环境胁迫影响三峡库区消落带草本和木本植物生理生态特征的meta分析

doi:10.17521/cjpe.2023.0127

摘要/Abstract

摘要：

三峡库区消落带适生草本与木本植物对环境胁迫的响应是生态系统结构与功能演变的重要机制。然而, 目前对消落带胁迫环境下草本和木本植物的响应特征及是否存在类群差异尚不清楚。该研究基于meta分析, 探讨了三峡消落带典型环境胁迫(水淹、干旱胁迫)对适生草本和木本植物生理生态特征的影响, 从物种和生活型的角度揭示了消落带适生植物类群对水淹、干旱胁迫的响应规律。结果表明: (1)草本植物和木本植物在环境胁迫下均表现出明显的生长发育抑制以及生理生化策略的适应性响应, 主要表现在各组织器官生物量、光合速率的下降, 丙二醛和脯氨酸含量的增加, 以及抗氧化酶(超氧化物歧化酶、过氧化物酶、过氧化氢酶)活性增强。(2)水淹胁迫对草本植物生物量负效应最强, 干旱胁迫次之; 水淹胁迫对木本植物总叶绿素含量和光合速率负效应最大, 干旱胁迫次之, 表现出环境胁迫作用的生活型效应; 但草本与木本植物的总叶绿素含量均对水淹胁迫响应最敏感, 而木本植物对干旱胁迫表现出一定的内稳态。木本植物在生理生化方面对水淹胁迫的响应比草本植物更显著, 且主要体现在气孔导度、丙二醛含量、脯氨酸含量和过氧化物酶活性方面的响应与草本植物存在差异; 而草本植物对干旱胁迫的响应比木本植物更显著, 主要表现为总叶绿素含量、水分利用效率、抗氧化酶(超氧化物歧化酶、过氧化物酶、过氧化氢酶)活性方面的响应与木本植物存在差异。(3)草本和木本植物对不同环境胁迫的形态响应具有组织特异性差异。草本植物主要以根部、茎部和叶片形态的变化适应水淹和干旱胁迫, 木本植物则主要以根部形态的变化为主。

关键词: 三峡库区, 植物, 生理生态, 环境胁迫, meta分析

Abstract:

Aims The response of herbaceous and woody plants to environmental stress is an important mechanism for the evolution of ecosystem structure and function in hydro-fluctuation belt of Three Gorges Reservoir. However, the pertinent response characteristics and whether there are group differences in these stress responses are still unclear.

Methods Here, we undertook a meta-analysis of the effects of typical environmental stresses (flooding and drought stress) on the physiological and ecological characteristics of suitable herbaceous and woody plants in the Three Gorges hydro-fluctuation belt, with an aim to reveal the response of suitable plant groups to flooding and drought stress from the perspective of species and life forms.

Important findings The results showed that: (1) Herbaceous plants and woody plants showed obvious growth and development inhibition and adaptive response in physiological and biochemical strategies under environmental stress, mainly manifested in the decrease of biomass and photosynthetic rate of each tissue and organ, the increase of malondialdehyde and proline content, and the initiation of antioxidant enzyme system (superoxide dismutase, peroxidase, catalase). (2) Flooding stress had the strongest negative effect on herbaceous plant biomass, followed by drought stress. Flooding stress had the greatest negative effect on total chlorophyll content and photosynthetic rate of woody plants, followed by drought stress, showing the life-form effect of environmental stress. However, the total chlorophyll content of herbaceous and woody plants was most sensitive to flooding stress, while woody plants showed a certain homeostasis to drought stress. The physiological and biochemical responses of woody plants to flooding stress were more significant than those of herbaceous plants, and the responses were mainly different from those of herbaceous plants in stomatal conductance, malondialdehyde and proline content, and peroxidase activity. The herbaceous plants showed stronger response to drought stress than did woody plants, and the response of total chlorophyll content, water use efficiency and antioxidant enzyme (superoxide dismutase, peroxidase, catalase) activity was different from that of woody plants. (3) The morphological responses of herbaceous and woody plants to different environmental stresses have tissue-specific differences. Herbaceous plants mainly adapt to flooding and drought stress by morphological changes of roots, stems and leaves, while woody plants mainly adapt via morphological changes of roots. In summary, this study presents an integrated view of the morphological and physiological characteristics of herbaceous and woody plants in response to environmental stress in the hydro-fluctuation belt of the Three Gorges Reservoir, which provided a scientific basis for evaluating the potential of different life-form vegetation in the ecological structure and functional restoration in this ecologically-important region.

Key words: Three Gorges Reservoir, plant, physiological ecology, environmental stress, meta-analysis

邓蓓, 王晓锋, 廖君. 环境胁迫影响三峡库区消落带草本和木本植物生理生态特征的meta分析. 植物生态学报, 2024, 48(5): 623-637. DOI: 10.17521/cjpe.2023.0127

DENG Bei, WANG Xiao-Feng, LIAO Jun. Ecophysiological responses of herbaceous and woody plants to environmental stresses in the riparian zone of Three Gorges Reservoir: a meta-analysis. Chinese Journal of Plant Ecology, 2024, 48(5): 623-637. DOI: 10.17521/cjpe.2023.0127

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2023.0127

https://www.plant-ecology.com/CN/Y2024/V48/I5/623

图/表 9

表1 三峡库区消落带常见草本植物和木本植物

Table 1 Common herbs and woody plants in the riparian zone of the Three Gorges Reservoir

生活型 Life form	科名 Family	种名 Species
草本 Herb	莎草科 Cyperaceae	香附子 Cyperus rotundus
	禾本科 Poaceae	芦苇 Phragmites australis
		扁穗牛鞭草 Hemarthria compressa
		狗牙根 Cynodon dactylon
		狗尾草 Setaria viridis
		野古草 Arundinella hirta
		香根草 Chrysopogon zizanioides
		双穗雀稗 Paspalum distichum
	蓼科 Polygonaceae	水蓼 Persicaria hydropiper
		酸模叶蓼 Persicaria lapathifolia
	菊科 Asteraceae	苍耳 Xanthium sibiricum
		马兰 Aster indicus
	苋科 Amaranthaceae	喜旱莲子草 Alternanthera philoxeroides
	三白草科 Saururaceae	蕺菜 Houttuynia cordata
	天南星科 Araceae	菖蒲 Acorus Calamus
木本 Wood	胡桃科 Juglandaceae	枫杨 Pterocarya stenoptera
	壳斗科 Fagaceae	栓皮栎 Quercus variabilis
	松科 Pinaceae	湿地松 Pinus elliottii
	杉科 Taxodiaceae	水松 Glyptostrobus pensilis
		水杉 Metasequoia glyptostroboides
		池杉 Taxodium ascendens
		落羽杉 Taxodium distichum
		中山杉 Taxodium ‘Zhongshanshan’
	大戟科 Euphorbiaceae	乌桕 Triadica sebifera
	桑科 Moraceae	桑 Morus alba
	桦木科 Betulaceae	水桦 Betula nigra
	杨柳科 Salicaceae	旱柳 Salix matsudana
		秋华柳 Salix variegate
		南川柳 Salix rosthornii
	樟科 Lauraceae	樟 Camphora officinarum
	含羞草科 Mimosaceae	银合欢 Leucaena leucocephala
	金缕梅科 Hamamelidaceae	中华蚊母树 Distylium chinen
	黄杨科 Buxaceae	宜昌黄杨 Buxus ichangensis

图1 水淹条件下草本植物与木本植物的生物量、形态变化。水平虚线表示响应比为0, 方块和误差棒表示平均值和95%置信区间。误差线与0线相交表示处理与对照不存在显著差异(p > 0.05); 误差线在0线以上表示为显著正相关, 误差线在0线以下表示为显著负相关(p < 0.05)。括号内数字表示物种样本数量。Ac, 菖蒲; Ap, 喜旱莲子草; Cd, 狗牙根; Cr, 香附子; Cz, 香根草; Dc, 中华蚊母树; Hc, 扁穗牛鞭草; Mg, 水杉; Pe, 湿地松; Ph, 水蓼; Ps, 枫杨; Sr, 南川柳; Sv, 狗尾草; Tz, 中山杉。

Fig. 1 Biomass and morphological changes of herbaceous plants and woody plants under flooding conditions. The horizontal dotted line indicates that the response ratio is 0, and the square and the error bar indicate the mean and the 95% confidence interval. There was no significant difference between the treatment and the control in the intersection of the 0 line (p > 0.05). The error line is expressed as a significant positive correlation above the 0 line, and the error line is expressed as a significant negative correlation below the zero line (p < 0.05). Numbers in brackets indicate sample size. Ac, Acorus calamus; Ap, Alternanthera philoxeroides; Cd, Cynodon dactylon; Cr, Cyperus rotundus; Cz, Chrysopogon zizanioides; Dc, Distylium chinen; Hc, Hemarthria compressa; Mg, Metasequoia glyptostroboides; Pe, Pinus elliottii; Ph, Persicaria hydropiper; Ps, Pterocarya stenoptera; Sr, Salix rosthornii; Sv, Setaria viridis; Tz, Taxodium ‘Zhongshanshan’.

图2 水淹条件下草本植物与木本植物的光合色素含量和光合参数含量变化。水平虚线表示响应比为0, 方块和误差棒表示平均值和95%置信区间。误差线与0线相交表示处理与对照不存在显著差异(p > 0.05); 误差线在0线以上表示为显著正相关, 误差线在0线以下表示为显著负相关(p < 0.05)。括号内数字表示物种样本数量。空缺位置表示该物种无对应数据(A-E)。Cd, 狗牙根; Cr, 香附子; Cz, 香根草; Dc, 中华蚊母树; Hc, 扁穗牛鞭草; Ma, 桑; Pa, 芦苇; Pe, 湿地松; Ph, 水蓼; Ps, 枫杨; Sv, 狗尾草; Sva, 秋华柳; Ta, 池杉; Td, 落羽杉。

Fig. 2 Changes of photosynthetic pigment content and photosynthetic parameters of herbaceous plants and woody plants under flooding conditions. The horizontal dotted line indicates that the response ratio is 0, and the square and the error bar indicate the mean and the 95% confidence interval. There was no significant difference between the treatment and the control in the intersection of the 0 line (p > 0.05). The error line is expressed as a significant positive correlation above the 0 line, and the error line is expressed as a significant negative correlation below the zero line (p < 0.05). Numbers in brackets indicate sample size. The vacancy position indicates that the species has no corresponding data (A-E). Cd, Cynodon dactylon; Cr, Cyperus rotundus; Cz, Chrysopogon zizanioides; Dc, Distylium chinen; Hc, Hemarthria compressa; Ma, Morus alba; Pa, Phragmites australis; Pe, Pinus elliottii; Ph, Persicaria hydropiper; Ps, Pterocarya stenoptera; Sv, Setaria viridis; Sva, Salix variegate; Ta, Taxodium ascendens; Td, Taxodium distichum. Ci, intercellular CO2 concentration; Gs, stomatal conductivity; Pn, net photosynthetic rate; Tr, transpiration rate; T Chl, total chlorophyll.

图3 水淹条件下草本植物与木本植物的各项生理指标变化。水平虚线表示响应比为0, 方块和误差棒表示平均值和95%置信区间。误差线与0线相交表示处理与对照不存在显著差异(p > 0.05); 误差线在0线以上表示为显著正相关, 误差线在0线以下表示为显著负相关(p < 0.05)。括号内数字表示物种样本数量。空缺位置表示该物种无对应数据(A、B)。Ai, 马兰; Cd, 狗牙根; Co, 樟; Cr, 香附子; Dc, 中华蚊母树; Hc, 扁穗牛鞭草; Hco, 蕺菜; Mg, 水杉; Pd, 双穗雀稗; Pe, 湿地松; Pl, 酸模叶蓼; Sr, 南川柳; Sva, 秋华柳; Xs, 苍耳。CAT activity, 过氧化氢酶活性; MDA content, 丙二醛含量; POD activity, 过氧化物酶活性; Pro content, 脯氨酸含量; SOD activity, 超氧化物歧化酶活性。

Fig. 3 Changes of physiological indexes of herbaceous plants and woody plants under flooding. The horizontal dotted line indicates that the response ratio is 0, and the square and the error bar indicate the mean and the 95% confidence interval. There was no significant difference between the treatment and the control in the intersection of the 0 line (p > 0.05). The error line is expressed as a significant positive correlation above the 0 line, and the error line is expressed as a significant negative correlation below the zero line (p < 0.05). Numbers in brackets indicate sample size.The vacancy position indicates that the species has no corresponding data (A, B). Ai, Aster indicus; Cd, Cynodon dactylon; Co, Camphora officinarum; Cr, Cyperus rotundus; Dc, Distylium chinen; Hc, Hemarthria compressa; Hco, Houttuynia cordata; Mg, Metasequoia glyptostroboides; Pd, Paspalum distichum; Pe, Pinus elliottii; Pl, Persicaria lapathifolia; Sr, Salix rosthornii; Sva, Salix variegate; Xs, Xanthium sibiricum. CAT, catalase; MDA, malondialdehyde; Pro, proline; POD, peroxidase; SOD, superoxide dismutase.

图4 干旱条件下草本植物与木本植物的生物量、形态变化。水平虚线表示响应比为0, 方块和误差棒表示平均值和95%置信区间。误差线与0线相交表示处理与对照不存在显著差异(p > 0.05); 误差线在0线以上表示为显著正相关, 误差线在0线以下表示为显著负相关(p < 0.05)。括号内数字表示物种样本数量。Ar, 根表面积; As, 比根面积; H, 株高; No., 叶片数; L, 叶长; Lr, 侧根数; W, 叶宽。

Fig. 4 Biomass changes and morphological changes of herbaceous plants and woody plants under drought conditions. The horizontal dotted line indicates that the response ratio is 0, and the square and the error bar indicate the mean and the 95% confidence interval. There was no significant difference between the treatment and the control in the intersection of the 0 line (p > 0.05). The error line is expressed as a significant positive correlation above the 0 line, and the error line is expressed as a significant negative correlation below the zero line (p < 0.05). Numbers in brackets indicate sample size. Ar, root surface; As, specific root area; H, plant height; No., blade number; L, leaf length; Lr, lateral root number; W, leaf width.

图5 干旱条件下草本植物与木本植物的光合色素、光合参数、水分效率变化量。水平虚线表示响应比为0, 方块和误差棒表示平均值和95%置信区间。误差线与0线相交表示处理与对照不存在显著差异(p > 0.05); 误差线在0线以上表示为显著正相关, 误差线在0线以下表示为显著负相关(p < 0.05)。括号内数字表示物种样本数量。空缺位置表示该物种无对应数据(A-F)。Cd, 狗牙根; Hc, 扁穗牛鞭草; Ll, 银合欢; Ma, 桑; Pa, 芦苇; Pe, 湿地松; Ps, 枫杨; Ta, 池杉; Td, 落羽杉。

Fig. 5 Changes of photosynthetic pigments, photosynthetic parameters and water efficiency of herbaceous plants and woody plants under drought conditions. The horizontal dotted line indicates that the response ratio is 0, and the square and the error bar indicate the mean and the 95% confidence interval. There was no significant difference between the treatment and the control in the intersection of the 0 line (p > 0.05). The error line is expressed as a significant positive correlation above the 0 line, and the error line is expressed as a significant negative correlation below the zero line (p < 0.05). Numbers in brackets indicate sample size. The vacancy position indicates that the species has no corresponding data (A-F). Cd, Cynodon dactylon; Hc, Hemarthria compressa; Ll, Leucaena leucocephala; Ma, Morus alba; Pa, Phragmites australis; Pe, Pinus elliottii; Ps, Pterocarya stenoptera; Ta, Taxodium ascendens; Td, Taxodium distichum. Ci, intercellular CO2 concentration; Gs, stomatal conductivity; Pn, net photosynthetic rate; Tr, transpiration rate; T Chl, total chlorophyll; WUE, water use efficiency.

图6 干旱条件下草本植物与木本植物的各项生理指标变化。水平虚线表示响应比为0, 方块和误差棒表示平均值和95%置信区间。误差线与0线相交表示处理与对照不存在显著差异(p > 0.05); 误差线在0线以上表示为显著正相关, 误差线在0线以下表示为显著负相关(p < 0.05)。括号内数字表示物种样本数量。空缺位置表示该物种无对应数据(A、B)。Ac, 菖蒲; Cd, 狗牙根; Cz, 香根草; Hc, 扁穗牛鞭草; Ma, 桑; Mg, 水杉; Ps, 枫杨; Sr, 南川柳; Sva, 秋华柳; Tz, 中山杉。CAT activity, 过氧化氢酶活性; MDA content, 丙二醛含量; POD activity, 过氧化物酶活性; Pro content, 脯氨酸含量; SOD activity, 超氧化物歧化酶活性。

Fig. 6 Changes of physiological indexes of herbaceous plants and woody plants under drought. The horizontal dotted line indicates that the response ratio is 0, and the square and the error bar indicate the mean and the 95% confidence interval. There was no significant difference between the treatment and the control in the intersection of the 0 line (p > 0.05). The error line is expressed as a significant positive correlation above the 0 line, and the error line is expressed as a significant negative correlation below the zero line (p < 0.05). Numbers in brackets indicate sample size.The vacancy position indicates that the species has no corresponding data (A, B). Ac, Acorus Calamus; Cd, Cynodon dactylon; Cz, Chrysopogon zizanioides; Hc, Hemarthria compressa; Ma, Morus alba; Mg, Metasequoia glyptostroboides; Ps, Pterocarya stenoptera; Sr, Salix rosthornii; Sv, Setaria viridis; Tz, Taxodium ‘Zhongshanshan’. CAT, catalase; MDA, malondialdehyde; Pro, proline; POD, peroxidase; SOD, superoxide dismutase.

表2 不同胁迫下草本植物与木本植物的生物量、总叶绿素含量和净光合速率的变化量(%)和95%置信区间

Table 2 Changes (%) and 95% confidence interval of biomass, total chlorophyll (T Chl) content and net photosynthetic rate (Pn) of herbaceous plants and woody plants under different stresses

生活型 Life form	胁迫 Stress	生物量 Biomass	总叶绿素含量 T Chl content	净光合速率 P_n
草本 Herb	水淹胁迫 Flooding stress	-53.48 [-61.22, -44.19]	-12.93 [-20.28, -4.92]	-17.50 [-30.41, -2.20]
草本 Herb	干旱胁迫 Drought stress	-20.49 [-33.89, -4.39]	-11.64 [-18.00, -4.77]	-17.99 [-23.10, -12.54]
木本 Wood	水淹胁迫 Flooding stress	-35.68 [-44.96, -24.84]	-28.66 [-35.39, -21.23]	-36.06 [-42.01, -29.50]
木本 Wood	干旱胁迫 Drought stress	-19.92 [-28.89, -9.82]	1.23 [-3.68, 6.37]	-14.32 [-24.15, -3.23]

图7 草本植物与木本植物在环境胁迫下的形态差异及共同响应特征示意图。红色箭头表示减少或抑制, 绿色箭头表示增加或促进。绿色和红色箭头并列表示先促进后抑制。

Fig. 7 Schematic diagram of morphological differences and common response characteristics of herbaceous plants and woody plants under environmental stress. Red arrows indicate decrease or inhibition, and green arrows indicate increase or promotion. The green and red arrows are juxtaposed to indicate first promotion and then inhibition. CAT, catalase; POD, peroxidase; SOD, superoxide dismutase.

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