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

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

Abstract

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

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[J]. Chin J Plant Ecol, 2008, 32(2): 392-401.

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URL: https://www.plant-ecology.com/EN/10.3773/j.issn.1005-264x.2008.02.017

https://www.plant-ecology.com/EN/Y2008/V32/I2/392

Figures/Tables 9

References 40

[1]	An SQ, Gu BH, Zhou CF, Wang ZS, Deng ZF, Zhi YB, Li HL, Chen L, Yu DH, Liu YH (2007). Spartina invasion in China: implications for invasive species management and future research. Weed Research, 47, 183-191.
[2]	Bastlová D, Cízková H, Bastl M, Květ J (2004). Growth of Lythrum salicaria and Phragmites australis plants originating from a wide geographical area: response to nutrient and water supply. Global Ecology and Biogeography, 13, 259-271.
[3]	Callaway JC, Josselyn MN (1992). The introduction and spread of smooth cordgrass (Spartina alterniflora) in South San Francisco Bay. Estuaries, 15, 218-226.
[4]	Chambers RM, Mozdzer TJ, Ambrose JC (1998). Effects of salinity and sulfide on the distribution of Phragmites australis and Spartina alterniflora in a tidal saltmarsh. Aquatic Botany, 62, 161-169.
[5]	Chen HL, Li B, Fang CM, Chen JK, Wu JH (2007). Exotic plant influences soil nematode communities through litter input. Soil Biology & Biochemistry, 39, 1782-1793.
[6]	Chen ZY (陈中义), Gao H (高慧), Wu H (吴涵), Li B (李博) (2005). Effects of simulated canopy shade on seed germination and seedlings growth of Spartina alterniflora and Scirpus mariqueter. Hubei Agricultural Sciences (湖北农业科学), (2), 82-84. (in Chinese with English abstract)
[7]	Chen YN (陈一宁), Gao S (高抒), Jia JJ (贾建军), Wang AJ (王爱军) (2005). Tidal flat ecological changes by transplanting Spartina anglica and Spartina alterniflora, northern Jiangsu coast. Oceanologia et Limnologia Sinica (海洋与湖泊), 36, 394-403. (in Chinese with English abstract)
[8]	Cox EF (1972). A photoelectric digital scanner for measuring leaf area. New Phytologist, 71, 819-823.
[9]	Dai T, Wiegert RG (1997). A field study of photosynthetic capacity and its response to nitrogen fertilization in Spartina alterniflora. Estuarine, Coastal and Shelf Science, 45, 273-283.
[10]	Deng ZF (邓自发), An SQ (安树青), Zhi YB (智颖飙), Zhou CF (周长芳), Chen L (陈琳), Zhao CJ (赵聪蛟), Fang SB (方淑波), Li HL (李红丽) (2006). Preliminary studies on invasive model and outbreak mechanism of exotic species, Spartina alterniflora Loisel. Acta Ecologica Sinica (生态学报), 26, 2678-2686. (in Chinese with English abstract)
[11]	Duan XN (段晓男), Wang XK (王效科), Ouyang ZY (欧阳志云), Miao H (苗鸿), Guo R (郭然) (2004). The biomass of Phragmites australis and its influencing factors in Wuliangsuhai. Acta Phytoecologica Sinica (植物生态学报), 28, 246-251. (in Chinese with English abstract)
[12]	Emery NC, Ewanchuk PJ, Bertness MD (2001). Competition and salt-marsh plant zonation: stress tolerators may be dominant competitors. Ecology, 82, 2471-2485.
[13]	Esselink P, Zijlstra W, Dijkema KS, van Diggenlen R (2000). The effects of decreased management on plant-species distribution patterns in a salt marsh nature reserve in the Wadden Sea. Biological Conservation, 93, 61-76.
[14]	Folgi S, Marchesini R, Gerdol R (2002). Reed (Phragmites australis) decline in a brackish wetland in Italy. Marine Environment Research, 53, 465-479.
[15]	He WS, Feagin R, Lu JJ, Liu WL, Yan Q, Xie ZF (2007). Impacts of introduced Spartina alterniflora along an elevation gradient at the Jiuduansha Shoals in the Yangtze Estuary, suburban Shanghai, China. Ecological Engineering, 29, 245-248.
[16]	Knight DH (1973). Leaf area dynamics of a shortgrass Prairie in Colorado. Ecology, 54, 891-896.
[17]	Levine JM, Brewer JS, Bertness MD (1998). Nutrients, competition and plant zonation in a New England salt marsh. Journal of Ecology, 86, 285-292.
[18]	Li BG (李宝光), Tao XH (陶秀花), Ni GP (倪国平), Huang DH (黄冬华), Song XM (宋小民) (2006). Study on determining plant leaf area by scanning image pixels method. Acta Agriculturae Jiangxi (江西农业学报), 18(3), 78-81. (in Chinese with English abstract)
[19]	Liang X (梁霞), Zhang LQ (张利权), Zhao GQ (赵广琦) (2006). A comparison of photosynthetic characteristics between Spartina alterniflora and Phragmites australis under different CO₂ concentrations. Acta Ecologica Sinica (生态学报), 26, 842-848. (in Chinese with English abstract)
[20]	Liu JW (刘金文), Sha W (沙伟) (2004). Origin, expansion and decline of Phragmites australis. Guizhou Science (贵州科学), 22(2), 65-68, 82. (in Chinese with English abstract)
[21]	Marks M, Lapin B, Randall J (1994). Phragmites australis (P. communis): threats, management, and monitoring. Natural Areas Journal, 14, 285-294.
[22]	Marshll JD, Waring RH (1986). Comparison of methods of estimating leaf-area index in old-growth Douglas-fir. Ecology, 67, 975-979.
[23]	Minchinton TE (2002). Disturbance by wrack facilitates spread of Phragmites australis in a coastal marsh. Journal of Experimental Marine Biology and Ecology, 281, 89-107.
[24]	Ostendorp W (1989). 'Die-back' of reeds in Europe—a critical review of literature. Aquatic Botany, 35, 5-26.
[25]	Perry LG, Galatowitsch SM (2006). Light competition for invasive species control: a model of cover crop—weed competition and implications for Phalaris arundinacea control in sedge meadow wetlands. Euphytica, 148, 121-134.
[26]	Pezeshki SR, DeLaune RD, Lindau CW (1988). Interaction among sediment anaerobiosis, nitrogen uptake and photosynthesis of Spartina alterniflora. Physiologia Plantarum, 74, 561-565.
[27]	Philipp KR, Field RT (2005). Phragmites australis expansion in Delaware Bay salt marshes. Ecological Engineering, 25, 275-291.
[28]	Poulin B, Lefebvre G, Mauchamp A (2002). Habitat requirements of passerines and reedbed management in Southern France. Biological Conservation, 107, 315-325.
[29]	Qin WH (秦卫华), Wang Z (王智), Jiang MK (蒋明康) (2004). Invasion of Spartina alterniflora Loisel to two wetland nature reserve in Changjiang Estuary. Weed Science (杂草科学), (4), 15-16. (in Chinese)
[30]	Rice D, Rooth J, Stevenson JC (2000). Colonization and expansion of Phragmites australis in upper Chesapeake Bay tidal marshes. Wetlands, 20, 280-299.
[31]	Rickey MA, Anderson RC (2004). Effects of nitrogen addition on the invasive grass Phragmites australis and a native competitor Spartina pectinata. Journal of Applied Ecology, 41, 888-896.
[32]	Seliskar DM, Gallagher JL, Burdick DM, Mutz LA (2002). The regulation of ecosystem functions by ecotypic variation in the dominant plant: a Spartina alterniflora salt-marsh case study. Journal of Ecology, 90, 1-11.
[33]	Tang CJ (唐承佳), Lu JJ (陆健健) (2003). Studies on plant community on the Jiuduansha Shoal at the Yangtze Estuary. Acta Ecologica Sinica (生态学报), 23, 399-403. (in Chinese with English abstract)
[34]	Vasquez EA, Glenn EP, Guntenspergen GR, Brown JJ, Nelson SG (2006). Salt tolerance and osmotic adjustment of Spartina alterniflora (Poaceae) and the invasive M haplotype of Phragmites australis (Poaceae) along a salinity gradient. American Journal of Botany, 93, 1784-1790.
[35]	Wang Q (王卿), An SQ (安树青), Ma ZJ (马志军), Zhao B (赵斌), Chen JK (陈家宽), Li B (李博) (2006). Invasive Spartina alterniflora: biology, ecology and management. Acta Phytotaxonomica Sinica (植物分类学报), 44, 559-588. (in Chinese with English abstract)
[36]	Wang Q, Wang CH, Zhao B, Ma ZJ, Luo YQ, Chen JK, Li B (2006). Effects of growing conditions on the growth of and interactions between salt marsh plants: implications for invasibility of habitats. Biological Invasion, 8, 1547-1560.
[37]	Weis JS, Weis P (2003). Is the invasion of the common reed, Phragmites australis, into tidal marshes of the eastern US an ecological disaster? Marine Pollution Bulletin, 46, 816-820.
[38]	Xiao Q (肖强), Ye WJ (叶文景), Zhu Z (朱珠), Chen Y (陈瑶), Zheng HL (郑海雷) (2005). A simple non-destructive method to measure leaf area using digital camera and Photoshop software. Chinese Journal of Ecology (生态学杂志), 24, 711-714. (in Chinese with English abstract)
[39]	Yuan ZM (袁哲明), Fu LC (傅凌才) (1997). Scanner measurement of insect-damaged leaf area. Journal of Hunan Agricultural University (湖南农业大学学报), 23, 472-473. (in Chinese with English abstract)
[40]	Zhao GQ (赵广琦), Zhang LQ (张利权), Liang X (梁霞) (2005). A comparison of photosynthetic characteristics between an invasive plant Spartina alterniflora and an indigenous plant Phragmites australis. Acta Ecologica Sinica (生态学报), 25, 1604-1611. (in Chinese with English abstract)

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

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

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^*

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^*

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^*