Abstract
Background and aims
In recent years, ecologists have tried to determine the importance of intraspecific trait variation (ITV) versus community composition change (CCC) in shifts in community functional composition. However, results to date have not provided generality.
Methods
A six-year nitrogen (N) fertilization experiment was conducted in species-rich alpine meadow communities. We focused on five key functional traits (height, leaf dry mass content LDMC, specific leaf area SLA, leaf nitrogen content LNC, and leaf phosphorus content LPC) that belong to three trait categories (whole-plant level trait, leaf morphology trait and leaf chemical trait) and measured biomass of each species in community. A sum of squares decomposition method was used to distinguish the relative contribution of ITV versus CCC to community weighted mean (CWM) traits.
Results
Our results showed that N fertilization led to increasing heightCWM and LNCCWM due to light competition intensification and soil nutrient enrichment. However, the responses of community-wide SLA, LDMC and LPC were highly inconsistent and depended on the balance of different opposing processes. Moreover, during short-term fertilization, ITV played a more important role in mediating functional composition changes in all traits, but the effects of CCC overwhelmed ITV and became more important in determining community-wide whole-plant level trait and leaf morphology traits (height, LDMC and SLA) in subsequent fertilization years. On the other hand, ITV always played a more important role than CCC in determining the community-wide leaf chemical traits (LNC and LPC), but CCC had a greater contribution than ITV in terms of explaining shifts in whole-plant traits (height) and leaf morphology traits (LDMC and SLA). In addition, both positive and negative covariations appeared in our study, indicating that community-wide trait shifts due to ITV and CCC may either reinforce or oppose one another, depending on different trait categories.
Conclusions
These findings highlight the importance of ITV, CCC and their covariation in mediating community functional composition, and the relative importance of ITV effects and CCC effects depended on fertilization duration and trait category. This study significantly improve our understanding of the mechanisms governing the shifts in community functional composition under N deposition scenarios.
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References
Albert CH, Thuiller W, Yoccoz NG, Soudant A, Boucher F, Saccone P, Lavorel S (2010) Intraspecific functional variability: extent, structure and sources of variation. J Ecol 98:604–613
Auger S, Shipley B (2013) Inter-specific and intra-specific trait variation along short environmental gradients in an old-growth temperate forest. J Veg Sci 24:419–428
Bai W, Guo D, Tian Q, Liu N, Cheng W, Li L, Zhang WH (2015) Differential responses of grasses and forbs led to marked reduction in below-ground productivity in temperate steppe following chronic N deposition. J Ecol 103:1570–1579
Carmona CP, Guerrero I, Morales MB, Oñate JJ, Peco B, Llorens L (2017) Assessing vulnerability of functional diversity to species loss: a case study in Mediterranean agricultural systems. Funct Ecol 31:427–435
Cheplick G (1989) Nutrient availability, dimorphic seed production, and reproductive allocation in the annual grass Amphicarpum purshii. Can J Bot 67:2514–2521
Clark CM, Tilman D (2008) Loss of plant species after chronic low-level nitrogen deposition to prairie grasslands. Nature 451:712–715
Cornelissen J, Lavorel S, Garnier E, Diaz S, Buchmann N, Gurvich D, Reich P, Ter Steege H, Morgan H, Van Der Heijden M (2003) A handbook of protocols for standardised and easy measurement of plant functional traits worldwide. Aust J Bot 51:335–380
Craine JM, Froehle J, Tilman DG, Wedin DA, Chapin IFS (2001) The relationships among root and leaf traits of 76 grassland species and relative abundance along fertility and disturbance gradients. Oikos 93:274–285
de Bello F, Lavorel S, Albert CH, Thuiller W, Grigulis K, Dolezal J, Janeček Š, Lepš J (2011) Quantifying the relevance of intraspecific trait variability for functional diversity. Methods Ecol Evol 2:163–174
de la Riva EG, Pérez Ramos IM, Tosto A, Navarro Fernández CM, Olmo M, Marañón T, Villar R (2016) Disentangling the relative importance of species occurrence, abundance and intraspecific variability in community assembly: a trait-based approach at the whole-plant level in Mediterranean forests. Oikos 125:354–363
Fajardo A, Piper FI (2011) Intraspecific trait variation and covariation in a widespread tree species (Nothofaguspumilio) in southern Chile. New Phytol 189:259–271
Grime JP (2001) Plant strategies, vegetation processes, and ecosystem properties, 2nd edn. John Wiley and Sons, Chichester 417 p
Hautier Y, Niklaus PA, Hector A (2009) Competition for light causes plant biodiversity loss after eutrophication. Science 324:636–638
Isbell F, Reich PB, Tilman D, Hobbie SE, Polasky S, Binder S (2013) Nutrient enrichment, biodiversity loss, and consequent declines in ecosystem productivity. Proc Natl Acad Sci 110:11911–11916
Jung V, Albert CH, Violle C, Kunstler G, Loucougaray G, Spiegelberger T (2014) Intraspecific trait variability mediates the response of subalpine grassland communities to extreme drought events. J Ecol 102:45–53
Kazakou E, Violle C, Roumet C, Navas ML, Vile D, Kattge J, Garnier E (2014) Are trait-based species rankings consistent across data sets and spatial scales? J Veg Sci 25:235–247
Knops JM, Reinhart K (2000) Specific leaf area along a nitrogen fertilization gradient. Am Midl Nat 144:265–272
Lambers H, Poorter H (1992) Inherent variation in growth rate between higher plants: a search for physiological causes and ecological consequences. Adv Ecol Res 23:187–261
Lefcheck JS, Duffy JE (2015) Multitrophic functional diversity predicts ecosystem functioning in experimental assemblages of estuarine consumers. Ecology 96:2973–2983
Lepš J, de Bello F, Šmilauer P, Doležal J (2011) Community trait response to environment: disentangling species turnover vs intraspecific trait variability effects. Ecography 34:856–863
Li W, Cheng JM, Yu KL, Epstein HE, Guo L, Jing GH, Zhao J, Du GZ (2015) Plant functional diversity can be independent of species diversity: observations based on the impact of 4-yrs of nitrogen and phosphorus additions in an alpine meadow. PLoS One 10:e0136040
Li H, Yu K, Ratajczak Z, Nippert JB, Tondrob D, Xu D, Li W, Du G (2016) When variability outperforms the mean: trait plasticity predicts plant cover and biomass in an alpine wetland. Plant Soil 407:401–415
Luo Y, Qin G, Du G (2006) Importance of assemblage-level thinning: a field experiment in an alpine meadow on the Tibet plateau. J Veg Sci 17:417–424
Marks CO (2007) The causes of variation in tree seedling traits: the roles of environmental selection versus chance. Evolution 61:455–469
McGill BJ, Enquist BJ, Weiher E, Westoby M (2006) Rebuilding community ecology from functional traits. Trends Ecol Evol 21:178–185
Münzbergová Z, Hadincová V, Skálová H, Vandvik V, Bonser S (2017) Genetic differentiation and plasticity interact along temperature and precipitation gradients to determine plant performance under climate change. J Ecol 105:1358–1373
Newman E (1973) Competition and diversity in herbaceous vegetation. Nature 244:310
Niu K, Zhang S, Zhao B, Du G (2010) Linking grazing response of species abundance to functional traits in the Tibetan alpine meadow. Plant Soil 330:215–223
Niu K, Schmid B, Choler P, Du G (2012) Relationship between reproductive allocation and relative abundance among 32 species of a Tibetan alpine meadow: effects of fertilization and grazing. PLoS One 7:e35448
Niu KC, He JS, Lechowicz MJ (2016) Grazing-induced shifts in community functional composition and soil nutrient availability in Tibetan alpine meadows. J Appl Ecol 53:1554–1564
Pérez-Harguindeguy N, Díaz S, Garnier E, Lavorel S, Poorter H, Jaureguiberry P, Bret-Harte MS, Cornwell WK, Craine JM, Gurvich DE, Urcelay C, Veneklaas EJ, Reich PB, Poorter L, Wright IJ, Ray P, Enrico L, Pausas JG, de Vos AC, Buchmann N, Funes G, Quétier F, Hodgson JG, Thompson K, Morgan HD, ter Steege H, Sack L, Blonder B, Poschlod P, Vaieretti MV, Conti G, Staver AC, Aquino S, Cornelissen JHC (2013) New handbook for standardised measurement of plant functional traits worldwide. Aust J Bot 61:167
Poorter H, Nagel O (2000) The role of biomass allocation in the growth response of plants to different levels of light, CO2, nutrients and water: a quantitative review. Funct Plant Biol 27:1191–1191
R Development Core Team (2013) R: A Language and Environment for Statistical Computing. In: R Foundation for Statistical Computing, Vienna, Austria. http://www.R-project.org/
Riva EG, Pérez Ramos IM, Tosto A, Navarro Fernández CM, Olmo M, Marañón T, Villar R (2015) Disentangling the relative importance of species occurrence, abundance and intraspecific variability in community assembly: a trait-based approach at the whole-plant level in Mediterranean forests. Oikos 125(3):354–363
Rozendaal DMA, Hurtado VH, Poorter L (2006) Plasticity in leaf traits of 38 tropical tree species in response to light: relationships with light demand and adult stature. Funct Ecol 20:207–216
Ryser P, Lambers H (1995) Root and leaf attributes accounting for the performance of fast- and slow-growing grasses at different nutrient supply. Plant Soil 170:251–265
Schellberg J, Pontes LS (2012) Plant functional traits and nutrient gradients on grassland. Grass Forage Sci 67:305–319
Schneider CA, Rasband WS, Eliceiri KW (2012) NIH image to image J: 25 years of image analysis. Nat Methods 9:671–675
Schwinning S, Sala OE (2004) Hierarchy of responses to resource pulses in arid and semi-arid ecosystems. Oecologia 141:211–220
Siefert A, Ritchie ME (2016) Intraspecific trait variation drives functional responses of old-field plant communities to nutrient enrichment. Oecologia 181:245–255
Siefert A, Violle C, Chalmandrier L, Albert CH, Taudiere A, Fajardo A, Aarssen LW, Baraloto C, Carlucci MB, Cianciaruso MV, de LDantas V, de Bello F, Duarte LDS, Fonseca CR, Freschet GT, Gaucherand S, Gross N, Hikosaka K, Jackson B, Jung V, Kamiyama C, Katabuchi M, Kembel SW, Kichenin E, Kraft NJB, Lagerström A, Bagousse Pinguet YL, Li Y, Mason N, Messier J, Nakashizuka T, Overton JM, Peltzer DA, Pérez-Ramos IM, Pillar VD, Prentice HC, Richardson S, Sasaki T, Schamp BS, Schöb C, Shipley B, Sundqvist M, Sykes MT, Vandewalle M, Wardle DA (2015) A global meta-analysis of the relative extent of intraspecific trait variation in plant communities. Ecol Lett 18:1406–1419
Smith MD, Knapp AK, Collins SL (2009) A framework for assessing ecosystem dynamics in response to chronic resource alterations induced by global change. Ecology 90:3279–3289
Treseder KK, Vitousek PM (2001) Effects of soil nutrient availability on investment in acquisition of n and p in hawaiian rain forests. Ecology 82:946–954
Vicente-Chandler J, Silva S, Figarella J (1959) The effect of nitrogen fertilization and frequency of cutting on the yield and composition of three tropical grasses. Agron J 51:202–206
Volf M, Redmond C, Albert ÁJ, Bagousse-Pinguet Y, Biella P, Götzenberger L, Hrázský Z, Janeček Š, Klimešová J, Lepš J, Šebelíková L, Vlasatá T, Bello F (2016) Effects of long- and short-term management on the functional structure of meadows through species turnover and intraspecific trait variability. Oecologia 180(4):1–10
Weiher E, van der Werf A, Thompson K, Roderick M, Garnier E, Eriksson O (1999) Challenging Theophrastus: a common core list of plant traits for functional ecology. J Veg Sci 10:609–620
Westoby M, Wright IJ (2006) Land-plant ecology on the basis of functional traits. Trends Ecol Evol 21:261–268
Wright IJ, Reich PB, Westoby M, Ackerly DD, Baruch Z, Bongers F, Cavender-Bares J, Chapin T, Cornelissen JHC, Diemer M, Flexas J, Garnier E, Groom PK, Gulias J, Hikosaka K, Lamont BB, Lee T, Lee W, Lusk C, Midgley JJ, Navas M-L, Niinemets U, Oleksyn J, Osada N, Poorter H, Poot P, Prior L, Pyankov VI, Roumet C, Thomas SC, Tjoelker MG, Veneklaas EJ, Villar R (2004) The worldwide leaf economics spectrum. Nature 428:821–827
Yang Z, Ruijven J, Du G (2011) The effects of long-term fertilization on the temporal stability of alpine meadow communities. Plant Soil 345:315–324
Yang Z, Guo H, Zhang J, Du G (2013) Stochastic and deterministic processes together determine alpine meadow plant community composition on the Tibetan plateau. Oecologia 171:495–504
Zhang P, Zhou X, Li J, Guo Z, Du G (2015) Space resource utilisation: a novel indicator to quantify species competitive ability for light. Sci Rep 5:16832
Zhou X, Wang Y, Zhang P, Guo Z, Chu C, Du G (2016) The effects of fertilization on the trait–abundance relationships in a Tibetan alpine meadow community. J Plant Ecol 9(2):144–152
Zhou X, Guo Z, Zhang P, Li H, Chu C, Li X, Du G (2017) Different categories of biodiversity explain productivity variation after fertilization in a Tibetan alpine meadow community. Ecol Evol 7:3464–3474
Acknowledgements
We would like to thank the staff of the Research Station of Alpine Meadow and Wetland Ecosystems at Lanzhou University for providing invaluable field assistance. The study was supported by the National Nature Science Foundation of China (Grant No.31700355), the Scientific Research Startup Foundation for Doctors of Xinjiang University (Grant No. BS160260).
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XLZ and GZD conceived and designed the experiments. XLZ, PFZ and ZG performed the experiments. XLZ analyzed the data. XLZ wrote the manuscript; other authors provided editorial advice.
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Zhou, X., Guo, Z., Zhang, P. et al. Shift in community functional composition following nitrogen fertilization in an alpine meadow through intraspecific trait variation and community composition change. Plant Soil 431, 289–302 (2018). https://doi.org/10.1007/s11104-018-3771-x
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DOI: https://doi.org/10.1007/s11104-018-3771-x