氮磷供应量及比例对灰绿藜种子性状的影响

doi:10.17521/cjpe.2018.0085

摘要/Abstract

摘要：

全球氮沉降不仅改变土壤氮和磷的有效性, 同时也改变氮磷比例。氮磷供应量、比例及其交互作用可能会影响植物种子性状。该研究在内蒙古草原基于沙培盆栽实验种植灰绿藜(Chenopodium glaucum), 设置3个氮磷供应量水平和3个氮磷比例的正交实验来探究氮磷供应量、比例及其交互作用对灰绿藜种子性状的影响。结果发现氮磷供应量对种子氮浓度、磷浓度和萌发率影响的相对贡献(15%-24%)大于氮磷比例(3%-7%), 而种子大小只受氮磷比例的影响。同时氮磷供应量和比例之间的交互作用显著影响种子氮浓度和磷浓度。同等氮磷比例情况下, 低量养分供应提高种子氮浓度、磷浓度和萌发率。氮磷比例只有在养分匮乏的环境中才会对种子大小和萌发率产生显著影响。总之, 灰绿藜种子不同性状对氮或磷限制的敏感性不同, 同时种子性状也对养分限制表现出适应性和被动响应。

关键词: 种子性状, 氮供应水平, 磷供应水平, 氮磷比例, 沙培盆栽实验, 适应性响应

Abstract:

Aims Global nitrogen (N) deposition not only alters soil N and phosphorus (P) availability, but also changes their ratio. The levels and ratios of N and P supply and their interaction may simultaneously influence plant seed traits. However, so far there has been no experiments to distinguish these complex impacts on plant seed traits in the field.

Methods A pot experiment with a factorial design of three levels and ratios of N and P supply was conducted in the Nei Mongol grassland to explore the effects of levels and ratios of N and P supply and their interaction on seed traits of Chenopodium glaucum.

Important findings We found that the relative contribution (15%-24%) of N and P supply levels in affecting the N concentrations, P concentrations and germination rates of seeds was larger than that (3%-7%) of N:P supply ratios, whereas seed size was only significantly influenced by N:P. Simultaneously, seed N and P concentrations were impacted by the interaction of N and P supply levels and ratios. At the same N:P, decrease in nutrient supply levels increased seed N concentrations, P concentrations and germination rates. N:P supply ratios only had a significant effect on seed size and germination rates under low nutrient levels. Overall, these results indicate that different seed traits of C. glaucum show different sensitivities to N or P limitations, leading to adaptive and passive responses under different nutrient limitations. This study presents the the first field experiment to distinguish the effects of nutrient supply levels, ratios and their interactions on plant seed traits, which provides a new case study on the influences of global N deposition on future dynamics of plant population and community.

Key words: seed trait, nutrient supply level, phosphorus supply level, N:P, sand cultured pot experiment, adaptive response

田大栓. 氮磷供应量及比例对灰绿藜种子性状的影响. 植物生态学报, 2018, 42(9): 963-970. DOI: 10.17521/cjpe.2018.0085

TIAN Da-Shuan. Effects of the supply levels and ratios of nitrogen and phosphorus on seed traits of Chenopodium glaucum. Chinese Journal of Plant Ecology, 2018, 42(9): 963-970. DOI: 10.17521/cjpe.2018.0085

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2018.0085

https://www.plant-ecology.com/CN/Y2018/V42/I9/963

图/表 5

表1 所有处理的氮磷供应量

Table 1 Levels and ratios of nitrogen and phosphorus supply in each treatment

N:P	氮/磷供应量 N/P supply amount (mg·pot^-1)
N:P	低量 Low	中量 Middle	高量 High
5	8.80/1.76	26.55/5.28	79.15/15.83
15	15.15/1.01	45.75/3.05	137.11/9.14
45	26.55/0.59	79.15/1.76	237.62/5.28

表2 氮磷供应量、比例及其交互作用对种子大小、氮浓度、磷浓度、萌发率和萌发速度的影响

Table 2 Effects of nitrogen (N) and phosphorus (P) supply levels, ratios and their interactions on the size, N concentrations, P concentrations, germination rates and germination speed of seeds

种子性状 Seed trait	氮磷供应水平 N and P supply level	氮磷供应比例 N:P supply ratio	交互作用 Their interaction
自由度 d.f.	2	2	4
种子大小 Seed size	0.74%^ns	4.41%^*	5.38%^ns
氮浓度 N concentration	23.58%^***	2.90%^ns	13.77%^**
磷浓度 P concentration	23.21%^***	7.07%^**	17.45%^***
萌发率 Germination rate	15.13%^***	5.86%^**	2.79%^ns
萌发速度 Germination speed	0.20%^ns	2.00%^ns	4.07%^ns

图1 氮磷供应量或比例对灰绿藜种子大小的影响(平均值±标准误差, n = 20)。*表示相同氮磷供应水平下不同氮磷比例的效应是显著的。

Fig. 1 Effects of N and P supply levels or ratios on seed size of Chenopodium glaucum (mean ± SE, n = 20). * represents a significant impact of N:P at the same nutrient supply level.

图2 氮磷供应量或比例对灰绿藜种子氮浓度和磷浓度的影响(平均值±标准误差, n = 20)。*表示相同氮磷供应水平下不同氮磷比例的影响是显著的。不同字母表示相同氮磷供应比例情况下不同氮磷量的效应是显著的。

Fig. 2 Effects of nitrogen (N) and phosphorus (P) supply levels or ratios on seed N and P contents of Chenopodium glaucum (mean ± SE, n = 20). * represents a significant effect of N:P at the same nutrient supply level. Different letters indicate a significant effect of nutrient supply level at the same N:P.

图3 氮磷供应量或比例对灰绿藜种子萌发率和萌发速度的影响(平均值±标准误差, n = 20)。*表示相同氮磷供应水平下不同氮磷比例的影响是显著的。不同字母表示相同氮磷供应比例情况下不同氮磷供应量的效应是显著的。

Fig. 3 Effects of nitrogen (N) and phosphorus (P) supply levels or ratios on seed germination rate and speed of Chenopodium glaucum (mean ± SE, n = 20). * indicates a significant effect of N:P at the same nutrient supply level. Different letters indicate a significant effect of nutrient supply level at the same N:P.

参考文献 57

[1]	Aarssen LW, Burton SM ( 1990). Maternal effects at 4 levels in Senecio vulgaris(Asteraceae) grown on a soil nutrient gradient. American Journal of Botany, 77, 1231-1240. DOI URL
[2]	Bai Y, Wu J, Clark CM, Naeem S, Pan Q, Huang J, Zhang L, Han X ( 2010). Tradeoffs and thresholds in the effects of nitrogen addition on biodiversity and ecosystem functioning: Evidence from inner Mongolia Grasslands. Global Change Biology, 16, 358-372. DOI URL
[3]	Balestri E, Gobert S, Lepoint G, Lardicci C ( 2009). Seed nutrient content and nutritional status of Posidonia oceanica seedlings in the northwestern Mediterranean Sea. Marine Ecology Progress Series, 388, 99-109. DOI URL
[4]	Bobbink R, Hicks K, Galloway J, Spranger T, Alkemade R, Ashmore M, Bustamante M, Cinderby S, Davidson E, Dentener F, Emmett B ( 2010). Global assessment of nitrogen deposition effects on terrestrial plant diversity: A synthesis. Ecological Applications, 20, 30-59. DOI URL
[5]	Breen AN, Richards JH ( 2008). Irrigation and fertilization effects on seed number, size, germination and seedling growth: Implications for desert shrub establishment. Oecologia, 157, 13-19. DOI URL
[6]	Cornwell WK, Cornelissen JHC, Amatangelo K, Dorrepaal E, Eviner VT, Godoy O, Hobbie SE, Hoorens B, Kurokawa H, Pérez-Harguindeguy N, Quested HM ( 2008). Plant species traits are the predominant control on litter decomposition rates within biomes worldwide. Ecology Letters, 11, 1065-1071. DOI URL PMID
[7]	Defalco LA, Bryla DR, Smith-Longozo V, Nowak RS ( 2003). Are Mojave Desert annual species equal? Resource acquisition and allocation for the invasive grass Bromus madritensis subsp rubens (Poaceae) and two native species. American Journal of Botany, 90, 1045-1053.
[8]	Fortunel C, Violle C, Roumet C, Buatois B, Navas ML, Garnier E ( 2009). Allocation strategies and seed traits are hardly affected by nitrogen supply in 18 species differing in successional status. Perspectives in Plant Ecology Evolution and Systematics, 11, 267-283. DOI URL
[9]	Fujita Y, de Ruiter PC, Wassen MJ, Heil GW ( 2010). Time-dependent, species-specific effects of N:P stoichiometry on grassland plant growth. Plant and Soil, 334, 99-112. DOI URL
[10]	Galloway LF ( 2001). Parental environmental effects on life history in the herbaceous plant Campanula americana. Ecology, 82, 2781-2789.
[11]	Grant CA, Flaten DN, Tomasiewicz DJ, Sheppard SC ( 2001). The importance of early season phosphorus nutrition. Canadian Journal of Plant Science, 81, 211-224. DOI URL
[12]	Groom PK, Lamont BB ( 2010). Phosphorus accumulation in Proteaceae seeds: A synthesis. Plant and Soil, 334, 61-72. DOI URL
[13]	Gusewell S ( 2004). N:P ratios in terrestrial plants: Variation and functional significance. New Phytologist, 164, 243-266. DOI URL
[14]	Gusewell S ( 2005a ). High nitrogen: phosphorus ratios reduce nutrient retention and second-year growth of wetland sedges. New Phytologist, 166, 537-550. DOI URL
[15]	Gusewell S ( 2005b ). Responses of wetland graminoids to the relative supply of nitrogen and phosphorus. Plant Ecology, 176, 35-55. DOI URL
[16]	Gusewell S, Bollens U ( 2003). Composition of plant species mixtures grown at various N:P ratios and levels of nutrient supply. Basic and Applied Ecology, 4, 453-466. DOI URL
[17]	Hejcman M, Kristalova V, Cervena K, Hrdlickova J, Pavlu V ( 2012). Effect of nitrogen, phosphorus and potassium availability on mother plant size, seed production and germination ability of Rumex crispus. Weed Research, 52, 260-268. DOI URL
[18]	Horak MJ, Wax LM ( 1991). Germination and seedling development of bigroot morningglory ( Ipomoea pandurata). Weed Science, 39, 390-396. DOI URL
[19]	Howard TG, Goldberg DE ( 2001). Competitive response hierarchies for germination, growth, and survival and their influence on abundance. Ecology, 82, 979-990. DOI URL
[20]	Hrdlickova J, Hejcman M, Kristalova V, Pavlu V ( 2011). Production, size, and germination of broad-leaved dock seeds collected from mother plants grown under different nitrogen, phosphorus, and potassium supplies. Weed Biology and Management, 11, 190-201. DOI URL
[21]	Hu XY, Sun ZG, Zhang DY, Sun WG, Zhu H, Ren P ( 2017). Germination and seedling growth of different N-substrate seeds of Suaeda salsa subjected to salinity stress and nitrogen loading in the newly created marshes of the Yellow River estuary, China. Acta Ecologica Sinica, 37, 8499-8510.
	[ 胡星云, 孙志高, 张党玉, 孙文广, 祝贺, 任鹏 ( 2017). 黄河口不同氮基质碱蓬种子萌发及幼苗生长对盐分及氮输入的响应. 生态学报, 37, 8499-8510.]
[22]	Lavorel S, Garnier E ( 2002). Predicting changes in community composition and ecosystem functioning from plant traits: Revisiting the Holy Grail. Functional Ecology, 16, 545-556. DOI URL
[23]	Leishman MR ( 2001). Does the seed size/number trade-off model determine plant community structure? An assessment of the model mechanisms and their generality. Oikos, 93, 294-302. DOI URL
[24]	Li Y, Hou L, Song B, Yang L, Li L ( 2017). Effects of increased nitrogen and phosphorus deposition on offspring performance of two dominant species in a temperate steppe ecosystem. Scientific Reports, 7, 40951. DOI: 10.1038/?srep40951. DOI URL PMID
[25]	Li Y, Niu S, Yu G ( 2016). Aggravated phosphorus limitation on biomass production under increasing nitrogen loading: A meta-analysis. Global Change Biology, 22, 934-943. DOI URL PMID
[26]	Liu L, Greaver TL ( 2010). A global perspective on belowground carbon dynamics under nitrogen enrichment. Ecology Letters, 13, 819-828. DOI URL
[27]	Liu QY, Jiang M, Wang GD, Lu XG, Wang M, Lou YJ, Yuan YX ( 2013). Effect of exogenous phosphorus inputs on seed germination of soil seed bank in marshes in Xingkai lake. Wetland Science, 11, 41-47. DOI URL
	[ 刘庆艳, 姜明, 王国栋, 吕宪国, 王铭, 娄彦景, 袁宇翔 ( 2013). 外源磷输入对兴凯湖沼泽土壤种子库种子萌发的影响. 湿地科学, 11, 41-47.] DOI URL
[28]	Liu X, Zhang Y, Han W, Tang A, Shen J, Cui Z, Vitousek P, Erisman JW, Goulding K, Christie P, Fangmeier A ( 2013). Enhanced nitrogen deposition over China. Nature, 494, 459-462. DOI URL PMID
[29]	Liu Y, Gao P, Zhang L, Niu X, Wang B ( 2016). Spatial heterogeneity distribution of soil total nitrogen and total phosphorus in the Yaoxiang watershed in a hilly area of northern China based on geographic information system and geostatistics. Ecology and Evolution, 6, 6807-6816. DOI URL PMID
[30]	Liu ZM, Li RP, Li XH, Luo YM, Wang HM, Jiang DM, Nan YH ( 2004a ). A comparative study of seed weight of 69 plant species in Horqin Sandyland, China. Acta Phytoecologica Sinica, 28, 225-230. DOI URL
	[ 刘志民, 李荣平, 李雪华, 骆永明, 王红梅, 蒋德明, 南寅镐 ( 2004a ). 科尔沁沙地69种植物种子重量比较研究. 植物生态学报, 28, 225-230.] DOI URL
[31]	Liu ZM, Li XH, Li RP, Jiang DM, Cao CY, Chen XL ( 2004b ). A comparative of seed germination for 31 annual species of the Horqin steppe. Acta Ecologica Sinica, 24, 648-653. DOI URL
	[ 刘志民, 李雪华, 李荣平, 蒋德明, 曹成有, 常学礼 ( 2004b ). 科尔沁沙地31种1年生植物萌发特性比较研究. 生态学报, 24, 648-653.] DOI URL
[32]	Lu M, Yang Y, Luo Y, Fang C, Zhou X, Chen J, Yang X, Li B ( 2011). Responses of ecosystem nitrogen cycle to nitrogen addition: A meta-analysis. New Phytologist, 189, 1040-1050. DOI URL PMID
[33]	Luo X, Mazer SJ, Guo H, Zhang N, Weiner J, Hu S ( 2016). Nitrogen: phosphorous supply ratio and allometry in five alpine plant species. Ecology and Evolution, 6, 8881-8892. DOI URL PMID
[34]	Ma QF, Longnecker N, Atkins C ( 2002). Varying phosphorus supply and development, growth and seed yield in narrow-leafed lupin. Plant and Soil, 239, 79-85. DOI URL
[35]	Manning P, Houston K, Evans T ( 2009). Shifts in seed size across experimental nitrogen enrichment and plant density gradients. Basic and Applied Ecology, 10, 300-308. DOI URL
[36]	Milberg P, Lamont BB ( 1997). Seed/cotyledon size and nutrient content play a major role in early performance of species on nutrient-poor soils. New Phytologist, 137, 665-672. DOI URL
[37]	Moles AT, Westoby M ( 2004). Seedling survival and seed size: A synthesis of the literature. Journal of Ecology, 92, 372-383. DOI URL
[38]	Nadeem M, Mollier A, Morel C, Vives A, Prud’homme L, Pellerin S ( 2011). Relative contribution of seed phosphorus reserves and exogenous phosphorus uptake to maize ( Zea mays L.) nutrition during early growth stages. Plant and Soil, 346, 231-244. DOI URL
[39]	Naegle ER, Burton JW, Carter TE, Rufty TW ( 2005). Influence of seed nitrogen content on seedling growth and recovery from nitrogen stress. Plant and Soil, 271, 329-340. DOI URL
[40]	Navarro L, Guitian J ( 2003). Seed germination and seedling survival of two threatened endemic species of the northwest Iberian peninsula. Biological Conservation, 109, 313-320. DOI URL
[41]	Pe?uelas J, Poulter B, Sardans J, Ciais P, van der Velde M, Bopp L, Boucher O, Godderis Y, Hinsinger P, Llusia J, Nardin E ( 2013). Human-induced nitrogen-phosphorus imbalances alter natural and managed ecosystems across the globe. Nature Communications, 4, 2934. DOI: 10.1038/?ncomms3934. DOI URL PMID
[42]	Schmid B, Dolt C ( 1994). Effects of maternal and paternal environment and genotype on offspring phenotype in Solidago altissima L. Evolution, 48, 1525-1549. DOI URL PMID
[43]	Stevens CJ, Dise NB, Mountford JO, Gowing DJ ( 2004). Impact of nitrogen deposition on the species richness of grasslands. Science, 303, 1876-1879. DOI URL PMID
[44]	Sultan SE ( 1996). Phenotypic plasticity for offspring traits in Polygonum persicaria. Ecology, 77, 1791-1807.
[45]	Sun ZG, Song HL, Hu XY ( 2017). Response of germination and seedling growth of Suaeda salsa seeds from tidal marshes in the Yellow River estuary to N/P ratio of soil. Wetland Science, 15, 10-19.
	[ 孙志高, 宋红丽, 胡星云 ( 2017). 黄河口潮滩碱蓬种子萌发与幼苗生长对土壤中氮磷比的响应. 湿地科学, 15, 10-19.]
[46]	Tripathi RS, Khan ML ( 1990). Effects of seed weight and microsite characteristics on germination and seedling fitness in 2 species of Quercus in a subtropical wet hill forest. Oikos, 57, 289-296. DOI URL
[47]	Tungate KD, Burton MG, Susko DJ, Sermons SM, Rufty TW ( 2006). Altered weed reproduction and maternal effects under low-nitrogen fertility. Weed Science, 54, 847-853. DOI URL
[48]	Verdu M, Traveset A ( 2005). Early emergence enhances plant fitness: A phylogenetically controlled meta-analysis. Ecology, 86, 1385-1394. DOI URL
[49]	Vergeer P, Rengelink R, Ouborg NJ, Roelofs JGM ( 2003). Effects of population size and genetic variation on the response of Succisa pratensis to eutrophication and acidification. Journal of Ecology, 91, 600-609. DOI URL
[50]	Violle C, Castro H, Richarte J, Navas ML ( 2009). Intraspecific seed trait variations and competition: Passive or adaptive response? Functional Ecology, 23, 612-620. DOI URL
[51]	Violle C, Navas ML, Vile D, Kazakou E, Fortunel C, Hummel I, Garnier E ( 2007). Let the concept of trait be functional! Oikos, 116, 882-892. DOI URL
[52]	Walck JL, Hidayati SN, Dixon KW, Thompson K, Poschlod P ( 2011). Climate change and plant regeneration from seed. Global Change Biology, 17, 2145-2161. DOI URL
[53]	Walters MB, Reich PB ( 2000). Seed size, nitrogen supply, and growth rate affect tree seedling survival in deep shade. Ecology, 81, 1887-1901. DOI URL
[54]	Wulff RD, Causin HF, Benitez O, Bacalini PA ( 1999). Intraspecific variability and maternal effects in the response to nutrient addition in Chenopodium album. Canadian Journal of Botany, 77, 1150-1158. DOI URL
[55]	Zhang H, Chang R, Guo X, Liang X, Wang R, Liu J ( 2017). Shifts in growth and competitive dominance of the invasive plant Alternanthera philoxeroides under different nitrogen and phosphorus supply. Environmental and Experimental Botany, 135, 118-125. DOI URL
[56]	Zhang M, Nyborg M, Mcgill WB ( 1990). Phosphorus concentration in barley ( Hordeum vulgare L.) seed: Influence on seedling growth and dry-matter production. Plant and Soil, 122, 79-83. DOI URL
[57]	Zhu YG, Smith SE ( 2001). Seed phosphorus (P) content affects growth, and P uptake of wheat plants and their association with arbuscular mycorrhizal (AM) fungi. Plant and Soil, 231, 105-112. DOI URL