Effects of the supply levels and ratios of nitrogen and phosphorus on seed traits of Chenopodium glaucum

TIAN Da-Shuan1

(1.Key Laboratory of Ecosystem Network Observation and Modeling, Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of Sciences, Beijing, China 100101)

【Abstract】Aims Global nitrogen (N) deposition not only alters the 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 experiment 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 contributions (15%–24%) of N and P supply levels in affecting the N concentrations, P concentrations and germination rates of seeds were larger than that (3%–7%) of N:P supply ratio, whereas seed size was only significantly influenced by N:P. Simultaneously, the seed N and P concentrations were impacted by the interaction of N and P supply levels and ratio. At the same N:P, the decrease in nutrient supply levels increased the seed N concentration, P concentration and germination rate. The N:P supply ratio only had a significant effect on the seed size and germination rate 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 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.

【Keywords】 seed trait; nitrogen supply level; phosphorus supply level; N:P; sand cultured pot experiment; adaptive response;


【Funds】 National Natural Science Foundation of China (31600356) National Basic Research Program of China (2017YFA0604801)

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    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.

    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.

    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.

    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.

    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.

    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.

    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.

    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.

    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.

    Galloway LF (2001). Parental environmental effects on life history in the herbaceous plant Campanula americana. Ecology, 82, 2781–2789.

    Grant CA, Flaten DN, Tomasiewicz DJ, Sheppard SC (2001). The importance of early season phosphorus nutrition. Canadian Journal of Plant Science, 81, 211–224.

    Groom PK, Lamont BB (2010). Phosphorus accumulation in Proteaceae seeds: A synthesis. Plant and Soil, 334, 61–72.

    Gusewell S (2004). N:P ratios in terrestrial plants: Variation and functional significance. New Phytologist, 164, 243–266.

    Gusewell S (2005a). High nitrogen:phosphorus ratios reduce nutrient retention and second-year growth of wetland sedges. New Phytologist, 166, 537–550.

    Gusewell S (2005b). Responses of wetland graminoids to the relative supply of nitrogen and phosphorus. Plant Ecology, 176, 35–55.

    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.

    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.

    Horak MJ, Wax LM (1991). Germination and seedling development of bigroot morningglory (Ipomoea pandurata). Weed Science, 39, 390–396.

    Howard TG, Goldberg DE (2001). Competitive response hierarchies for germination, growth, and survival and their influence on abundance. Ecology, 82, 979–990.

    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.

    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 (in Chinese).

    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.

    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.

    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.

    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.

    Liu L, Greaver TL (2010). A global perspective on belowground carbon dynamics under nitrogen enrichment. Ecology Letters, 13, 819–828.

    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.

    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.

    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.

    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.

    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.

    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.

    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.

    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.

    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.

    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.

    Moles AT, Westoby M (2004). Seedling survival and seed size: A synthesis of the literature. Journal of Ecology, 92, 372–383.

    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.

    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.

    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.

    Peñuelas 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.

    Schmid B, Dolt C (1994). Effects of maternal and paternal environment and genotype on offspring phenotype in Solidago altissima L. Evolution, 48, 1525–1549.

    Stevens CJ, Dise NB, Mountford JO, Gowing DJ (2004). Impact of nitrogen deposition on the species richness of grasslands. Science, 303, 1876–1879.

    Sultan SE (1996). Phenotypic plasticity for offspring traits in Polygonum persicaria. Ecology, 77, 1791–1807.

    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.

    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.

    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.

    Verdu M, Traveset A (2005). Early emergence enhances plant fitness: A phylogenetically controlled meta-analysis. Ecology, 86, 1385–1394.

    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.

    Violle C, Castro H, Richarte J, Navas ML (2009). Intraspecific seed trait variations and competition: Passive or adaptive response?Functional Ecology, 23, 612–620.

    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.

    Walck JL, Hidayati SN, Dixon KW, Thompson K, Poschlod P (2011). Climate change and plant regeneration from seed. Global Change Biology, 17, 2145–2161.

    Walters MB, Reich PB (2000). Seed size, nitrogen supply, and growth rate affect tree seedling survival in deep shade. Ecology, 81, 1887–1901.

    Wulff RD, Causin HF, Benitez O, Bacalini PA (1999). Intras-pecific variability and maternal effects in the response to nutrient addition in Chenopodium album. Canadian Journal of Botany, 77, 1150–1158.

    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.

    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.

    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.

This Article



Vol 42, No. 09, Pages 963-970

September 2018


Article Outline


  • 1 Materials and methods
  • 2 Results and analysis
  • 3 Discussion
  • References