哀牢山山地森林不同附生地衣功能群的水分关系和光合生理特征

doi:10.17521/cjpe.2015.0445

摘要/Abstract

摘要：

附生地衣是热带和亚热带山地森林生态系统中重要的结构性组分, 在生物多样性保护、环境监测、养分循环中发挥着重要作用。附生地衣按共生藻、生活型和繁殖策略的不同可划分为不同的功能群, 不同附生地衣功能群的分布格局存在较大的差异, 然而其生理生态机制仍不清楚。该研究以我国西南地区哀牢山亚热带山地森林中的附生地衣优势类群为研究对象, 对该地区蓝藻地衣、阔叶地衣、狭叶地衣及枝状地衣4种功能群的8种附生地衣的水分关系、光合生理特征等进行了测定分析, 结果显示: 不同功能群附生地衣的持水力和失水速率均存在差异, 其中蓝藻地衣具有较高的最大水分含量, 而枝状地衣的失水速率较快; 过高和过低的水分含量都会抑制附生地衣的光合作用, 但抑制程度有所差异; 蓝藻地衣的光合作用最适水分含量比较高, 表明它们的光合生理活动对水分条件要求较高, 所以它们偏好潮湿的生境, 同时蓝藻地衣的光补偿点比较低但光饱和点却不低, 反映出它们具有较宽的光强适应范围, 所以蓝藻地衣能够同时分布于强光和弱光生境中; 枝状地衣的光合最适水分含量较低, 表明它们的光合生理活动对水分条件要求不是很高, 能够适应较为干旱的环境, 同时枝状地衣的光补偿点和光饱和点都很高, 说明它们的光合生理活动对光照条件要求比较高, 所以它们广泛分布于强光生境中; 阔叶和狭叶地衣的光补偿点比较高, 说明它们更适应有充足光照条件的生境。

关键词: 附生地衣, 功能群, 水分关系, 光合作用, 变水植物

Abstract:

Aims There are abundant epiphytic lichens in the tropical and subtropical montane forest ecosystems, which are important components of forest canopy and play a vital role in biodiversity conservation, environmental monitoring and nutrient cycling. In accordance with photobiont type, growth form and reproductive strategy, the epiphytic lichens can be divided into different functional groups, with different distribution patterns. In this study we aim to explain this phenomenon from the perspective of physiological ecology. Methods The maximum water content, water loss curves, photosynthetic water and light response curves were determined in four epiphytic lichen functional groups, including cyanolichens, fruticose lichens, broadly lobed foliose lichens and narrowly lobed foliose lichens. Important findings The functional characteristics of epiphytic lichens influence their maximum water-holding capacity and rate of water loss. The cyanolichens have higher maximum water content, while the fruticose lichens have a faster water loss. The cyanolichens that are widely distributed in the moist habitats require particularly high moisture for their photosynthetic activities; their optimal water content for photosynthesis is higher in comparison with other groups. They also have a low light compensation point and a high light saturation point, which explain the wide range of light intensity of the habitat. The fruticose lichens, widely distributed in the relatively arid habitats with high irradiance, have high light compensation point and light saturation point, and low optimum water content for photosynthesis. The broadly lobed foliose lichens and the narrowly lobed foliose lichens have a high light compensation point and light saturation point; they preferably occur in habitats with strong light.

Key words: epiphytic lichen, functional groups, water relation, photosynthesis, poikilohydry

胡涛, 李苏, 柳帅, 刘文耀, 陈曦, 宋亮, 陈泉. 哀牢山山地森林不同附生地衣功能群的水分关系和光合生理特征. 植物生态学报, 2016, 40(8): 810-826. DOI: 10.17521/cjpe.2015.0445

Tao HU, Su LI, Shuai LIU, Wen-Yao LIU, Xi CHEN, Liang SONG, Quan CHEN. Water relations and photosynthetic characteristics in different functional groups of epiphytic lichens in montane forest of Ailaoshan. Chinese Journal of Plant Ecology, 2016, 40(8): 810-826. DOI: 10.17521/cjpe.2015.0445

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2015.0445

https://www.plant-ecology.com/CN/Y2016/V40/I8/810

图/表 10

表1 附生地衣功能群的划分

Table 1 Classification of functional groups of epiphytic lichens

功能群 Functional group	简称 Abbreviation	功能群特征 Functional traits
蓝藻地衣 Cyanolichens	CYL	共生藻为蓝藻或包含蓝藻 With cyanobacteria
枝状地衣 Fruticose lichens	FRL	枝状或灌丛状, 共生藻为绿藻 Fruticose, with green algae
阔叶地衣 Broadly lobed foliose lichens	BFL	大型阔叶状, 共生藻为绿藻 Broadly lobed foliose, with green algae
狭叶地衣 Narrowly lobed foliose lichens	NFL	小型或中型细叶状, 共生藻为绿藻 Narrowly lobed foliose, with green algae

表2 不同附生地衣的持水量与失水速率(平均值±标准误差)

Table 2 Water-holding and rate of water loss in different epiphytic lichens (mean ± SE)

物种 Species	单位干质量持水量 Water-holding capacity per dry mass (n = 10)	最大含水量 Maximum water content (% dry mass)	T₈₀ (h)	T₅₀ (h)	T₂₀ (h)
皮革肾岛衣 Nephromopsis pallescens	3.01 ± 0.09^cd	308.8	0.59	1.77	3.78
针芽肺衣 Lobaria isidiophora	2.84 ± 0.16^d	315.1	0.61	1.83	3.86
网肺衣 Lobaria retigera	3.76 ± 0.20^bcd	373.1	0.63	1.87	3.82
猫耳衣 Leptogium menziesii	10.49 ± 0.49^a	1 104.1	1.03	2.94	5.64
云南袋衣 Hypogymnia yunnanensis	5.01 ± 0.30^abc	473.6	0.56	1.69	3.69
白绵腹衣 Anzia leucobatoides	5.70 ± 0.19^ab	514.2	1.83	4.85	8.28
多花松萝 Usnea florida	3.22 ± 0.09^cd	193.6	0.4	1.26	3.14
裂髓树花 Ramalina conduplicans	6.07 ± 0.18^ab	449.2	0.48	1.48	3.41

图1 各附生地衣干质量和持水量之间的关系(n = 10)。

Fig. 1 Relationship between water-holding capacity (WHC) and dry mass in different epiphytic lichens (n = 10).

图2 不同附生地衣失水动力学曲线。A, 皮革肾岛衣。B, 针芽肺衣。C, 网肺衣。D, 猫耳衣。E, 云南袋衣。F, 白绵腹衣。G, 多花松萝。H, 裂髓树花。WC表示含水量。不同形状的图列符号表示不同的样品(n = 3-5)。

Fig. 2 Water release curves for different epiphytic lichens. A, Nephromopsis pallescens. B, Lobaria isidiophora. C, Lobaria retigera. D, Leptogium menziesii. E, Hypogymnia yunnanensis. F, Anzia leucobatoides. G, Usnea florida. H, Ramalina conduplicans. WC in graphs refers to water content. Different shapes of symbols represent different samples (n = 3-5).

图3 不同附生地衣失水过程中不同时刻的含水量。含水量表示为地衣含水量占最大含水量的百分比。

Fig. 3 Water content in different epiphytic lichens at different times during water loss. Hydration (%) refers to the percentage of maximum water content.

图4 不同附生地衣光合作用水分响应曲线。A, 皮革肾岛衣。B, 针芽肺衣。C, 网肺衣。D, 猫耳衣。E, 云南袋衣(R2 = 0.83, p < 0.001)。F, 白绵腹衣。G, 多花松萝。H, 裂髓树花。Pn和WC分别表示净光合速率和含水量。不同形状的图列符号表示不同的样品(n = 4, 5)。

Fig. 4 Photosynthetic water response curves for different epiphytic lichens. A, Nephromopsis pallescens. B, Lobaria isidiophora. C, Lobaria retigera. D, Leptogium menziesii. E, Hypogymnia yunnanensis. F, Anzia leucobatoides. G, Usnea florida. H, Ramalina conduplicans. Pn and WC in graphs refer to net photosynthesis rate and water content, respectively. Different shapes of symbols represent different samples (n = 4, 5).

图5 附生地衣光合最适水分含量和最大水分含量的关系(A), 水分补偿点和最适水分补偿点的关系(B), 水分含量和光合作用的关系(C)。图C中净光合速率比表示为占最大净光合速率的百分比。

Fig. 5 The relationship between optimal water content and maximum water content (A), the relationship between water compensation point and optimal water content (B), and the relationship between photosynthesis and water content in different epiphytic lichens (C). Photosynthesis is expressed as the percentage of max net photosynthesis.

表3 各附生地衣光合作用的最适水分含量和水分补偿点, 光合作用和水分含量关系曲线的初始斜率

Table 3 Optimal water content, water compensation point, and initial slope of water content-dependent net photosynthesis in different epiphytic lichens

物种 Species	最适水分含量 Optimal water content (% dry mass)	水分补偿点 Water compensation point (% dry mass)	初始斜率 Initial slope
皮革肾岛衣 Nephromopsis pallescens	232.6	24.2	0.75
针芽肺衣 Lobaria isidiophora	119.0	16.0	2.11
网肺衣 Lobaria retigera	172.9	44.4	1.11
猫耳衣 Leptogium menziesii	419.5	16.3	0.50
云南袋衣 Hypogymnia yunnanensis	175.0	22.3	0.98
白绵腹衣 Anzia leucobatoides	298.9	19.7	0.64
多花松萝 Usnea florida	93.0	27.0	2.59
裂髓树花 Ramalina conduplicans	139.4	33.7	1.92

图6 各附生地衣光响应曲线。A, 皮革肾岛衣。B, 针芽肺衣。C, 网肺衣。D, 猫耳衣。E, 云南袋衣(R2 = 0.83, p < 0.001)。F, 白绵腹衣。G, 多花松萝。H, 裂髓树花。不同形状的图列符号表示不同的样品(n = 4, 5)。

Fig. 6 Photosynthetic light response curves for different epiphytic lichens. A, Nephromopsis pallescens. B, Lobaria isidiophora. C, Lobaria retigera. D, Leptogium menziesii. E, Hypogymnia yunnanensis. F, Anzia leucobatoides. G, Usnea florida. H, Ramalina conduplicans. Pn and I in graphs refer to net photosynthesis and light intensity, respectively. Different shapes of symbols represent different samples (n = 4, 5).

表4 各附生地衣的光响应曲线参数(平均值±标准误差, n = 5)

Table 4 Parameters of light response curves for different epiphytic lichens (mean ± SE, n = 5)

物种 Species	光补偿点 LCP (μmol·m^-2·s^-1)	光饱和点 LSP (μmol·m^-2·s^-1)	最大净光合速率 P_nmax (μmol·m^-2·s^-1)	暗呼吸速率 R_d (μmol·m^-2·s^-1)
皮革肾岛衣 Nephromopsis pallescens	46.6 ± 2.3^a	407.5 ± 14.6^b	0.88 ± 0.03^e	0.66 ± 0.04^ab
针芽肺衣 Lobaria isidiophora	43.4 ± 2.6^ab	466.4 ± 22.7^b	1.77 ± 0.06^c	0.83 ± 0.03^a
网肺衣 Lobaria retigera	20.3 ± 2.2^c	416.3 ± 25.4^b	2.85 ± 0.05^a	0.63 ± 0.03^ab
猫耳衣 Leptogium menziesii	23.2 ± 0.3^c	322.4 ± 25.4^b	1.61 ± 0.02^c	0.60 ± 0.02^ab
云南袋衣 Hypogymnia yunnanensis	41.1 ± 3.7^ab	410.7 ± 40.6^b	1.72 ± 0.05^c	0.94 ± 0.16^a
白绵腹衣 Anzia leucobatoides	36.0 ± 2.1^b	441.7 ± 37.6^b	2.17 ± 0.07^b	0.90 ± 0.04^a
多花松萝 Usnea florida	35.5 ± 2.2^b	610.9 ± 54.3^a	1.29 ± 0.07^d	0.37 ± 0.01^b
裂髓树花 Ramalina conduplicans	38.6 ± 2.3^ab	613.4 ± 42.3^a	2.10 ± 0.08^b	0.95 ± 0.03^a

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