淹水和干旱生境下铅对芦苇生长、生物量分配和光合作用的影响

doi:10.17521/cjpe.2017.0218

植物生态学报 ›› 2018, Vol. 42 ›› Issue (2): 229-239.DOI: 10.17521/cjpe.2017.0218

淹水和干旱生境下铅对芦苇生长、生物量分配和光合作用的影响

张娜^1,^2,³,朱阳春¹,李志强⁵,卢信¹,范如芹¹,刘丽珠¹,童非¹,陈静³,穆春生^4,^*(),张振华^1,^*>()

¹江苏省农业科学院农业资源与环境研究所, 南京 210014
²江苏省农业科学院江苏省食品质量安全重点实验室, 南京 210014
³淮阴工学院江苏省凹土资源利用重点实验室, 江苏淮安 223003
⁴东北师范大学草地科学研究所植被生态教育部重点实验室, 长春 130024
⁵南京信息工程大学公共管理学院/气候变化与公共政策研究院, 南京 210044

出版日期:2018-02-20 发布日期:2018-04-16
通讯作者: 穆春生,张振华
基金资助:
中国博士后科学基金(2017M621670);国家重点基础研究发展计划973计划(2017M621670);江苏省农业科技自主创新资金(CX(16)1051)

Effect of Pb pollution on the growth, biomass allocation and photosynthesis of Phragmites australis in flood and drought environment

ZHANG Na^1,^2,³,ZHU Yang-Chun¹,LI Zhi-Qiang⁵,LU Xin¹,FAN Ru-Qin¹,LIU Li-Zhu¹,TONG Fei¹,CHEN Jing³,MU Chun-Sheng^4,^*(),ZHANG Zhen-Hua^1,^*()

¹Institute of Agricultural Resource and Environment, Jiangsu Academy of Agricultural Sciences, Nanjing 210014, China

²Key Laboratory of Food Quality and Safety of Jiangsu Province, Jiangsu Academy of Agricultural Sciences, Nanjing 210014, China

³Key Laboratory for Palygorskite Science and Applied Technology of Jiangsu Province, Huaiyin Institute of Technology, Huai’an, Jiangsu 223003, China;

⁴Key Laboratory of Vegetation Ecology, Ministry of Education, Institute of Grassland Science, Northeast Normal University, Changchun 130024, China

⁵Institution of Climate Change and Public Polices, School of Public Administration, Nanjing University of Information Science and Technology, Nanjing 210044, China

Online:2018-02-20 Published:2018-04-16
Contact: Chun-Sheng MU,Zhen-Hua ZHANG
Supported by:
Supported by the China Postdoctoral Science Foundation.(2017M621670);the National Basic Research Program of China.(2017M621670);the Agricultural Science and Technology Independent Innovation Fund of Jiangsu Province.(CX(16)1051)

摘要/Abstract

摘要：

芦苇(Phragmites australis)作为典型的根茎型多年生湿地植物, 具有广泛的环境耐受性。该研究采用盆栽实验, 采取裂区实验设计, 水分处理为主区, 包括淹水和干旱两个水平, 铅(Pb)为副区, 包括0、500、1500、3β000、4β500 mg·kg^-1 5个水平, 共10个处理, 每个处理12个重复, 研究淹水和干旱条件下Pb污染对芦苇生长、生物量分配及光合作用的影响, 以期明确不同生境下芦苇适应或忍耐重金属污染而采取的策略, 为芦苇应用于湿地恢复和污染修复提供理论依据。结果表明, 在淹水处理中, Pb显著抑制地下芽形成和根茎生长, 但对子株数没有影响; 与母株相比子株具有高的日生长速率、光合速率和生物量(母株的3-7倍)。在干旱环境中, Pb显著抑制根、地下芽和根茎生长, 母株和子株生物量积累及光合作用, 且这些指标均小于淹水处理的。无论在淹水还是干旱环境中, 芦苇体内绝大部分Pb积累在根中, 根茎和子株中Pb含量较少, 被转运至母株中的Pb大约是子株的3倍。淹水条件下子株体内Pb含量小于干旱处理的。结果表明, 干旱和Pb的协同作用显著抑制芦苇生长、生物量积累和光合作用, 可能导致子株生产力和种群密度减小甚至种群衰退。但淹水芦苇能够采取相应的Pb分配策略减缓Pb污染对芦苇生长、生理和繁殖的负面影响, 有利于芦苇种群的繁衍和稳定。

关键词: 铅污染, 水分, 生长, 生物量分配, 光合作用

Abstract:

Aims Reed (Phragmites australis) is a typical perennial rhizomatic plant with extensive tolerance to environmental stress. In order to better understand the adaptation and tolerance of reeds subjected to heavy metal pollution in different levels of water, we conducted a study on the effects of Pb pollution on growth, biomass and photosynthesis of reeds in flood and drought environment. This research would provide theoretical basis for application of reeds in wetland restoration and remediation.

Methods We conducted a pot experiment with destructive sampling after 90 days of growth. The water treatments were main plot, including two water levels. The Pb treatments were secondary plot (nested within water treatments), including five levels (0, 500, 1 500, 3 000, 4 500 mg·kg^-1). There were 10 treatments with 12 replicates per treatment.

Important findings In the flood environment, Pb pollution significantly inhibited the growth of buds and rhizomes, but had no significant effect on the number of offspring shoots. The offspring shoots had higher growth rate per day, net photosynthetic rate and biomass compared to the parent shoots. In the drought environment, Pb pollution inhibited the growth of roots, buds and rhizomes, and biomass accumulation of parent and offspring shoots as well as photosynthetic parameters. These parameters were lower under the drought condition than in the flood environment. The Pb was mostly concentrated in roots compared to rhizomes and offspring shoots. In both flood and drought environments, the concentration of Pb in parent shoots was about three times of that in offspring shoots. The Pb concentration in offspring shoots under the flood condition was less than that in the drought environment. Overall, these results indicated that the synergistic effect of Pb and drought significantly inhibited the growth, biomass accumulation and photosynthesis of reeds, which might result in reduced offspring productivity and population density and may lead to population decline. However, the flooded reeds could adopt some strategies of Pb allocation to alleviate the negative effect of Pb on the growth, physiology and clonal propagation, benefiting the population reproduction and stabilization.

Key words: Pb pollution, water, growth, biomass allocation, photosynthesis

张娜, 朱阳春, 李志强, 卢信, 范如芹, 刘丽珠, 童非, 陈静, 穆春生, 张振华. 淹水和干旱生境下铅对芦苇生长、生物量分配和光合作用的影响. 植物生态学报, 2018, 42(2): 229-239. DOI: 10.17521/cjpe.2017.0218

ZHANG Na, ZHU Yang-Chun, LI Zhi-Qiang, LU Xin, FAN Ru-Qin, LIU Li-ZhuTONG , Fei, CHEN Jing, MU Chun-Sheng, ZHANG Zhen-Hua. Effect of Pb pollution on the growth, biomass allocation and photosynthesis of Phragmites australis in flood and drought environment. Chinese Journal of Plant Ecology, 2018, 42(2): 229-239. DOI: 10.17521/cjpe.2017.0218

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https://www.plant-ecology.com/CN/Y2018/V42/I2/229

图/表 12

参考文献 26

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	重复测量分析 Repeated measure (p)
	铅 Pb	水 Water	铅 × 水 Pb × water		时间 Time		时间 × 铅 Time × Pb		时间 × 水 Time × Water		时间 × 铅 × 水 Time × Pb × Water
母株生长 Growth of parent shoot
茎长 Stem length	< 0.001	< 0.001	< 0.01		< 0.001		< 0.01		< 0.001		< 0.001
叶片死亡数 No. of dead leaves	0.366	< 0.001	< 0.05		< 0.001		< 0.001		< 0.001		< 0.05
子株生长 Growth of offspring shoot
茎长 Stem length	< 0.001	< 0.001	< 0.001		< 0.001		< 0.001		< 0.001		< 0.001
叶片死亡数 No. of dead leaves	0.168	< 0.001	0.913		< 0.001		< 0.05		< 0.001		0.742
生长指标 Growth parameters		双因素方差分析 Two-way AVONA
		铅 Pb				水 Water				铅×水 Pb × water
		F		p		F		p		F		p
母株叶片死亡率 Leaf mortality in parent shoots		2.496		0.054		2.461		0.127		0.932		0.459
子株叶片死亡率 Leaf mortality in offspring shoots		7.086		< 0.001		2.185		0.150		4.353		< 0.01

	重复测量分析 Repeated measure (p)
	铅 Pb	水 Water	铅 × 水 Pb × water		时间 Time		时间 × 铅 Time × Pb		时间 × 水 Time × Water		时间 × 铅 × 水 Time × Pb × Water
母株生长 Growth of parent shoot
茎长 Stem length	< 0.001	< 0.001	< 0.01		< 0.001		< 0.01		< 0.001		< 0.001
叶片死亡数 No. of dead leaves	0.366	< 0.001	< 0.05		< 0.001		< 0.001		< 0.001		< 0.05
子株生长 Growth of offspring shoot
茎长 Stem length	< 0.001	< 0.001	< 0.001		< 0.001		< 0.001		< 0.001		< 0.001
叶片死亡数 No. of dead leaves	0.168	< 0.001	0.913		< 0.001		< 0.05		< 0.001		0.742
生长指标 Growth parameters		双因素方差分析 Two-way AVONA
		铅 Pb				水 Water				铅×水 Pb × water
		F		p		F		p		F		p
母株叶片死亡率 Leaf mortality in parent shoots		2.496		0.054		2.461		0.127		0.932		0.459
子株叶片死亡率 Leaf mortality in offspring shoots		7.086		< 0.001		2.185		0.150		4.353		< 0.01

水分 Water level	处理时间 Treatment time (d)	指标 Parameter	铅处理浓度 Pb concentration of treatments (mg·kg^-1)
水分 Water level	处理时间 Treatment time (d)	指标 Parameter	对照 Control	500	1 500	3 000	4 500
淹水 Flood	30	PSL (cm)	18.13 ± 0.81^a	16.64 ± 0.75^ab	15.54 ± 0.40^b	13.10 ± 0.52c	13.24 ± 0.29^c
		PDLN (No.)	1.28 ± 0.05^b	1.54 ± 0.12^ab	1.73 ± 0.11^a	1.90 ± 0.06^a	1.84 ± 0.13^a
		PGR (cm·d^-1)	0.44 ± 0.03^a	0.39 ± 0.02^ab	0.35 ± 0.01^b	0.27 ± 0.02^c	0.28 ± 0.01^c
	60	PSL (cm)	23.86 ± 0.83^a	22.34 ± 1.52^ab	22.71 ± 0.72^ab	19.22 ± 1.04^b	18.17 ± 0.54^b
		PDLN (No.)	3.57 ± 0.15^b	3.65 ± 0.22^b	4.38 ± 0.14^a	4.50 ± 0.24^a	4.56 ± 0.08^a
		PGR (cm·d^-1)	0.19 ± 0.02^a	0.19 ± 0.03^a	0.24 ± 0.01^a	0.21 ± 0.04^a	0.16 ± 0.02^a
	90	PSL (cm)	26.00 ± 0.96^a	24.03 ± 1.84^a	24.21 ± 0.97^a	22.80 ± 0.57^a	21.68 ± 1.20^a
		PDLN (No.)	7.02 ± 0.06^a	6.59 ± 0.43^a	6.61 ± 0.22^a	6.80 ± 0.16^a	6.43 ± 0.15^a
		PGR (cm·d^-1)	0.07 ± 0.03^a	0.06 ± 0.01^a	0.05 ± 0.01^a	0.12 ± 0.04^a	0.12 ± 0.03^a
干旱 Drought	30	PSL (cm)	12.17 ± 0.88^a*	11.90 ± 0.73^a*	9.30 ± 0.37^b*	7.87 ± 0.36^b*	7.72 ± 0.42^b*
		PDLN (No.)	1.64 ± 0.06^c*	2.08 ± 0.15^b*	2.13 ± 0.05^ab*	2.18 ± 0.10^ab*	2.42 ± 0.17^a*
		PGR (cm·d^-1)	0.24 ± 0.03^a*	0.23 ± 0.02^a*	0.15 ± 0.01^b*	0.10 ± 0.01^b*	0.09 ± 0.01^b*
	60	PSL (cm)	17.30 ± 0.54^a*	15.06 ± 0.89^b*	11.15 ± 0.57^c*	9.09 ± 0.42^d*	7.58 ± 0.61^d*
		PDLN (No.)	2.60 ± 0.04^b	3.19 ± 0.27^a*	3.33 ± 0.12^a*	3.51 ± 0.17^a*	3.65 ± 0.15^a*
		PGR (cm·d^-)	0.17 ± 0.03^a	0.11 ± 0.01^b*	0.06 ± 0.01^bc*	0.04 ± 0.00^c*	0.02 ± 0.01^c*
	90	PSL (cm)	20.29 ± 0.94^a*	18.14 ± 0.74^a*	13.61 ± 0.94^b*	9.43 ± 0.10^c*	8.31 ± 0.53^c*
		PDLN (No.)	7.32 ± 0.16^a	5.46 ± 0.15^b*	4.82 ± 0.13^bc*	4.73 ± 0.34^bc*	4.08 ± 0.36^c*
		PGR (cm·d^-1)	0.10 ± 0.02^a	0.10 ± 0.03^a	0.08 ± 0.03^ab	0.02 ± 0.01^b	0.03 ± 0.01^b*

水分 Water level	处理时间 Treatment time (d)	指标 Parameter	铅处理浓度 Pb concentration of treatments (mg·kg^-1)
水分 Water level	处理时间 Treatment time (d)	指标 Parameter	对照 Control	500	1 500	3 000	4 500
淹水 Flood	30	PSL (cm)	18.13 ± 0.81^a	16.64 ± 0.75^ab	15.54 ± 0.40^b	13.10 ± 0.52c	13.24 ± 0.29^c
		PDLN (No.)	1.28 ± 0.05^b	1.54 ± 0.12^ab	1.73 ± 0.11^a	1.90 ± 0.06^a	1.84 ± 0.13^a
		PGR (cm·d^-1)	0.44 ± 0.03^a	0.39 ± 0.02^ab	0.35 ± 0.01^b	0.27 ± 0.02^c	0.28 ± 0.01^c
	60	PSL (cm)	23.86 ± 0.83^a	22.34 ± 1.52^ab	22.71 ± 0.72^ab	19.22 ± 1.04^b	18.17 ± 0.54^b
		PDLN (No.)	3.57 ± 0.15^b	3.65 ± 0.22^b	4.38 ± 0.14^a	4.50 ± 0.24^a	4.56 ± 0.08^a
		PGR (cm·d^-1)	0.19 ± 0.02^a	0.19 ± 0.03^a	0.24 ± 0.01^a	0.21 ± 0.04^a	0.16 ± 0.02^a
	90	PSL (cm)	26.00 ± 0.96^a	24.03 ± 1.84^a	24.21 ± 0.97^a	22.80 ± 0.57^a	21.68 ± 1.20^a
		PDLN (No.)	7.02 ± 0.06^a	6.59 ± 0.43^a	6.61 ± 0.22^a	6.80 ± 0.16^a	6.43 ± 0.15^a
		PGR (cm·d^-1)	0.07 ± 0.03^a	0.06 ± 0.01^a	0.05 ± 0.01^a	0.12 ± 0.04^a	0.12 ± 0.03^a
干旱 Drought	30	PSL (cm)	12.17 ± 0.88^a*	11.90 ± 0.73^a*	9.30 ± 0.37^b*	7.87 ± 0.36^b*	7.72 ± 0.42^b*
		PDLN (No.)	1.64 ± 0.06^c*	2.08 ± 0.15^b*	2.13 ± 0.05^ab*	2.18 ± 0.10^ab*	2.42 ± 0.17^a*
		PGR (cm·d^-1)	0.24 ± 0.03^a*	0.23 ± 0.02^a*	0.15 ± 0.01^b*	0.10 ± 0.01^b*	0.09 ± 0.01^b*
	60	PSL (cm)	17.30 ± 0.54^a*	15.06 ± 0.89^b*	11.15 ± 0.57^c*	9.09 ± 0.42^d*	7.58 ± 0.61^d*
		PDLN (No.)	2.60 ± 0.04^b	3.19 ± 0.27^a*	3.33 ± 0.12^a*	3.51 ± 0.17^a*	3.65 ± 0.15^a*
		PGR (cm·d^-)	0.17 ± 0.03^a	0.11 ± 0.01^b*	0.06 ± 0.01^bc*	0.04 ± 0.00^c*	0.02 ± 0.01^c*
	90	PSL (cm)	20.29 ± 0.94^a*	18.14 ± 0.74^a*	13.61 ± 0.94^b*	9.43 ± 0.10^c*	8.31 ± 0.53^c*
		PDLN (No.)	7.32 ± 0.16^a	5.46 ± 0.15^b*	4.82 ± 0.13^bc*	4.73 ± 0.34^bc*	4.08 ± 0.36^c*
		PGR (cm·d^-1)	0.10 ± 0.02^a	0.10 ± 0.03^a	0.08 ± 0.03^ab	0.02 ± 0.01^b	0.03 ± 0.01^b*

水分 Water level	处理时间 Treatment time (d)	指标 Parameter	铅处理浓度 Pb concentration of treatments (mg·kg^-1)
水分 Water level	处理时间 Treatment time (d)	指标 Parameter	对照 Control	500	1 500	3 000	4 500
淹水 Flood	30	OSL (cm)	15.08 ± 0.88^a	13.20 ± 0.99^ab	11.28 ± 0.95^b	8.42 ± 0.26^c	7.92 ± 0.35^c
		ODLN (No.)	0.27 ± 0.09^a	0.28 ± 0.05^a	0.06 ± 0.03^b	0.06 ± 0.03^b	0.00 ± 0.00^b
		OGR (cm·d^-1)	0.76 ± 0.05^a	0.66 ± 0.05^ab	0.57 ± 0.05^b	0.42 ± 0.01^c	0.40 ± 0.02^c
	60	OSL (cm)	31.06 ± 0.37^a	30.08 ± 0.42^a	28.53 ± 0.77^a	20.12 ± 0.73^b	15.24 ± 0.83^c
		ODLN (No.)	1.49 ± 0.10^a	1.50 ± 0.17^a	1.74 ± 0.34^a	1.82 ± 0.20^a	1.96 ± 0.22^a
		OGR (cm·d^-1)	0.53 ± 0.03^a	0.57 ± 0.02^a	0.58 ± 0.02^a	0.39 ± 0.03^b	0.24 ± 0.03^c
	90	OSL (cm)	41.22 ± 1.14^a	40.97 ± 1.19^a	40.65 ± 1.68^a	32.98 ± 1.78^b	22.41 ± 0.74^c
		ODLN (No.)	3.28 ± 0.15^a	2.99 ± 0.27^ab	2.95 ± 0.22^ab	3.10 ± 0.20^ab	2.53 ± 0.35^b
		OGR (cm·d^-1)	0.34 ± 0.05^ab	0.36 ± 0.03^a	0.40 ± 0.04^a	0.43 ± 0.04^a	0.24 ± 0.03^b
干旱Drought	30	OSL (cm)	4.50 ± 0.65^a*	3.38 ± 0.63^a*	3.38 ± 0.55^a*	3.50 ± 0.29^a*	1.50 ± 0.05^b*
		ODLN (No.)	0.00 ± 0.00^a*	0.00 ± 0.00^a*	0.00 ± 0.00^a	0.00 ± 0.00^a	0.00 ± 0.00^a
		OGR (cm·d^-1)	0.23 ± 0.03^a*	0.17 ± 0.03^a*	0.17 ± 0.03^a*	0.18 ± 0.01^a*	0.07 ± 0.00^b*
	60	OSL (cm)	19.95 ± 1.08^a*	11.23 ± 0.43^b*	5.55 ± 0.75^c*	4.44 ± 0.87^c*	3.65 ± 0.61^c*
		ODLN (No.)	0.10 ± 0.04^a*	0.14 ± 0.05^a*	0.15 ± 0.05^a*	0.25 ± 0.16^a*	0.06 ± 0.06^a*
		OGR (cm·d^-1)	0.52 ± 0.03^a	0.26 ± 0.03^b*	0.07 ± 0.04^cd*	0.03 ± 0.02^d*	0.14 ± 0.04^c
	90	OSL (cm)	28.82 ± 1.16^a*	24.56 ± 0.76^b*	14.46 ± 1.00^c*	8.45 ± 0.25^d*	4.13 ± 0.12^e*
		ODLN (No.)	2.23 ± 0.28^a*	1.83 ± 0.11^ab*	1.40 ± 0.26^ab*	1.86 ± 0.34^ab*	1.21 ± 0.44^b*
		OGR (cm·d^-1)	0.30 ± 0.06^b	0.44 ± 0.03^a	0.30 ± 0.05^b	0.13 ± 0.03^c*	0.03 ± 0.02^c*

淹水和干旱生境下铅对芦苇生长、生物量分配和光合作用的影响

Effect of Pb pollution on the growth, biomass allocation and photosynthesis of Phragmites australis in flood and drought environment

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生长指标 Growth parameters	双因素方差分析 Two-way AVONA
	铅 Pb		水 Water		铅 × 水 Pb × water
	F	p	F	p	F	p
根长 Root length	184.571	< 0.001	273.946	< 0.001	4.605	< 0.01
根茎长 Rhizome length	65.735	< 0.001	126.045	< 0.002	5.271	< 0.01
根茎数 No. of rhizomes	92.408	< 0.001	207.906	< 0.003	8.108	< 0.001
芽数 No. of buds	42.807	< 0.001	51.986	< 0.004	6.879	< 0.001
子株数 No. of offspring shoots	12.658	< 0.001	442.488	< 0.005	4.293	< 0.01

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光合参数 Photosynthetic parameters	双因素方差分析 Two-way AVONA
	铅 Pb		水 Water		铅 × 水 Pb × water
	F	p	F	p	F	p
净光合速率 Net photosynthetic rate	12.69	<0.001	20.38	<0.001	6.92	<0.001
气孔导度 Stomatal conductance	16.90	<0.001	95.36	<0.001	3.80	<0.01
胞间CO₂浓度 Intercellular CO₂ concentration	9.80	<0.001	111.35	<0.001	1.54	0.23
蒸腾速率 Transpiration rate	6.08	<0.001	88.80	<0.001	1.07	0.38

光合参数 Photosynthetic parameters	双因素方差分析 Two-way AVONA
	铅 Pb		水 Water		铅 × 水 Pb × water
	F	p	F	p	F	p
净光合速率 Net photosynthetic rate	12.69	<0.001	20.38	<0.001	6.92	<0.001
气孔导度 Stomatal conductance	16.90	<0.001	95.36	<0.001	3.80	<0.01
胞间CO₂浓度 Intercellular CO₂ concentration	9.80	<0.001	111.35	<0.001	1.54	0.23
蒸腾速率 Transpiration rate	6.08	<0.001	88.80	<0.001	1.07	0.38

	双因素方差分析 Two-way AVONA
	铅 Pb		水 Water		铅×水 Pb × water
	F	p	F	p	F	p
根 Roots	472.06	<0.001	1 061.42	<0.001	166.30	<0.001
根茎 Rhizomes	174.54	<0.001	317.95	<0.001	79.07	<0.001
母株 Parent shoots	3 304.20	<0.001	369.17	<0.001	434.26	<0.001
子株 Offspring shoots	2 392.36	<0.001	42.83	<0.001	31.27	<0.001

	双因素方差分析 Two-way AVONA
	铅 Pb		水 Water		铅×水 Pb × water
	F	p	F	p	F	p
根 Roots	472.06	<0.001	1 061.42	<0.001	166.30	<0.001
根茎 Rhizomes	174.54	<0.001	317.95	<0.001	79.07	<0.001
母株 Parent shoots	3 304.20	<0.001	369.17	<0.001	434.26	<0.001
子株 Offspring shoots	2 392.36	<0.001	42.83	<0.001	31.27	<0.001