Reproductive allocation of Carex flava reacts differently to competition and resources in a designed plant mixture of five species
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Abstract
In natural plant communities, reproductive allocation can be affected by complex interactions among abiotic resources, species competition and plant size. This topic was addressed using a variety of designed mixed stands of five species (Carex elata, Carex flava, Lycopus europaeus, Lysimachia vulgaris and Mentha aquatica) under four abiotic conditions to investigate how competition and abiotic resources influence the reproductive allocation of one of the five species, C. flava. The plant mixtures varied systematically in both the relative abundance of the five species and the absolute density, and were each established with two levels of water and nutrients. In total, 176 mixtures were maintained for two growing seasons in large pots in an experimental garden. Reproductive allocation of C. flava increased from 6.8% to 9.7% under high nutrient application; however, for both nutrient levels, reproductive allocation was independent of shoot mass (size-independent allocation). Under low competition, reproductive allocation of C. flava decreased as its shoot mass increased, indicating a relatively high investment in vegetative structures under higher light availability. However, under strong competition, the allocation pattern changed and a constant reproductive allocation for different plant sizes was observed. Different water levels did not influence the shoot mass, seed mass or reproductive allocation of C. flava, indicating that the species was not stressed under dryer conditions. When under competitive pressure, however, the species responded with reduced shoot and seed production under more favourable water conditions. This behaviour indicates a trade-off between the ability to tolerate stress and the competitive and reproductive response of C. flava. In conclusion, C. flava was adversely affected by competition with some of the species, and competition, mediated by plant size, indirectly affected reproductive allocation. C. flava was able to modify its allocation pattern depending on the available resources and retained its reproductive allocation even under unfavourable conditions for varying plant sizes, which is interpreted as an advantageous reaction to greater competition pressure.
Keywords
Fen meadow species Nutrient water supply Simplex design Stress tolerance Trade-offNotes
Acknowledgements
I am grateful to T. Steffen for weighing the seed and biomass samples. I would also like to thank M. Fotsch, P. Borer and P. Kadelbach for assistance in practical work and K. Seipel for linguistic improvements. J. Connolly, P. Edwards, S. Guesewell, D. Ramseier and two anonymous reviewers provided helpful comments on an earlier version of the manuscript. The project was funded by the Swiss Federal Institute of Technology, Zurich (Grant No. 0-20891-01).
Supplementary material
References
- Aeschimann D, Heitz C (2005) Synonymie-Index der Schweizer Flora. Zentrum des Datenverbundnetzes der Schweizer Flora, BernGoogle Scholar
- Arenas F, Viejo RM, Fernandez C (2002) Density-dependent regulation in an invasive seaweed: responses at plant and modular levels. J Ecol 90:820–829. doi: 10.1046/j.1365-2745.2002.00720.x CrossRefGoogle Scholar
- Campbell BD, Grime JP (1992) An experimental test of plant strategy theory. Ecology 73:15–29. doi: 10.2307/1938717 CrossRefGoogle Scholar
- Cheplick GP (2001) Quantitative genetics of mass allocation and the allometry of reproduction in Amaranthus albus: relation to soil nutrients. Int J Plant Sci 162:807–816. doi: 10.1086/320778 CrossRefGoogle Scholar
- Chiariello NR, Gulmon SL (1991) Stress effects on plant reproduction. In: Mooney HA, Winner WE, Pell EJ et al (eds) Response of plants to multiple stresses. Academic Press, San Diego, pp 161–188Google Scholar
- Cornell JA (2002) Experiments with mixtures. Wiley, New York, USAGoogle Scholar
- Edelkraut KA, Guesewell S (2006) Progressive effects of shading on experimental wetland communities over three years. Plant Ecol 183:315–327. doi: 10.1007/s11258-005-9042-y CrossRefGoogle Scholar
- Ellenberg H (1996) Vegetation Mitteleuropas mit den Alpen. Ulmer, Stuttgart, DeutschlandGoogle Scholar
- Emery SM, Gross KL (2007) Dominant species identity, not community evenness, regulates invasion in experimental grassland plant communities. Ecology 88:954–964. doi: 10.1890/06-0568 PubMedCrossRefGoogle Scholar
- Grime JP (2001) Plant strategies, vegetation processes, and ecosystem properties. Wiley, Chichester, UKGoogle Scholar
- Hara T, Kawano S, Nagai Y (1988) Optimum reproductive strategy of plants, with special reference to the modes of reproductive resource allocation. Plant Species Biol 3:43–59. doi: 10.1111/j.1442-1984.1988.tb00170.x CrossRefGoogle Scholar
- Harper JL, Ogden J (1970) The reproductive strategy of higher plants. I. The concept of strategy with special reference to Senecio vulgaris L. J Ecol 58:681–698. doi: 10.2307/2258529 CrossRefGoogle Scholar
- Hendriks AJ, Mulder C (2008) Scaling of offspring number and mass to plant and animal size: model and meta-analysis. Oecologia 155:705–716. doi: 10.1007/s00442-007-0952-3 PubMedCrossRefGoogle Scholar
- Karlsson PS, Méndez M (2005) The resource economy in plant reproduction. In: Reekie EG, Bazzaz FA (eds) Reproductive allocation in plants. Elsevier, Amsterdam, pp 1–49CrossRefGoogle Scholar
- Klinkhamer PGL, de Jong TJ, Nell HW (1994) Limiting factors for seed production and phenotypic gender in the gynodioecious species Echium vulgare (Boraginaceae). Oikos 71:469–478. doi: 10.2307/3545835 CrossRefGoogle Scholar
- Lauber K, Wagner G (2007) Flora Helvetica. Paul Haupt, Bern, SchweizGoogle Scholar
- Liancourt P, Callaway RM, Michalet R (2005) Stress tolerance and competitive-response ability determine the outcome of biotic interactions. Ecology 86:1611–1618. doi: 10.1890/04-1398 CrossRefGoogle Scholar
- Méndez M, Karlsson PS (2004) Between-population variation in size-dependent reproduction and reproductive allocation in Pinguicula vulgaris (Lentibulariaceae) and its environmental correlates. Oikos 104:59–70. doi: 10.1111/j.0030-1299.2004.12335.x CrossRefGoogle Scholar
- Pauli D, Peintinger M, Schmid B (2002) Nutrient enrichment in calcareous fens: effects on plant species and community structure. Basic Appl Ecol 3:255–266. doi: 10.1078/1439-1791-00096 CrossRefGoogle Scholar
- Ramseier D, Connolly J, Bazzaz FA (2005) Carbon dioxide regime, species identity and influence of species initial abundance as determinants of change in stand biomass composition in five-species communities: an investigation using a simplex design and RGRD analysis. J Ecol 93:502–511. doi: 10.1111/j.1365-2745.2005.00999.x CrossRefGoogle Scholar
- R Development Core Team (2007) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. ISBN 3-900051-07-0, http://www.R-project.org
- Reekie EG (1998) An explanation for size-dependent reproductive allocation in Plantago major. Can J Bot 76:43–50. doi: 10.1139/cjb-76-1-43 CrossRefGoogle Scholar
- Reekie EG, Bazzaz FA (eds) (2005) Reproductive allocation in plants. Elsevier, Amsterdam, NetherlandsGoogle Scholar
- Sadras VO, Bange MP, Milroy SP (1997) Reproductive allocation of cotton in response to plant and environmental factors. Ann Bot (Lond) 80:75–81. doi: 10.1006/anbo.1997.0402 CrossRefGoogle Scholar
- Schmid B (1984) Life histories in clonal plants of the Carex flava group. J Ecol 72:93–114. doi: 10.2307/2260008 CrossRefGoogle Scholar
- Schmid B (1986) Colonizing plants with persistent seeds and persistent seedlings (Carex flava group). Bot Helv 96:19–26Google Scholar
- Sletvold N (2002) Effects of plant size on reproductive output and offspring performance in the facultative biennial Digitalis purpurea. J Ecol 90:958–966. doi: 10.1046/j.1365-2745.2002.00725.x CrossRefGoogle Scholar
- Suding KN, Goldberg DE, Hartman KM (2003) Relationships among species traits: separating levels of response and identifying linkages to abundance. Ecology 84:1–16. doi: 10.1890/0012-9658(2003)084[0001:RASTSL]2.0.CO;2 CrossRefGoogle Scholar
- Sugiyama S, Bazzaz FA (1998) Size dependence of reproductive allocation: the influence of resource availability, competition and genetic identity. Funct Ecol 12:280–288. doi: 10.1046/j.1365-2435.1998.00187.x CrossRefGoogle Scholar
- Susko DJ, Lovett DL (2000) Plant-size and fruit-position effects on reproductive allocation in Alliaria petiolata (Brassicaceae). Can J Bot 78:1398–1407. doi: 10.1139/cjb-78-11-1398 CrossRefGoogle Scholar
- Suter M, Ramseier D, Guesewell S et al (2007) Convergence patterns and multiple species interactions in a designed plant mixture of five species. Oecologia 151:499–511. doi: 10.1007/s00442-006-0594-x PubMedCrossRefGoogle Scholar
- van der Hoek D, van Mierlo AJE, van Groenendael J (2004) Nutrient limitation and nutrient-driven shifts in plant species composition in a species-rich fen meadow. J Veg Sci 15:389–396. doi: 10.1658/1100-9233(2004)015[0389:NLANSI]2.0.CO;2 CrossRefGoogle Scholar
- Weiner J (1988) The influence of competition on plant reproduction. In: Lovett Doust L (ed) Plant reproductive ecology. Oxford University Press, New York, pp 228–245Google Scholar
- Whitfield CP, Davison AW, Ashenden TW (1998) The effects of nutrient limitation on the response of Plantago major to ozone. New Phytol 140:219–230. doi: 10.1046/j.1469-8137.1998.00277.x CrossRefGoogle Scholar