模拟氮沉降和干旱对准噶尔盆地两种一年生荒漠植物生长和光合生理的影响

doi:10.3773/j.issn.1005-264x.2010.12.005

摘要/Abstract

摘要：

氮素和水分是荒漠生态系统的两个主要限制因子, 研究两者对荒漠植物的效应有助于深入了解荒漠生态系统对全球变化的响应。该文选择准噶尔盆地荒漠地区两种常见的一年生植物涩荠(Malcolmia africana)和钩刺雾冰藜(Bassia hyssopifolia), 设置0、0.18和0.72 g N·m ^-2·week ^-13个施氮浓度和湿润与干旱两个土壤水分处理, 研究模拟氮沉降增加和干旱对其生长和光合生理的影响。结果表明: (1)两种植物的根长、根重、叶片数、叶面积、总生物量和冠根比均随着施氮浓度的增加而增加, 干旱能够抑制氮对植物生长的促进作用, 但是, 氮的增加同时也能部分缓解干旱对植物生长的影响。与钩刺雾冰藜相比, 涩荠的根长、生物量和冠根比更易受氮增加和干旱的影响。(2)两种植物的最大净光合速率、叶绿素含量、可溶性蛋白含量随着氮浓度增加而增加, 但涩荠和钩刺雾冰藜对氮增加和干旱的生理响应也有所不同, 涩荠的响应更加敏感。两种植物对氮沉降和干旱胁迫响应的差异可能是其生活型等生物学特性差异所引起。通过对两种一年生植物的生长和光合生理分析表明, 在古尔班通古特沙漠, 春季丰富的降水和氮素增加将有利于涩荠和钩刺雾冰藜的生长和生产力的增加, 相对地下生长, 地上部分增加更显著。当干旱季节来临时, 氮的增加又能够在一定程度上降低干旱对这两种植物的负效应, 说明其对干旱具有一定的生态补偿作用。

关键词: 一年生荒荒漠植物, 生物量, 干旱胁迫, 氮沉降, 可溶性蛋白

Abstract:

Aims The two primary limiting factors for biological activities in desert ecosystems are nitrogen and water. Our study, which examined their combined effects, can provide insight into the responses of arid ecosystems to global climate change. We selected two typical annual desert plants, Malcolmia africana and Bassia hyssopifolia to determine the combined effects of nitrogen deposition and drought stress on their growth and photosynthetic physiological responses.
Methods Three levels of N addition (0, 0.18 and 0.72 g N·m ^-2·week ^-1) and two soil watering regimes (60%-70% and 30%-40% of field capacity) were randomly provided in order to simulate nitrogen deposition and drought stress. Changes in plant growth and photosynthetic physiological traits were measured shortly before flowering.
Important findings N supply and drought stress significantly affected growth of both species. With the enhancement of N supply, we found an increase in growth parameters (including root length, root weight, leaf number, leaf area, total biomass and shoot/root (S/R)). At the same N level, increased drought stress could counteract the positive effects of N supply on plant growth. Increased N, however, could also alleviate the negative effects caused by drought stress. With the increasing N addition, we also observed increase of physiological indices (net photosynthetic rate, content of chlorophyll and soluble protein). Malcolmia africana was more sensitive to N supply and drought stress than B. hyssopifolia. The different responses of the two species may due to their different biological characteristics, such as life form. The results indicated that N and water pulses in spring in this desert would be beneficial to the growth and productivity of M. africana and B. hyssopifolia, especially for aboveground parts. Moreover, N deposition could partially alleviate the negative effects caused by drought stress during the severe dry season.

Key words: annual desert plant, biomass, drought stress, N deposition, soluble protein

周晓兵, 张元明, 王莎莎, 张丙昌. 模拟氮沉降和干旱对准噶尔盆地两种一年生荒漠植物生长和光合生理的影响. 植物生态学报, 2010, 34(12): 1394-1403. DOI: 10.3773/j.issn.1005-264x.2010.12.005

ZHOU Xiao-Bing, ZHANG Yuan-Ming, WANG Sha-Sha, ZHANG Bing-Chang. Combined effects of simulated nitrogen deposition and drought stress on growth and photosynthetic physiological responses of two annual desert plants in Junggar Basin, China. Chinese Journal of Plant Ecology, 2010, 34(12): 1394-1403. DOI: 10.3773/j.issn.1005-264x.2010.12.005

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链接本文: https://www.plant-ecology.com/CN/10.3773/j.issn.1005-264x.2010.12.005

https://www.plant-ecology.com/CN/Y2010/V34/I12/1394

图/表 6

表1 土壤基质的物理和化学特征(平均值±标准偏差)

Table 1 Physical and chemical characteristics of substrate soil (mean ± SD)

pH	体积含水量 Volumetric water content (%)	有机碳 Organic C (g·kg^-1)	总氮 Total N (g·kg^-1)	总磷 Total P (g·kg^-1)	总钾 Total K (g·kg^-1)	有效氮 Available N (mg·kg^-1)	有效磷 Available P (mg·kg^-1)	有效钾 Available K (mg·kg^-1)
8.36 ± 0.16	7.74 ± 2.18	1.69 ± 0.55	0.19 ± 0.09	0.40 ± 0.04	10.94 ± 0.78	36.31 ± 9.44	6.31 ± 1.53	167.75 ± 20.72

表2 氮和水分处理对植物生长影响的双因素方差分析

Table 2 Two-way ANOVA analysis of N and water treatment effects on plant growth

物种 Species	处理 Treatment	根长 Root length	根重 Root weight	叶片数 Leaf number	叶面积 Leaf area	生物量 Biomass
涩荠 Malcolmia africana	水分 water	10.56^**	485.71^**	22.26^**	58.32^**	2 158.61^**
	氮 N	4.43^*	62.19^**	7.43^**	68.45^**	804.31^**
	水分×氮 Water × N	0.77	11.77^**	5.29^**	21.31^**	235.69^**
钩刺雾冰藜 Bassia hyssopifolia	水分 water	3.11	272.14^**	19.19^**	49.64^**	7 319.02^**
	氮 N	2.25	544.34^**	54.98^**	82.15^**	2 4217.89^**
	水分×氮 Water × N	0.08	21.99^**	31.87^**	23.42^**	4 463.67^**

图1 不同水分条件下涩荠和钩刺雾冰藜的根长、根重、叶片数(平均值±标准偏差, n = 5)。不同的字母代表处理间的差异显著(p < 0.05)。

Fig. 1 The root length, root weight, leaf number of Malcolmia africana and Bassia hyssopifolia under different water regimes (mean ± SD, n = 5). DS, drought stress; WW, well watered. Different letters indicate significance among treatments (p < 0.05).

图2 不同水分条件下涩荠和钩刺雾冰藜的叶面积、生物量、冠根比(平均值±标准偏差, n = 5)。图注同图1。

Fig. 2 The leaf area, biomass, shoot/root (S/R) of Malcolmia africana and Bassia hyssopifolia under different water regimes (mean ± SD, n = 5). Notes see Fig. 1.

表3 氮和水分处理对植物生理影响的双因素方差分析

Table 3 Two-way ANOVA analysis for the effects of N and water treatment on physiological traits

物种 Species	处理 Treatment	最大净光合速率 Max net photosynthetic rate	叶绿素 Chlorophyll	可溶性蛋白 Soluble protein
涩荠 Malcolmia africana	水分 water	148.18^**	17.76^**	6.85^*
	氮 N	69.02^**	59.05^**	51.36^*
	水分×氮 Water × N	1.33	15.24^**	0.13
钩刺雾冰藜 Bassia hyssopifolia	水分 water	1.97^**	2.54	84.34^**
	氮 N	23.38	201.16^**	60.16^**
	水分×氮 Water × N	2.35	4.92^**	13.67^**

图3 不同处理下涩荠和钩刺雾冰藜的最大净光合速率、叶绿素、可溶性蛋白含量(平均值±标准偏差, n = 3)。不同字母表示涩荠或钩刺雾冰藜之间差异显著(p < 0.05)。

Fig. 3 The maximum net photosynthetic rate (Pmax), chlorophyll (Chl), soluble protein (SP) of Malcolmia africana and Bassia hyssopifolia under different treatments (mean ± SD, n = 3). Different letters indicate significance of M. africana or B. hyssopifolia under different treatments (p < 0.05). CW, control + well watered; MW, medium N + well watered; HW, high N + well watered; CD, control + drought stress; MD, medium N + drought stress; HD, high N + drought stress.

参考文献 45

[1]	Anderson GQA, Andrews M, Percival SM, Kirby JS (1997). Nitrogen nutrition of brant (Branta bernicla L.) grazing on saltmarsh and pasture species. Proceedings of the XVIII International Grasslands Congress. Winnipeg and Saskatoon, Canada. 263-264.
[2]	Andrews M (1993). Nitrogen effects on the partitioning of dry matter between shoot and root of higher plants. Current Topics in Plant Physiology, 1, 119-126.
[3]	Andrews M, Sprent JI, Raven JA, Eady PE (1999). Relationships between shoot to root ratio, growth and leaf soluble protein concentration of Pisum sativum, Phaseolus vulgaris and Triticum aestivum under different nutrient deficiencies. Plant, Cell & Environment, 22, 949-958.
[4]	Baez S, Fargione J, Moore DI, Collins SL, Gosz JR (2007). Atmospheric nitrogen deposition in the northern Chihuahuan desert: temporal trends and potential consequences. Journal of Arid Environments, 68, 640-651.
[5]	Barathi P, Sundar D, Reddy AR (2001). Changes in mulberry leaf metabolism in response to water stress. Biologia Plantarum, 44, 83-87.
[6]	Barker DH, Vanier C, Naumburg E, Charlet TN, Nielsen KM, Newingham BA, Smith SD (2006). Enhanced monsoon precipitation and nitrogen deposition affect leaf traits and photosynthesis differently in spring and summer in the desert shrub Larrea tridentata. New Phytologist, 169, 799-808.
[7]	Bradford MM (1976). A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Analytical Biochemistry, 72, 248-254. DOI URL PMID
[8]	Brooks ML (2003). Effects of increased soil nitrogen on the dominance of alien annual plants in the Mojave Desert. Journal of Applied Ecology, 40, 344-353. DOI URL
[9]	Chen SP, Bai YF, Zhang LX, Han XG (2005). Comparing physiological responses of two dominant grass species to nitrogen addition in Xilin River Basin of China. Environmental and Experimental Botany, 53, 65-75. DOI URL
[10]	Compton JE, Watrud LS, Porteous LA, DeGrood S (2004). Response of soil microbial biomass and community composition to chronic nitrogen additions at Harvard forest. Forest Ecology and Management, 196, 143-158.
[11]	DaMatta FM, Loos RA, Silva EA, Loureiro ME, Ducatti C (2002). Effects of soil water deficit and nitrogen nutrition on water relations and photosynthesis of pot-grown Coffea canephora Pierre. Trees-Structure and Function, 16, 555-558.
[12]	Egli P, Schmid B (1999). Relationships between leaf nitrogen and limitations of photosynthesis in canopies of Solidago altissima. Acta Oecologica-International Journal of Ecology, 20, 559-570.
[13]	Evans J (1989). Partitioning of nitrogen between and within leaves grown under different irradiances. Australian Journal of Plant Physiology, 16, 533-548.
[14]	Fan HB (樊后保), Liu WF (刘文飞), Li YY (李燕燕), Liao YC (廖迎春), Yuan YH (袁颖红), Xu L (徐雷) (2007). Tree growth and soil nutrients in response to nitrogen deposition in a subtropical Chinese fir plantation. Acta Ecologica Sinica (生态学报), 27, 4630-4642. (in Chinese with English abstract)
[15]	Fenn ME, Haeuber R, Tonnesen GS, Baron JS, Grossman- Clarke S, Hope D, Jaffe DA, Copeland S, Geiser L, Rueth HM, Sickman JO (2003). Nitrogen emissions, deposition, and monitoring in the western United States. BioScience, 53, 391-403.
[16]	Garg BK, Kathju S, Burman U (2001). Influence of water stress on water relations, photosynthetic parameters and nitrogen metabolism of moth bean genotypes. Biologia Plantarum, 44, 289-292.
[17]	Hooper DU, Johnson L (1999). Nitrogen limitation in dryland ecosystems: responses to geographical and temporal variation in precipitation. Biogeochemistry, 46, 247-293.
[18]	Hyvonen R, Agren GI, Linder S, Persson T, Cotrufo MF, Ekblad A, Freeman M, Grelle A, Janssens IA, Jarvis PG, Kellomaki S, Lindroth A, Loustau D, Lundmark T, Norby RJ, Oren R, Pilegaard K, Ryan MG, Sigurdsson BD, Stromgren M, van Oijen M, Wallin G (2007). The likely impact of elevated CO₂, nitrogen deposition, increased temperature and management on carbon sequestration in temperate and boreal forest ecosystems: a literature review. New Phytologist, 173, 463-480.
[19]	Lajtha K, Schlesinger WH (1986). Plant response to variations in nitrogen availability in a desert shrubland community. Biogeochemistry, 2, 29-37.
[20]	Li DJ (李德军), Mo JM (莫江明), Fang YT (方运霆), Cai XA (蔡锡安), Xue JH (薛璟花), Xu GL (徐国良) (2004). Effects of simulated nitrogen deposition on growth and photosynthesis of Schima superba, Castanopsis chinensis and Cryptocarya concinna seedlings. Acta Ecologica Sinica (生态学报), 24, 876-882. (in Chinese with English abstract)
[21]	Li DJ (李德军), Mo JM (莫江明), Fang YT (方运霆), Li ZA (李志安) (2005). Effects of simulated nitrogen deposition on biomass production and allocation of Schima superba and Cryptocarya concinna seedlings in subtropical China. Acta Phytoecologica Sinica (植物生态学报), 29, 543-549. (in Chinese with English abstract)
[22]	Lichtenthaler HK, Wellburn AR (1983). Determinations of total carotenoids and chlorophylls a and b of leaf extracts in different solvents. Biochemical Society Transactions, 11, 591-592. DOI URL
[23]	Lu XK (鲁显楷), Mo JM (莫江明), Peng SL (彭少麟), Fang YT (方运霆) (2006). Effects of simulated N deposition on free amino acids and soluble protein of three dominant understory species in a monsoon evergreen broad-leaved forest of subtropical China. Acta Ecologica Sinica (生态学报), 26, 743-753. (in Chinese with English abstract)
[24]	Lü CQ (吕超群), Tian HQ (田汉勤), Huang Y (黄耀) (2007). Ecological effects of nitrogen deposition in terrestrial ecosystems. Journal of Plant Ecology (Chinese Version) (植物生态学报), 31, 205-218. (in Chinese with English abstract)
[25]	Ma X (马旭), Tian CY (田长彦), Feng G (冯固), Xiang XS (向祥盛) (2006). Spatio-temporal change of the application of chemical fertilizers in Xinjiang. Arid Land Geography (干旱区地理), 29, 439-444. (in Chinese with English abstract)
[26]	Mo JM, Zhang W, Zhu WX, Fang YT, Li DJ, Zhao P (2007). Response of soil respiration to simulated N deposition in a disturbed and a rehabilitated tropical forest in southern China. Plant and Soil, 296, 125-135. DOI URL
[27]	Nakaji T, Fukami M, Dokiya Y, Izuta T (2001). Effects of high nitrogen load on growth, photosynthesis and nutrient status of Cryptomeria japonica and Pinus densiflora seedlings. Trees-Structure and Function, 15, 453-461. DOI URL
[28]	Patterson TB, Guy RD, Dang QL (1997). Whole-plant nitrogen- and water-relations traits, and their associated trade-offs, in adjacent muskeg and upland boreal spruce species. Oecologia, 110, 160-168. DOI URL PMID
[29]	Phoenix GK, Hicks WK, Cinderby S, Kuylenstierna JCI, Stock WD, Dentener FJ, Giller KE, Austin AT, Lefroy RDB, Gimeno BS, Ashmore MR, Ineson P (2006). Atmospheric nitrogen deposition in world biodiversity hotspots: the need for a greater global perspective in assessing N deposition impacts. Global Change Biology, 12, 470-476. DOI URL
[30]	Poorter H, Nagel O (1999). The role of biomass allocation in the growth response of plants to different levels of light, CO₂, nutrients and water: a quantitative review. International Conference on Assimilate Transport and Partitioning (ICATP 99), Newcastle, Australia. 595-607.
[31]	Pregitzer KS, Burton AJ, Zak DR, Talhelm AF (2008). Simulated chronic nitrogen deposition increases carbon storage in Northern Temperate Forests. Global Change Biology, 14, 142-153. DOI URL
[32]	Schwinning S, Starr BI, Wojcik NJ, Miller ME, Ehleringer JE, Sanford RL (2005). Effects of nitrogen deposition on an arid grassland in the Colorado Plateau cold desert. Rangeland Ecology & Management, 58, 565-574.
[33]	Shangguan ZP, Shao MA, Dyckmans J (2000). Nitrogen nutrition and water stress effects on leaf photosynthetic gas exchange and water use efficiency in winter wheat. Environmental and Experimental Botany, 44, 141-149. DOI URL PMID
[34]	Throop HL (2005). Nitrogen deposition and herbivory affect biomass production and allocation in an annual plant. Oikos, 111, 91-100. DOI URL
[35]	Wan HW (万宏伟), Yang Y (杨阳), Bai SQ (白世勤), Xu YH (徐云虎), Bai YF (白永飞) (2008). Variations in leaf functional traits of six species along a nitrogen addition gradient in Leymus chinensis steppe in Inner Mongolia. Journal of Plant Ecology (Chinese Version) (植物生态学报), 32, 611-621. (in Chinese with English abstract)
[36]	Warren CR, Livingston NJ, Turpin DH (2004). Photosynthetic responses and N allocation in Douglas-fir needles following a brief pulse of nutrients. Tree Physiology, 24, 601-608. DOI URL PMID
[37]	Wu C (吴楚), Wang ZQ (王政权), Fan ZQ (范志强), Sun HL (孙海龙) (2003). Effects of different concentrations and form ratios of nitrogen on chlorophyll biosynthesis, photosynthesis, and biomass partitioning in Fraxinus mandshrica seedlings. Acta Phytoecologica Sinica (植物生态学报), 27, 771-779. (in Chinese with English abstract)
[38]	Wu FZ, Bao WK, Li FL, Wu N (2008). Effects of drought stress and N supply on the growth, biomass partitioning and water-use efficiency of Sophora davidii seedlings. Environmental and Experimental Botany, 63, 248-255. DOI URL
[39]	Wu FZ (吴福忠), Bao WK (包维楷), Wu N (吴宁) (2008). Growth, accumulation and partitioning of biomass, C, N of Sophora davidii seedlings in response to N supply in dry valley of upper Minjiang River. Acta Ecologica Sinica (生态学报), 28, 3817-3824. (in Chinese with English abstract)
[40]	Xu ZZ, Zhou GS, Wang YH (2007). Combined effects of elevated CO₂ and soil drought on carbon and nitrogen allocation of the desert shrub Caragana intermedia. Plant and Soil, 301, 87-97. DOI URL
[41]	Yuan XM (袁新民), Wang ZQ (王周琼) (1997). Studies on the NO₃^--N in the environment and soil. Arid Zone Research (干旱区研究), 14, 52-55. (in Chinese with English abstract)
[42]	Zavaleta ES, Shaw MR, Chiariello NR, Mooney HA, Field CB (2003). Additive effects of simulated climate changes, elevated CO₂, and nitrogen deposition on grassland diversity. Proceedings of the National Academy of Sciences of the United States of America, 100, 7650-7654. DOI URL PMID
[43]	Zhang LY (张立运), Chen CD (陈昌笃) (2002). On the general characteristics of plant diversity of Gurbantunggut sandy desert. Acta Ecologica Sinica (生态学报), 22, 1923-1932. (in Chinese with English abstract)
[44]	Zhang Y, Zheng LX, Liu XJ, Jickells T, Cape JN, Goulding K, Fangmeier A, Zhang FS (2008). Evidence for organic N deposition and its anthropogenic sources in China. Atmospheric Environment, 42, 1035-1041. DOI URL
[45]	Zhao DL, Reddy KR, Kakani VG, Reddy VR (2005). Nitrogen deficiency effects on plant growth, leaf photosynthesis, and hyperspectral reflectance properties of Sorghum. European Journal of Agronomy, 22, 391-403. DOI URL