Abstract
Little is known about the consequences of phenotypic plasticity in co-existing species for plant community structure. However, it has been proposed that the potential of plants to exhibit plastic responses to light availability could be a key factor determining the capability of individuals to co-exist at small scales. Our previous research demonstrated that morphological plasticity to light was positively related to small-scale species richness in a temperate grassland. However, it remained unclear whether this relationship was solely due to a higher shoot density in plastic assemblages, or whether diversity was directly related to the morphological plasticity of the co-inhabitants. We used two data sets to clarify this relationship: experimentally acquired estimates of plasticity to light availability for 45 herbaceous plant species, and species richness and ramet density data from a 2-year permanent plot study in a semi-natural calcareous grassland. There was little ramet mortality observed in the permanent plot study indicating that the link between plasticity and richness does not operate through reduced mortality in more morphologically plastic assemblages. The local density of ramets explained most of variation in small-scale richness, but there was also a significant direct density-independent effect of mean shoot plasticity on richness, showing that plasticity to light directly enhances the small-scale co-existence of species.
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References
Callaway RM, Pennings SC, Richards CL (2003) Phenotypic plasticity and interactions among plants. Ecology 84:1115–1128. doi:10.1890/0012-9658(2003)084[1115:ppaiap]2.0.co;2
Casal JJ, Mazzella MA (1998) Conditional synergism between cryptochrome 1 and phytochrome B is shown by the analysis of phyA, phyB, andhy4 simple, double, and triple mutants in Arabidopsis. Plant Physiol 118:19–25
de Bello F, Janeček Š, Lepš J, Doležal J, Macková J, Lanta V, Klimešová J, Bruun HH (2012) Different plant trait scaling in dry versus wet Central European meadows. J Veg Sci 23:709–720. doi:10.1111/j.1654-1103.2012.01389.x
Díaz S, Cabido M, Casanoves F (1998) Plant functional traits and environmental filters at a regional scale. J Veg Sci 9:113–122. doi:10.2307/3237229
Eek L, Zobel K (2001) Structure and diversity of a species-rich grassland community, treated with additional illumination, fertilization and mowing. Ecography 24:157–164. doi:10.1034/j.1600-0587.2001.240206.x
Franklin KA (2008) Shade avoidance. New Phytol 179:930–944. doi:10.1111/j.1469-8137.2008.02507.x
Götzenberger L, de Bello F, Bråthen K, Davison J, Dubuis A, Guisan A, Lepš J, Lindborg R, Moora M, Pärtel M, Pellissier L, Pottier J, Vittoz P, Zobel K, Zobel M (2012) Ecological assembly rules in plant communities-approaches, patterns and prospects. Biol Rev Camb Philos Soc 87:111–127. doi:10.1111/j.1469-185X.2011.00187.x
Herben T, Novoplansky A (2010) Fight or flight: plastic behavior under self-generated heterogeneity. Evol Ecol 24:1521–1536. doi:10.1007/s10682-010-9386-1
Jung V, Violle C, Mondy C, Hoffmann L, Muller S (2010) Intraspecific variability and trait-based community assembly. J Ecol 98:1134–1140. doi:10.1111/j.1365-2745.2010.01687.x
Keuskamp DH, Pollmann S, Voesenek LA, Peeters A, Pierik R (2010) Auxin transport through PIN-FORMED 3 (PIN3) controls shade avoidance and fitness during competition. Proc Natl Acad Sci USA 107:22740–22744. doi:10.1073/pnas.1013457108
Kline RB (2011) Principles and practice of structural equation modeling, 3rd edn. Guilford Press, New York
Kull K, Zobel M (1991) High species richness in an Estonian wooded meadow. J Veg Sci 2:711–714. doi:10.2307/3236182
Lankau RA (2011) Rapid evolutionary change and the coexistence of species. Annu Rev Ecol Evol Syst 42:335–354. doi:10.1146/annurev-ecolsys-102710-145100
Lepik M, Liira J, Zobel K (2005) High shoot plasticity favours plant coexistence in herbaceous vegetation. Oecologia 145:465–474. doi:10.1007/s00442-005-0142-0
Leyser O (2009) The control of shoot branching: an example of plant information processing. Plant Cell Environ 32:694–703. doi:10.1111/j.1365-3040.2009.01930.x
Maddonni GA, Otegui ME, Andrieu B, Chelle M, Casal JJ (2002) Maize leaves turn away from neighbors. Plant Physiol 130:1181–1189. doi:10.1104/pp.009738
McConnaughay K, Coleman J (1999) Biomass allocation in plants: ontogeny or optimality? A test along three resource gradients. Ecology 80:2581–2593. doi:10.1890/0012-9658(1999)080[2581:baipoo]2.0.co;2
Nagashima H, Hikosaka K (2011) Plants in a crowded stand regulate their height growth so as to maintain similar heights to neighbours even when they have potential advantages in height growth. Ann Bot 108:1–8. doi:10.1093/aob/mcr109
Novoplansky A (2009) Picking battles wisely: plant behaviour under competition. Plant Cell Environ 32:726–741. doi:10.1111/j.1365-3040.2009.01979.x
Pierik R, Mommer L, Voesenek LA (2012) Molecular mechanisms of plant competition: neighbour detection and response strategies. Funct Ecol 27:841–853. doi:10.1111/1365-2435.12010
Pigliucci M, Schmitt J (1999) Genes affecting phenotypic plasticity in Arabidopsis: pleiotropic effects and reproductive fitness of photomorphogenic mutants. J Evol Biol 12:551–562. doi:10.1046/j.1420-9101.1999.00057.x
R Core Team (2014) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. http://www.R-project.org/
Rosseel Y (2012) lavaan: an R package for structural equation modeling. J Stat Softw 48:1–36. http://www.jstatsoft.org/v48/i02/
Schmitt J, Wulff RD (1993) Light spectral quality, phytochrome and plant competition. Trends Ecol Evol 8:47–51. doi:10.1016/0169-5347(93)90157-k
Schwinning S, Weiner J (1998) Mechanisms determining the degree of size asymmetry in competition among plants. Oecologia 113:447–455. doi:10.1007/s004420050397
Semchenko M, Zobel K, Hutchings MJ (2010) To compete or not to compete: an experimental study of interactions between plant species with contrasting root behaviour. Evol Ecol 24:1433–1445. doi:10.1007/s10682-010-9401-6
Semchenko M, Lepik M, Götzenberger L, Zobel K (2012) Positive effect of shade on plant growth: amelioration of stress or active regulation of growth rate? J Ecol 100:459–466. doi:10.1111/j.1365-2745.2011.01936.x
Skálová H (2010) Potential and constraints for grasses to cope with spatially heterogeneous radiation environments. Plant Ecol 206:115–125. doi:10.1007/s11258-009-9628-x
Sorrensen-Cothern KA, Ford ED, Sprugel DG (1993) A model of competition incorporating plasticity through modular foliage and crown development. Ecol Monogr 63:277–304. doi:10.2307/2937102
Stoll P, Weiner J, Muller-Landau H, Müller E, Hara T (2002) Size symmetry of competition alters biomass–density relationships. Proc R Soc Lond Ser B Biol Sci 269:2191–2195. doi:10.1098/rspb.2002.2137
Stuefer JF, Huber H (1998) Differential effects of light quantity and spectral light quality on growth, morphology and development of two stoloniferous Potentilla species. Oecologia 117:1–8. doi:10.1007/s004420050624
Sultan SE (1995) Phenotypic plasticity and plant adaptation. Acta Bot Neerl 44:363–383. doi:10.1111/j.1438-8677.1995.tb00793.x
Vellend M (2010) Conceptual synthesis in community ecology. Q Rev Biol 85:183–206. doi:10.1086/652373
Vermeulen PJ, Anten NP, Schieving F, Werger MJ, During HJ (2008) Height convergence in response to neighbour growth: genotypic differences in the stoloniferous plant Potentilla reptans. New Phytol 177:688–697. doi:10.1111/j.1469-8137.2007.02301.x
Wacker L, Baudois O, Eichenberger-Glinz S, Schmid B (2009) Effects of plant species richness on stand structure and productivity. J Plant Ecol 2:95–106. doi:10.1093/jpe/rtp010
Weiner J, Berntson GM, Thomas SC (1990) Competition and growth form in a woodland annual. J Ecol 78:459–469. doi:10.2307/2261124
Weinig C (2000) Plasticity versus canalization: population differences in the timing of shade-avoidance responses. Evolution 54:441–451. doi:10.1554/0014-820(2000)054[0441:pvcpdi]2.0.co;2
Acknowledgments
We are grateful to all members of the working group for their help with the experiments and two anonymous referees for valuable comments. Special thanks to Anu Lepik and Oliver Parrest for assistance with fieldwork. Jaan Liira helped with the statistical analysis. Two anonymous reviewers and editors provided numerous comments that helped significantly to improve the manuscript. The study was supported by the Estonian Science Foundation (5535, 7576, 9269) and the University of Tartu (IUT 20-31). The experiments comply with the current laws of the Republic of Estonia.
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Communicated by Bryan Foster.
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Lepik, M., Zobel, K. Is the positive relationship between species richness and shoot morphological plasticity mediated by ramet density or is there a direct link?. Oecologia 178, 867–873 (2015). https://doi.org/10.1007/s00442-015-3288-4
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DOI: https://doi.org/10.1007/s00442-015-3288-4