甘肃小苏干湖盐沼湿地盐地风毛菊叶形态-光合生理特征对淹水的响应

doi:10.17521/cjpe.2019.0132

植物生态学报 ›› 2019, Vol. 43 ›› Issue (8): 685-696.DOI: 10.17521/cjpe.2019.0132

所属专题：光合作用

甘肃小苏干湖盐沼湿地盐地风毛菊叶形态-光合生理特征对淹水的响应

李群,赵成章(),王继伟,文军,李子琴,马俊逸

西北师范大学地理与环境科学学院, 甘肃省湿地资源保护与产业发展工程研究中心, 兰州 730070

收稿日期:2019-05-30 修回日期:2019-08-09 出版日期:2019-08-20 发布日期:2020-01-03
通讯作者: 赵成章 ORCID:0000-0002-0127-8405
作者简介:李群: ORCID: 0000-0002-1279-4739
基金资助:
国家自然科学基金(41861009);国家自然科学基金(41461013)

Morphological and photosynthetic physiological characteristics of Saussurea salsa in response to flooding in salt marshes of Xiao Sugan Lake, Gansu, China

LI Qun,ZHAO Cheng-Zhang(),WANG Ji-Wei,WEN Jun,LI Zi-Qin,MA Jun-Yi

College of Geography and Environmental Science, Northwest Normal University, Research Center of Wetland Resources Protection and Industrial Development Engineering of Gansu Province, Lanzhou 730070, China

Received:2019-05-30 Revised:2019-08-09 Online:2019-08-20 Published:2020-01-03
Contact: ZHAO Cheng-Zhang ORCID:0000-0002-0127-8405
Supported by:
National Natural Science Foundation of China(41861009);National Natural Science Foundation of China(41461013)

摘要/Abstract

摘要：

盐沼湿地植物叶片功能性状对淹水的响应分析, 有助于探究植物叶片可塑性机制与光合生理特征间的内在关联性, 对深入理解盐沼湿地植物的生境抗逆性策略具有重要意义。根据小苏干湖湖水泛滥区静水持留时间长短分别设置: 轻度淹水区(静水持留30-90天)、中度淹水区(静水持留90-150天)、重度淹水区(静水持留150-210天) 3个试验样地, 以盐地风毛菊(Saussurea salsa)为研究对象, 研究了小苏干湖盐沼湿地盐地风毛菊叶片功能性状对淹水的响应。结果表明: 随着静水持留时间的增加, 轻度淹水区盐地风毛菊形态上采用小比叶面积(SLA)的肉质化小叶模式, 光合生理上具有高实际光合效率(Y(II))和低调节性能量耗散的量子产额(Y(NPQ))的协同变异; 重度淹水区盐地风毛菊形态和光合生理上则采用与轻度淹水区完全相反的协同变异策略; 在3个样地中, SLA与Y(II)、光化学淬灭(QP)和Y(NPQ)间均呈极显著相关关系; 叶绿素a含量和叶绿素b含量与调节性能量耗散的量子产额(Y(NO))均呈显著正相关关系。小苏干湖湖水泛滥区静水时空演变格局影响下, 盐地风毛菊种群通过改变叶面积、叶厚度和SLA等叶片形态特征, 适时调整叶片Y(II)和Y(NPQ)等光合生理特征, 实现植物叶片光合碳同化产物的收支平衡, 表现出对水盐异质性环境较强的耐受性, 反映了盐沼湿地植物在极端生存环境下的叶片可塑性和抗逆性机制。

关键词: 叶性状, 比叶面积, 实际光化学效率, 风毛菊, 小苏干湖

Abstract:

Aims The response of plant leaf functional traits to flooding in salt marshes is not only helpful in exploring the internal correlation between leaf plasticity mechanism and photosynthetic characteristics, but also of vital significance for gaining a better understanding on the stress resistance strategies of plants in salt marsh wetlands. The aim of this study is to investigate the responses of leaf functional traits of Saussurea salsa to flooding with the changes of water-logging durations.
Methods The research site was located in provincial migratory bird nature reserve in Xiao Sugan Lake, Gansu Province, China (39.22°-39.35° N, 94.45°-94.59° E). A sample belt was selected from the edge of the lakeshore to the end of the perennial tidewater line in the low-lying area along the north side of Xiao Sugan River. The duration of water-logging in the salt marsh wetland was measured base on the water level marked by the flood rise and retreat marks in Xiao Sugan Lake over the years. The sample belt was divided into 3 plots according to the water-logging duration: low flooding area (water-logging duration: 30-90 days); medium flooding area (water- logging duration: 90-150 days); and deep flooding area (water-logging duration: 150-210 days). Six subplots (2 m × 2 m) of S. salsa were selected from each of the three plots for a total of 18 subplots. We investigated community characteristics and population traits of S. salsa, soil moisture and soil electrical conductivity (EC). Six plants per subplot were selected for photosynthesis and chlorophyll fluorescence measurements. Foliar samples collected from each of the six S. salsa were taken to the laboratory for measurements of leaf traits (leaf area, thickness, dry mass and chlorophyll content).
Important findings The results showed that S. salsa changed the covariation strategy of S. salsa foliar morphology and photosynthesis with the extension of water-logging duration. In the low flooding area, S. salsa adopted fleshy lobular pattern with small specific leaf area (SLA), high effective quantum yield of photosynthetic system II (PSII) photochemistry in light (Y(II)) and low quantum yield of regulated energy dissipation (Y(NPQ)). However, S. salsa grew in the deep flooding area adopted completely opposite covariation strategy in foliar morphology and photosynthesis compared with those grew in the low flooding area. We observed a significant correlation between SLA and Y(II), photochemical quenching (QP), and Y(NPQ), as well as a significant positive correlation between the quantum yield of non-regulated energy dissipation (Y(NO)) and chlorophyll a content (C_a), chlorophyll b content (C_b) in all of the three plots. Under the influence of the spatio-temporal evolution pattern of still water in the flooded area of Xiao Sugan Lake, S. salsa population achieved the balance of the photosynthetic carbon budget by changing the morphological characteristics of leaves, such as leaf area, leaf thickness and SLA, timely adjusting the photosynthetic characteristics, such as Y(NPQ) and Y(II). Saussurea salsa showed strong tolerance to water and salt heterogeneity, reflecting the leaf plasticity and resistance mechanism of salt marsh wetland plant under extreme environments.

Key words: leaf traits, specific leaf area, actual photochemical efficiency of PSII, Saussurea salsa, Xiao Sugan Lake

李群, 赵成章, 王继伟, 文军, 李子琴, 马俊逸. 甘肃小苏干湖盐沼湿地盐地风毛菊叶形态-光合生理特征对淹水的响应. 植物生态学报, 2019, 43(8): 685-696. DOI: 10.17521/cjpe.2019.0132

LI Qun, ZHAO Cheng-Zhang, WANG Ji-Wei, WEN Jun, LI Zi-Qin, MA Jun-Yi. Morphological and photosynthetic physiological characteristics of Saussurea salsa in response to flooding in salt marshes of Xiao Sugan Lake, Gansu, China. Chinese Journal of Plant Ecology, 2019, 43(8): 685-696. DOI: 10.17521/cjpe.2019.0132

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https://www.plant-ecology.com/CN/Y2019/V43/I8/685

图/表 9

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参数 Parameter	缩写 Abbreviation	单位 Unit
土壤含水量 Soil moisture content	SMC	%
土壤电导率 Soil electrical conductivity	EC	ms·cm^-1
地上生物量 Aboveground biomass	AB	g·m^-2
平均高度 Average height	AH	cm
密度 Density	D	株·m^-2
叶面积 Leaf area	LA	cm²
叶厚度 Leaf thickness	LT	mm
比叶面积 Specific leaf area	SLA	cm²·g^-1
叶干质量 Leaf dry mass	LDW	g
叶绿素a含量 Chlorophyll a content	C_a	mg·g^-1
叶绿素b含量 Chlorophyll b content	C_b	mg·g^-1
类胡萝卜素含量 Carotenoid content	C_car	mg·g^-1
净光合速率 Net photosynthetic rate	P_n	μmol CO₂·m^-2·s^-1
蒸腾速率 Transpiration rate	T_r	mmol H₂O·m^-2·s^-1
水分利用效率 Water use efficiency	WUE	μmol CO₂·mmol^-1 H₂O
实际光合效率 The actual photochemical efficiency of PSII	Y(II)	无纲量 No dimension
非调节性能量耗散的量子产量 The quantum yield of non-regulated energy dissipation	Y(NO)	无纲量 No dimension
调节性能量耗散的量子产额 The quantum yield of regulated energy dissipation	Y(NPQ)	无纲量 No dimension
光化学淬灭系数 Photochemical quenching	QP	无纲量 No dimension
非光化学猝灭系数 Non- Photochemical quenching	NPQ	无纲量 No dimension
电子传递速率 Electron transport rate	ETR	µmol·m^-2·s^-1

参数 Parameter	缩写 Abbreviation	单位 Unit
土壤含水量 Soil moisture content	SMC	%
土壤电导率 Soil electrical conductivity	EC	ms·cm^-1
地上生物量 Aboveground biomass	AB	g·m^-2
平均高度 Average height	AH	cm
密度 Density	D	株·m^-2
叶面积 Leaf area	LA	cm²
叶厚度 Leaf thickness	LT	mm
比叶面积 Specific leaf area	SLA	cm²·g^-1
叶干质量 Leaf dry mass	LDW	g
叶绿素a含量 Chlorophyll a content	C_a	mg·g^-1
叶绿素b含量 Chlorophyll b content	C_b	mg·g^-1
类胡萝卜素含量 Carotenoid content	C_car	mg·g^-1
净光合速率 Net photosynthetic rate	P_n	μmol CO₂·m^-2·s^-1
蒸腾速率 Transpiration rate	T_r	mmol H₂O·m^-2·s^-1
水分利用效率 Water use efficiency	WUE	μmol CO₂·mmol^-1 H₂O
实际光合效率 The actual photochemical efficiency of PSII	Y(II)	无纲量 No dimension
非调节性能量耗散的量子产量 The quantum yield of non-regulated energy dissipation	Y(NO)	无纲量 No dimension
调节性能量耗散的量子产额 The quantum yield of regulated energy dissipation	Y(NPQ)	无纲量 No dimension
光化学淬灭系数 Photochemical quenching	QP	无纲量 No dimension
非光化学猝灭系数 Non- Photochemical quenching	NPQ	无纲量 No dimension
电子传递速率 Electron transport rate	ETR	µmol·m^-2·s^-1

样地 Plot	群落特征 Community characteristics				盐地风毛菊 Saussurea salsa
样地 Plot	SMC (%)	EC (ms·cm^-1)	AH (cm)	AB (g·m^-2)	AH (cm)	D (株·m^-2)
I	31.79 ± 1.59^c	9.94 ± 0.50^a	6.16 ± 0.31^c	165.30 ± 8.27^a	7.48 ± 0.37^b	59 ± 2.95^c
II	39.12 ± 1.96^b	7.52 ± 0.30^b	9.73 ± 0.49^b	121.66 ± 6.08^b	13.8 ± 0.69^b	74 ± 3.70^a
III	59.63 ± 2.98^a	3.85 ± 0.19^c	30.88 ± 1.54^a	95.93 ± 4.80^c	22.8 ± 1.14^a	62 ± 3.10^b

样地 Plot	群落特征 Community characteristics				盐地风毛菊 Saussurea salsa
样地 Plot	SMC (%)	EC (ms·cm^-1)	AH (cm)	AB (g·m^-2)	AH (cm)	D (株·m^-2)
I	31.79 ± 1.59^c	9.94 ± 0.50^a	6.16 ± 0.31^c	165.30 ± 8.27^a	7.48 ± 0.37^b	59 ± 2.95^c
II	39.12 ± 1.96^b	7.52 ± 0.30^b	9.73 ± 0.49^b	121.66 ± 6.08^b	13.8 ± 0.69^b	74 ± 3.70^a
III	59.63 ± 2.98^a	3.85 ± 0.19^c	30.88 ± 1.54^a	95.93 ± 4.80^c	22.8 ± 1.14^a	62 ± 3.10^b

参数 Parameter	样地 Plot
参数 Parameter	I	II	III
LA (cm²)	7.14 ± 0.36^c	8.34 ± 0.42^b	17.61 ± 0.88^a
LT (mm)	0.15 ± 0.01^a	0.14 ± 0.01^a	0.10 ± 0.01^b
SLA (cm²·g^-1)	8.28 ± 0.41^b	8.82 ± 0.44^b	14.53 ± 0.73^a
LDW (g)	0.86 ± 0.04^a	0.95 ± 0.05^a	1.21 ± 0.06^b
C_a (mg·g^-1)	5.39 ± 0.27^a	3.11 ± 0.16^b	1.26 ± 0.06^c
C_b (mg·g^-1)	1.93 ± 0.10^a	1.15 ± 0.06^b	0.67 ± 0.03^c
C_car (mg·g^-1)	1.39 ± 0.07^a	0.67 ± 0.03^b	0.58 ± 0.03^c
P_n (μmol CO₂·m^-2·s^-1)	4.67 ± 0.23^b	5.17 ± 0.26^a	3.77 ± 0.19^c
T_r (mmol H₂O·m^-2·s^-1)	1.08 ± 0.05^a	1.19 ± 0.06^a	0.97 ± 0.05^b
WUE (μmol CO₂·mmol^-1 H₂O)	4.32 ± 0.22^a	4.34 ± 0.22^a	3.89 ± 0.19^b
Y(II)	0.33 ± 0.02^a	0.33 ± 0.02^a	0.23 ± 0.01^b
Y(NO)	0.38 ± 0.02^a	0.26 ± 0.01^b	0.23 ± 0.01^c
Y(NPQ)	0.29 ± 0.01^c	0.41 ± 0.02^b	0.54 ± 0.03^a
QP	0.73 ± 0.04^a	0.75 ± 0.04^a	0.52 ± 0.03^b
NPQ	0.19 ± 0.01^c	0.40 ± 0.02^b	0.59 ± 0.03^a
ETR (µmol·m^-2·s^-1)	60.02 ± 3.00^a	51.94 ± 2.60^b	49.02 ± 2.45^c

甘肃小苏干湖盐沼湿地盐地风毛菊叶形态-光合生理特征对淹水的响应

Morphological and photosynthetic physiological characteristics of Saussurea salsa in response to flooding in salt marshes of Xiao Sugan Lake, Gansu, China

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统计量 Statistic		轴1 Axis 1	轴2 Axis 2	总方差 Total variance
土壤含水量 Soil moisture content		0.936 6	0.038 9	1.000 0
土壤电导率 Soil electrical conductivity		-0.963 2	-0.052 4
静水持留时间 Water-logging duration		0.910 3	0.198 8
特征值 Eigenvalues		0.533 0	0.050 0
功能性状与环境的相关性 Function traits-environment correlations		0.977 0	0.565 0
累积百分比方差 Explained variation (cumulative)	功能性状数据 Function traits data	55.300 0	60.200 0
累积百分比方差 Explained variation (cumulative)	功能性状-环境关系 Relationship between function traits and environment relation	89.700 0	97.800 0
总特征值 All eigenvalues				1.000 0
总典范特征值 Canonical eigenvalues				0.616 0

功能性状 Functional traits	总离差平方和 SST	残差均方 MSE	未考虑协变量的均方 MS₁	F₁ (MS₁/MSE)	显著性水平p₁	判定系数R₁²	考虑协变量的均方 MS₂	F₂ (MS₂/MSE)	显著性水平 p₂	判定系数R₂²
SLA	6.311 1	2.129 5	3.155 6	1.481 9	0.266 0	0.852 0	29.425 5	13.818 3	0.000 1	0.790 0
C_a	4.432 7	0.572 2	2.216 3	3.873 3	0.050 4	0.889 0	10.950 6	19.137 5	0.000 0	0.842 0
C_b	0.349 8	0.080 4	0.174 9	2.175 4	0.156 3	0.854 0	1.132 9	14.090 5	0.000 1	0.879 0
Y(II)	0.004 6	0.000 6	0.002 3	4.174 3	0.042 1	0.877 0	0.009 4	17.093 1	0.000 0	0.826 0
Y(NPQ)	0.008 1	0.000 4	0.004 1	10.662 2	0.002 2	0.977 0	0.039 1	102.532 8	0.000 0	0.968 0
Y(NO)	0.016 0	0.000 1	0.008 0	136.047 6	0.000 0	0.991 0	0.015 2	259.079 1	0.000 0	0.987 0
QP	0.024 7	0.003 4	0.012 4	3.592 8	0.059 9	0.832 0	0.041 0	11.919 5	0.000 3	0.763 0
NPQ	0.023 4	0.000 5	0.011 7	22.825 4	0.000 1	0.987 0	0.093 1	181.178 3	0.000 0	0.981 0

	C_a	C_b	SLA	Y(II)	Y(NPQ)	Y(NO)	NPQ	QP
C_a	1.00
C_b	1.00^**	1.00
SLA	-0.14	-0.18	1.00
Y(II)	0.06	0.10	-1.00^**	1.00
Y(NPQ)	-0.53	-0.57	0.91^**	-0.88^*	1.00
Y(NO)	0.79^*	0.82^*	-0.71	0.65	-0.94^**	1.00
NPQ	-0.58	-0.61	0.89^*	-0.85^*	1.00^**	-0.95^**	1.00
QP	-0.02	0.02	-0.99^**	1.00^**	-0.84^*	0.59	-0.81^*	1.00

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