Regina BERJANO, Montserrat ARISTA, María TALAVERA, María Jesús ARIZA, Pedro Luis ORTIZ

Plasticity and within plant sex-ratio variation in monoecious Emex spinosa

Understanding how phenotype trait expression varies at different levels of biological organization is crucial to understanding the fundamental ecological, evolutionary, and developmental issues that contribute to adaptive plasticity. This is particularly important in plants because of their modular organization. Here, we study plasticity in flower production in response to light intensity at 3 levels of biological organization (between populations, among individuals, and within individuals) in Emex spinosa (L.) Compd., a Mediterranean amphicarpic annual. Plants exposed to high light intensity grew faster and produced more flowers than plants at low light intensity. Female flower production was the most labile trait. Within individuals, the production of female and male flowers, and the floral sex ratio strongly varied along the stems. Light intensity only significantly affected flower production at central and terminal positions in the inflorescence, while sex ratio was invariable to light intensity in each position, reflecting a sexual specialization independently of light, from initial femaleness towards maleness at the end of flowering. In Emex spinosa, the production of selfing fruits seems not only to concern ground fruits, but also early aerial fruits.

Anahtar Kelimeler:

Key words: Amphicarpy, flower production, modularity, phenotypic plasticity, sex ratio

Plasticity and within plant sex-ratio variation in monoecious Emex spinosa

Keywords:

Key words: Amphicarpy, flower production, modularity, phenotypic plasticity, sex ratio,

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Armbruster WS (1991). Multilevel analysis of morphometric data from natural plant populations: insights into ontogenetic, genetic, and selective correlations in Dalechampia scandens. Evolution, 45: 1229–1244.
Baker HG (1974). The evolution of weeds. Annu Rev Ecol Syst 5: 1–
Bell DL, Galloway LF (2007). Plasticity to neighbour shade: fitness consequences and allometry. Funct Ecol 21: 1146–1153.
Bell DL, Galloway LF (2008). Population differentiation for plasticity to light in an annual herb: adaptation and cost. Am J Bot 95: 59–
Bertin RI (2007). Sex allocation in Carex (Cyperaceae): effects of light, water, and nutrients. Can J Bot 85: 377–384.
Bertin RI, Kerwin MA (1998). Floral sex ratios and gynomonoecy in Aster (Asteraceae). Am J Bot 85: 235–244.
Bickel AM, Freeman DC (1993). Effects of pollen vector and plant geometry on floral sex ratio in monoecious plants. Am Midl Nat 130: 239–247.
Bissell EK, Diggle PK (2008). Floral morphology in Nicotiana: architectural and temporal effects on phenotypic integration. Int J Plant Sci, 169: 225–240.
Burd M, Allen TFH (1988). Sexual allocation strategy in wind pollinated plants. Evolution 42: 403–407.
Charlesworth D, Morgan MT (1991). Allocation of resources to sex functions in flowering plants. Philos T Roy Soc B 332: 91–102.
Charnov EL (1982). The theory of sex allocation. Princeton, NJ, USA: Princeton University Press.
Charnov EL, Bull J (1977). When is sex environmentally determined? Nature 266: 829–830.
Cheplick GP (1987). The ecology of amphicarpic plants. Trends Ecol Evol 2: 97–101.
Cheplick GP (1989). Nutrient availability, dimorphic seed production, and reproductive allocation in the annual grass Amphicarpum purshii. Can J Bot 67: 2514–2521.
Cid-Benevento CR (1987). Relative effects of light, soil moisture availability and vegetative size on sex ratio of two monoecious woodland annual herbs: Acalypha rhomboidea (Euphorbiaceae) and Pilea pumila (Urticaceae). B Torrey Bot Club 114: 293–306.
Cohen D (1966). Optimizing reproduction in a randomly varying environment. J Theor Biol 12: 119–129.
Coleman JS, McConnaughay KDM, Ackerly DD (1994). Interpreting phenotypic variation in plants. Trends Ecol Evol 9: 187–191.
Davidson AM, Jennions M, Nicotra AB (2011). Do invasive species show higher phenotypic plasticity than native species and, if so, is it adaptive? A meta-analysis. Ecol Lett 14: 419–431. de Kroon H, Huber H, Stuefer JF, Groenendael JM (2005). A modular concept of phenotypic plasticity in plants. New Phytol 166: 73–
DeWitt TJ, Langerhans RB (2004). Integrated solutions to environmental heterogeneity. In: DeWitt TJ, Scheiner SM, editors Phenotipic plasticity: functional and conceptual approaches. New York, USA: Oxford University Press, pp. 98–111.
Diggle PK (1994). The expression of andromonoecy in Solanum hirtum (Solanaceae): phenotypic plasticity and ontogenic contingency. Am J Bot 81: 1354–1365.
Dorken ME, Barrett SCH (2004). Phenotypic plasticity of vegetative and reproductive traits in monoecious and dioecious populations of Sagittaria latifolia (Alismataceae): a clonal aquatic plant. J Ecol 92: 32–44.
Dorn LA, Pyle EH, Schmitt J (2000). Plasticity to light cues and resources in Arabidopsis thaliana: testing for adaptive value and costs. Evolution 54: 1982–1994.
Droste T, Flory SL, Clay K (2010). Variation for phenotypic plasticity among populations of an invasive exotic grass. Plant Ecol 207: 297–306.
Franklin KA, Whitelam GC (2005). Phytochromes and shadeavoidance responses in plants. Ann Bot-London 96: 169–175.
Freeman DC, McArthur ED, Harper KT, Blauer AC (1981). Influence of environment on the floral sex-ratio of monoecious plants. Evolution 35: 194–197.
Friedman J, Barrett SCH (2011). Genetic and environmental control of temporal and size-dependent sex allocation in a windpollinated plant. Evolution 65: 2061–2074.
Funk JL (2008). Differences in plasticity between invasive and native plants from a low resource environment. J Ecol 96: 1162–1173.
Galloway LF, Etterson JR (2009). Plasticity to canopy shade in a monocarpic herb: within- and between-generation effects. New Phytol 182: 1003–1012.
Ghalambor CK, McKay JK, Carroll SP, Reznick DN (2007). Adaptive versus non-adaptive phenotypic plasticity and the potential for contemporary adaptation in new environments. Funct Ecol 21: 394–407.
Goldman DA, Willson MF (1986). Sex allocation in functionally hermaphroditic plants: a review and critique. Bot Rev 52: 157–194.
Gregg KB (1978). The interaction of light intensity, plant size, and nutrition in sex expression in Cycnoches (Orchidaceae). Selbyana: 212–223.
Guillespie JH (1977). Natural selection for variance in offspring numbers: a new evolutionary principal. Am Nat 111: 1010– 10
Gupta S, Narayan R (2012). Phenotypic plasticity of Chenopodium murale across contrasting habitat conditions in peri-urban areas in Indian dry tropics: is it indicative of its invasiveness? Plant Ecol 213: 493–503.
Hameed M, Nawaz T, Ashraf M, Naz N, Batool R, Ahmad MSA, Riaz A (2013). Physioanatomical adaptations in response to salt stress in Sporobolus arabicus (Poaceae) from the Salt Range, Pakistan. Turk J Bot 37: 715–724.
Harper JL (1977). Population biology of plants. London, UK: Academic Press.
Korpelainen H (1998). Labile sex expression in plants. Biol Rev 73: 157–180.
Kutbay HG, Uçkan F (1998). Phenotypic plasticity in Turkish Commelina communis L. (Commelinaceae) populations. Turk J Bot 22: 199–204.
Lemen C (1980). Allocation of reproductive effort to the male and female strategies in wind-pollinated plants. Oecologia 45: 156–159.
Liang KY, Zeger SL (1986). Longitudinal data analysis using generalized linear models. Biometrika 73: 13–22.
Lloyd DG (1979). Parental strategies of angiosperms. New Zeal J Bot 17: 595–606.
Lloyd DG, Bawa KS (1984). Modification of the gender of seed plants in varying conditions. Evol Biol 17: 255–338.
Mancuso E, Peruzzi L (2010). Male individuals in cultivated Fritillaria persica L. (Liliaceae): real androdioecy or gender disphasy. Turk J Bot 34: 435–440.
Menges ES (1990). Environmental correlations with male, female and clonal biomass allocation in the forest herb, Laportea canadensis. Am Midl Nat 124: 171–180.
Molina-Montenegro MA, Palma-Rojas C, Alcayaga-Olivares Y, Oses R, Corcuera LJ, Cavieres LA, Gianoli E (2013). Ecophysiological plasticity and local differentiation help explain the invasion success of Taraxacum officinale (dandelion) in South America. Ecography 36: 718–730.
Narbona E, Dirzo R (2010). Experimental defoliation affects male but not female reproductive performance of the tropical monoecious plant Croton suberosus (Euphorbiaceae). Ann Bot-London 106: 359–369.
Niklas KJ (1985). The aerodynamics of wind pollination. Bot Rev 51: 328–386.
Oliva G, Martinez A, Collantes M, Dubcovsky J (1993). Phenotypic plasticity and contrasting habitat colonization in Festuca pallescens. Can J Bot 71: 970–977.
Ortiz PL, Berjano R, Talavera M, Arista M (2009). The role of resources and architecture in modeling floral variability for the monoecious amphicarpic Emex spinosa (Polygonaceae). Am J Bot 96: 2062–2073.
Petit C, Thompson JD (1998). Phenotypic selection and population differentiation in relation to habitat heterogeneity in Arrhenatherum elatius. J Ecol 86: 829–840.
Price SC, Jain SK (1981). Are inbreeders better colonizers? Oecologia 49: 283–286.
Richards CL, Bossdorf O, Muth NZ, Gurevitch J, Pigliucci M (2006). Jack of all trades, master of some? On the role of phenotypic plasticity in plant invasions. Ecol Lett 9: 981–993.
Sadeh A, Guterman H, Gersani M, Ovadia O (2009). Plastic bethedging in an amphicarpic annual: an integrated strategy under variable conditions. Evol Ecol 23: 373–388.
Scheiner SM (1993). Genetics and evolution of phenotypic plasticity. Annu Rev Ecol Syst 24: 35–68.
Schlichting CD (1986). The evolution of phenotypic plasticity in plants. Annu Rev Ecol Syst 17: 667–693.
Schlichting CD, Pigliucci M (1995). Gene-regulation, quantitative genetics and the evolution of reaction norms. Evol Ecol 9: 154–168.
Schmitt J, Stinchcombe JR, Heschel MS, Huber H (2003). The adaptive evolution of plasticity: phytochrome-mediated shade avoidance responses. Integr Comp Biol 43: 459–469.
Shakarishvili N, Oshishvili L (2013). Sexual phenotype of Capparis herbacea (Capparaceae). Turk J Bot 37: 682–689.
Smith H, Whitelam GC (1997). The shade avoidance syndrome: Multiple responses mediated by multiple phytochromes. Plant Cell Environ 20: 840–844.
Solomon BP (1985). Environmentally influenced changes in sex expression in an andromonoecious plant. Ecology 66: 1321– 13
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.
Sultan SE (2001). Phenotypic plasticity for fitness components in Polygonum species of contrasting ecological breadth. Ecology 82: 328–343.
Sultan SE (2003). Phenotypic plasticity in plants: a case study in ecological development. Evol Dev 5: 25–33.
Sultan SE (2004). Promising directions in plant phenotypic plasticity. Perspect Plant Ecol 6: 227–233.
Sultan SE, Bazzaz FA (1993). Phenotypic plasticity in Polygonum persicaria. 1. Diversity and uniformity in genotypic norms of reaction to light. Evolution 47: 1009–1031.
Sultan SE, Stearns SC (2005). Environmentally contingent variation: phenotypic plasticity and norms of reaction. In: Hallgrimsson B, Hall B, editors. Variation: a hierarchical examination of a central concept in biology. Boston, USA: Elsevier Academic Press, pp. 303–327.
Talavera M, Ortiz PL, Arista M, Berjano R, Imbert E (2010). Disentangling sources of maternal effects in the heterocarpic species Rumex bucephalophorus. Perspect Plant Ecol 12: 295– 30
Thompson JD (1991). Phenotypic plasticity as a component of evolutionary change. Trends Ecol Evol: 246–249. van Kleunen M, Fischer M (2005). Constraints on the evolution of adaptive phenotypic plasticity in plants. New Phytol 166: 49–
Venable DL, Dyreson E, Morales E (1995). Population-dynamic consequences and evolution of seed traits of Heterosperma pinnatum (Asteraceae). Am J Bot 82: 410–420.
Weijschede J, Martinkova J, de Kroon H, Huber H (2006). Shade avoidance in Trifolium repens: costs and benefits of plasticity in petiole length and leaf size. New Phytol 172: 655–666.
Weiss PW (1980). Germination, reproduction and interference in the amphicarpic annual Emex spinosa (L) Campd. Oecologia 45: 244–251.
Zeide B (1978). Reproductive behavior of plants in time. Am Nat 112: 636–639.
Zhao, ZG, Du, GZ, and Huang, SQ (2010). The effect of flower position on variation and covariation in floral traits in a wild hermaphrodite plant. BMC Plant Biology 10: 91.