贡献率 Cumulative proportion	52.66%	79.60%

亚热带森林附生地衣压力-体积曲线分析及其适用性

Pressure-volume curve analysis of epiphytic lichens and its applicability in subtropical forests

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