氮水平和竞争对互花米草与芦苇叶特征的影响

doi:10.3773/j.issn.1005-264x.2008.02.017

植物生态学报 ›› 2008, Vol. 32 ›› Issue (2): 392-401.DOI: 10.3773/j.issn.1005-264x.2008.02.017

氮水平和竞争对互花米草与芦苇叶特征的影响

赵聪蛟¹, 邓自发¹^,², 周长芳¹, 关保华¹, 安树青¹^,^*(), 陈琳¹, 陆霞梅¹

1 南京大学生命科学学院湿地生态研究所,南京 210093
2 南通大学生命科学学院,江苏南通 226007

收稿日期:2007-01-24 接受日期:2007-07-05 出版日期:2008-01-24 发布日期:2008-03-30
通讯作者: 安树青
作者简介:* E-mail: anshq@nju.edu.cn
第一联系人：
英文摘要部分在修改过程中得到了美国路易斯安那州立大学Irving A. Mendelssohn教授和南京大学外国语学院李长生副教授的帮助,在此一并致谢
基金资助:
国家自然科学基金项目(30400054);教育部跨世纪优秀人才培养计划基金项目(0208200202)

EFFECTS OF NITROGEN AVAILABILITY AND COMPETITION ON LEAF CHARACTERISTICS OF SPARTINA ALTERNIFLORA AND PHRAGMITES AUSTRALIS

ZHAO Cong-Jiao¹, DENG Zi-Fa¹^,², ZHOU Chang-Fang¹, GUAN Bao-Hua¹, AN Shu-Qing¹^,^*(), CHEN Lin¹, LU Xia-Mei¹

¹Institute of Wetland Ecology, School of Life Science, Nanjing University, Nanjing 210093, China
²School of Life Sciences, Nantong University, Nantong, Jiangsu 226007, China

Received:2007-01-24 Accepted:2007-07-05 Online:2008-01-24 Published:2008-03-30
Contact: AN Shu-Qing

摘要/Abstract

摘要：

互花米草 (Spartina alterniflora) 和芦苇 (Phragmites australis) 是滨海盐沼湿地的多年生草本植物,从世界范围来看,它们二者具有区域性的相互入侵特征,因此研究生境条件对两物种互侵机制的影响是一个十分有意义的生态学命题。该文运用随机区组实验设计方法,模拟海滩环境、构建人工种群、控制可变因子,研究了外来种互花米草与本地种芦苇分别单种和混种时,叶特征对不同氮水平、不同植株密度的响应。结果表明:随着氮水平的升高,互花米草和芦苇的叶面积无论是在单种还是混种情况下都显著增加 (p<0.05),但混种条件下芦苇的叶面积在高氮水平下增幅减少,这与高氮状况下互花米草与芦苇的竞争加剧有关;氮水平对单种中两种植物的叶数影响最显著 (p<0.01),对混种中互花米草的叶数和芦苇的叶宽影响最大 (p<0.05)。植株密度增加导致种内和种间竞争加剧,无论在单种还是混种处理下,都造成两种植物叶面积的显著减少 (p<0.05)。单种处理中,两物种的叶数受密度的响应最显著 (p<0.05);而混种处理中芦苇对互花米草的竞争显著减小了互花米草的叶宽和叶数(p<0.05),互花米草对芦苇的竞争则显著减小了芦苇的叶长、叶宽和叶数 (p<0.05)。两种植物的竞争结果受到氮营养的调控,低、高氮水平下互花米草的种间竞争能力大于芦苇,中氮水平下则是芦苇的种间竞争能力大于互花米草。高氮水平下互花米草通过叶面积的快速增加抑制了芦苇的叶生长,使其叶面积减少,从而在竞争中占据优势,这可能是互花米草入侵我国海滩芦苇种群的机制之一。

关键词: 氮水平, 植株密度, 竞争, 互花米草, 芦苇, 叶形态

Abstract:

Aims Spartina alterniflora, originating from North America, has become an invasive species in Europe and China. Meanwhile, Phragmites australis, a species experiencing 'die-back' in Europe, has invaded coastal ecosystems in North America. Each species is invading the other's native habitat. We studied changes of leaf characters for the two species under different nitrogen and planting densities in the greenhouse to 1) compare the relative competitiveness and invasive capacity of the two species and 2) reveal potential mechanisms that determine successful invasion in different regions.
Methods We grew artificial populations of Spartina alterniflora (S) and Phragmites australis (P) at three different densities in monoculture (S, SS, SSS and P, PP, PPP) and mixed-culture (SP, SPP and PSS), and under three levels of nitrogen (0, 60 and 120 mg·kg^-1). Plants were harvested after 15 weeks, and their leaf characteristics, including area, length, width, thickness and number were measured.
Important findings Nitrogen addition increased leaf area in both species whether in monoculture or mixed-culture (p<0.05), but the change in leaf area of P. australis in mixed-culture decreased with high nitrogen level, which may be due to greater interspecific competition from S. alterniflora. In monoculture, the effects of nitrogen addition on leaf number were greater than on the other leaf traits (p<0.01), while the effects on leaf number (S. alterniflora) or leaf width (P. australis) were greatest (p<0.05) in mixed-culture. Plant densities decreased leaf area of the two species in all treatments (p<0.05). In monoculture, the effects of plant densities on leaf number were greatest (p<0.05). However, in mixed-culture, P. australis mainly reduced leaf width and leaf number of S. alterniflora (p<0.05), while S. alterniflora reduced all the parameters of P. australis (p<0.05). The intensity of competition which S. alterniflora imposed on P. australis was greater than the reverse with low and high nitrogen levels, but this outcome was reversed with medium nitrogen level. At high nitrogen levels, S. alterniflora dominated interspecific competition, with its increased leaf area restraining the leaf growth and reducing the leaf area of P. australis; this may be a mechanism for the successful invasion of S. alterniflora into P. australis populations.

Key words: nitrogen level, plant density, competition, Spartina alterniflora, Phragmites australis, leaf morphology

赵聪蛟, 邓自发, 周长芳, 关保华, 安树青, 陈琳, 陆霞梅. 氮水平和竞争对互花米草与芦苇叶特征的影响. 植物生态学报, 2008, 32(2): 392-401. DOI: 10.3773/j.issn.1005-264x.2008.02.017

ZHAO Cong-Jiao, DENG Zi-Fa, ZHOU Chang-Fang, GUAN Bao-Hua, AN Shu-Qing, CHEN Lin, LU Xia-Mei. EFFECTS OF NITROGEN AVAILABILITY AND COMPETITION ON LEAF CHARACTERISTICS OF SPARTINA ALTERNIFLORA AND PHRAGMITES AUSTRALIS. Chinese Journal of Plant Ecology, 2008, 32(2): 392-401. DOI: 10.3773/j.issn.1005-264x.2008.02.017

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图/表 9

表1 每个氮水平下单种种群和混合种群中的物种组合与密度设计

Table 1 Species combination and density series at each nitrogen level

种植方式 Planting treatment	物种组合 Species combination	植株密度 Plant density in each pot
种植方式 Planting treatment	物种组合 Species combination	1株 One individual	2株 Two individuals	3株 Three individuals
单种系列	S+S (SS)	S+0	S+S	S+S+S
Monoculture	P+P (PP)	P+0	P+P	P+P+P
混种系列	S+P (SP)	S+0	S+P	S+P+P
Mixed-culture	P+S (PS)	P+0	P+S	P+S+S

表2 互花米草、芦苇单种种群叶特征二维方差分析

Table 2 Results of Two-way ANOVA of leaf characters for Spartina alterniflora and Phragmites australis in monoculture

叶特征 Leaf characters	变异来源 Variation source	互花米草 S. alterniflora			芦苇 P. australis
叶特征 Leaf characters	变异来源 Variation source	F	p		F	p
叶长	D	1.831 4		0.176 1	10.657 6	0.000 3^**
Leaf length	N	11.898 4		0.000 1^**	22.768 6	0.000 0^**
	D × N	1.287 1		0.295 0	0.965 6	0.439 3
叶宽	D	10.615 5		0.000 3^**	11.321 4	0.000 2^**
Leaf width	N	24.700 2		0.000 0^**	18.568 5	0.000 0^**
	D × N	1.454 3		0.238 4	2.105 1	0.102 4
叶片厚度	D	2.305 2		0.115 6	0.624 8	0.541 6
Leaf thickness	N	0.019 7		0.980 5	3.561 3	0.039 8^*
	D × N	2.211 2		0.089 2	0.952 6	0.446 3
叶数	D	39.405 2		0.000 0^**	7.307 5	0.002 4^**
Leaf number	N	56.688 5		0.000 0^**	4.321 8	0.021 5^*
	D × N	3.132 6		0.027 4^*	0.859 3	0.498 4
叶面积	D	152.858 6		0.000 0^**	77.101 2	0.000 0^**
Leaf area	N	142.586 1		0.000 0^**	38.637 7	0.000 0^**
	D × N	8.408 8		0.000 0^**	3.634 6	0.014 0^*

图1 互花米草 (a)、芦苇 (b) 单种种群中单株植物的叶面积变化 N0、N1、N2: 低、中、高3个氮水平N0 (0 mg·kg-1), N1 (60 mg·kg-1), N2 (120 mg·kg-1) three nitrogen levels S、SS、SSS: 不同密度的互花米草Different density series of S. alterniflora P、PP、PPP: 不同密度的芦苇Different density series of P. australis 1个字母代表1株植物 Each letter corresponds to 1 individual plant of that species e, f, g: 表示相同氮水平的不同植株密度之间差异显著 (p<0.05) Indicating significantly density effect differences at the same nitrogen level (p<0.05) E, F, G: 表示相同植株密度的不同氮水平之间差异显著 (p<0.05) Indicating significant nitrogen effect differences in the same density (p<0.05)

Fig.1 Leaf area changes of individual plant of Spartina alterniflora (a) and Phragmites australis (b) in monoculture

图2 互花米草单种种群中单株植物的叶长 (a)、叶宽 (b) 与叶数 (c) 的变化图注同图1

Fig. 2 Changes of leaf length (a), leaf width (b) and leaf number (c) of individual Spartina alterniflora in monoculture Notes see Fig. 1

图3 芦苇单种种群中单株植物的叶长 (a)、叶宽 (b)、叶片厚度 (c) 与叶数 (d) 的变化图注同图1

Fig. 3 Changes of leaf length (a), leaf width (b), leaf thickness (c) and leaf number (d) of individual Phragmites australis in monoculture Notes see Fig. 1

表3 互花米草、芦苇混合种群叶特征二维方差分析

Table 3 Results of Two-way ANOVA of leaf characters for Spartina alterniflora and Phragmites australis in mixed-culture

叶特征 Leaf characters	变异来源 Variation source	互花米草 S. alterniflora			芦苇 P. australis
叶特征 Leaf characters	变异来源 Variation source	F	p		F	p
叶长	MT	0.743 9		0.489 3	9.669 1	0.001 4^**
Leaf length	N	1.494 5		0.250 9	10.241 5	0.001 1^**
	MT × N	0.400 6		0.805 6	1.662 6	0.202 3
叶宽	MT	6.822 2		0.006 2^**	16.083 3	0.000 1^**
Leaf width	N	7.755 6		0.003 7^**	9.255 2	0.001 7^**
	MT × N	1.444 4		0.260 2	0.645 8	0.636 9
叶片厚度	MT	1.496 4		0.250 5	1.704 1	0.210 0
Leaf thickness	N	0.899 1		0.424 4	0.216 9	0.807 1
	MT × N	0.424 4		0.789 0	1.197 4	0.345 9
叶数	MT	1.888 4		0.180 1	3.787 3	0.042 4^*
Leaf number	N	9.654 7		0.001 4^**	5.190 6	0.016 6^*
	MT × N	1.460 2		0.255 5	0.711 3	0.594 8
叶面积	MT	77.436 4		0.000 0^**	54.776 8	0.000 0^**
Leaf area	N	102.774 8		0.000 0^**	28.698 9	0.000 0^**
	MT × N	21.881 2		0.000 0^**	5.848 0	0.014 0^*

图4 互花米草与芦苇混合种群中单株植物的叶面积变化 (a) 随芦苇密度的增加互花米草的单株叶面积变化Leaf area changes of individual S. alterniflora under different density series of P. australis (b) 随互花米草密度的增加芦苇的单株叶面积变化Leaf area changes of individual P. australis under different density series of S. alterniflora S, P, SP or PS, SPP, PSS:不同的混种方式Different mixed-culture treatments of S. alterniflora (S) and P. australis (P) 1个字母代表1株植物Each letter corresponds to 1 individual plant of that species e, f, g, E, F, G, N0, N1, N2:见图1 See Fig. 1 SP, PS: 见表1 See Table 1

Fig. 4 Leaf area changes of individual plant of Spartina alterniflora and Phragmites australis in mixed-culture

图5 混合种群中单株互花米草在不同密度芦苇影响下叶宽 (a) 与叶数 (b) 的变化图注同图4

Fig. 5 Changes of leaf width (a) and leaf number (b) of individual Spartina alterniflora under different density series of Phragmites australis Notes see Fig. 4

图6 混合种群中单株芦苇在不同密度互花米草影响下叶长 (a)、叶宽 (b) 与叶数 (c) 的变化图注同图4

Fig. 6 Changes of leaf length (a), leaf width (b) and leaf number (c) of individual Phragmites australis under different density series of Spartina alterniflora Notes see Fig. 4

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氮水平和竞争对互花米草与芦苇叶特征的影响

EFFECTS OF NITROGEN AVAILABILITY AND COMPETITION ON LEAF CHARACTERISTICS OF SPARTINA ALTERNIFLORA AND PHRAGMITES AUSTRALIS

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