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Allocation Strategies and Seed Traits Are Hardly Affected by Nitrogen

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Allocation Strategies and Seed Traits Are Hardly Affected by Nitrogen ARTICLE IN PRESS Perspectives in Plant Ecology, Evolution and Systematics Perspectives in Plant Ecology, Evolution and Systematics 11 (2009) 267–283 www.elsevier.de/ppees RESEARCH ARTICLE Allocation strategies and seed traits are hardly affected by nitrogen supply in 18 species differing in successional status Claire Fortunela,Ã, Cyrille Viollea, Catherine Roumeta, Bruno Buatoisa, Marie-Laure Navasb, Eric Garniera aCNRS, Centre d’Ecologie Fonctionnelle et Evolutive (UMR 5175), 1919 Route de Mende, 34293 Montpellier Cedex 5, France bMontpellier SupAgro, Centre d’Ecologie Fonctionnelle et Evolutive (UMR 5175), 1919 Route de Mende, 34293 Montpellier Cedex 5, France Received 9 May 2008; received in revised form 7 April 2009; accepted 8 April 2009 Abstract Species performance depends on ecological strategies, revealed by suites of traits, conferring different relative ecological advantages in different environments. Although current knowledge on plant strategies along successional gradients is derived from studies conducted in situ, actually quantifying these strategies requires disentangling the effects of environmental factors from intrinsic differences between species. Here we tested whether allocation strategies and seed traits differ among successional stages and nitrogen levels. To this aim, we assessed biomass and nitrogen allocations and seed traits variations for 18 species, differing in life history and belonging to three stages of a Mediterranean old-field succession. These species were grown as monocultures in an experimental garden under limiting and non-limiting nitrogen supply. Early successional species allocated allometrically more nitrogen and proportionally more biomass to reproduction, and set more seeds than later successional species. Seed mass increased with successional status and was negatively related to seed number. Early successional species thus produced more but less-provisioned seeds, suggesting better colonization abilities. These patterns were not the sole consequence of the replacement of annuals by perennials along the successional gradient, since comparable trends were also observed within each life history. Allocation patterns were generally not altered by nitrogen supply and the higher nitrogen content in vegetative organs of plants grown under high nitrogen supply was not retranslocated from leaves to seeds during seed development. We therefore conclude that differences in plant ecological strategies in species characteristics from contrasting successional stages appear to be intrinsic properties of the studied species, and independent from environmental conditions. r 2009 Ru¨bel Foundation, ETH Zu¨rich. Published by Elsevier GmbH. All rights reserved. Keywords: Allometry; Reproductive output; Seed mass; Nitrogen concentration of organs; Succession; Nitrogen supply Introduction Biomass and nutrient allocation patterns reflect the ÃCorresponding author. Present address: INRA Kourou-UMR EcoFoG (Ecologie des Foreˆts de Guyane), Avenue de France, BP 709, way species interact with their environment (Antono- 97387 Kourou Cedex, French Guiana, France. vics, 1980; Bazzaz et al., 1987). In successional environ- E-mail address: [email protected] (C. Fortunel). ments, the proportion of total plant biomass in leaf, 1433-8319/$ - see front matter r 2009 Ru¨bel Foundation, ETH Zu¨rich. Published by Elsevier GmbH. All rights reserved. doi:10.1016/j.ppees.2009.04.003 ARTICLE IN PRESS 268 C. Fortunel et al. / Perspectives in Plant Ecology, Evolution and Systematics 11 (2009) 267–283 stem and reproductive organs tend to decrease while allocation patterns observed in standard conditions that in root tends to increase as succession proceeds (e.g. were consistent with those observed in situ. Newell and Tramer, 1978; Abrahamson, 1979; Hancock In situations where nutrients strongly limit plant and Pritts, 1987). These patterns point to a decreasing growth, nutrient allocation patterns better reflect con- investment in growth and colonization in advanced straints on plant resource economy than do biomass successional stages, but an increasing investment in allocation patterns (Reekie and Bazzaz, 1987b). Yet, structures contributing to below-ground competitive and though nitrogen often limits plant growth in ability (Gleeson and Tilman, 1990). Early successional successional environments (Gleeson and Tilman, 1990; species also produce many small well-dispersed seeds, cf. Garnier et al., 2007 for the Mediterranean sere whereas late successional species invest in few large concerned here), only few studies have examined how offspring with higher probability of survival and nitrogen availability might affect the biomass and establishment (Stewart and Thompson, 1982; Gleeson nitrogen allocation patterns of species differing in and Tilman, 1990; Schippers et al., 2001; Fenner and successional status (e.g. Olff, 1992). Nitrogen availabil- Thompson, 2005). These changes in species character- ity influences allocation patterns between shoots and istics are usually concomitant with the replacement of roots, and affects nitrogen concentration of plant organs annual species by perennial species over succession (e.g. Reynolds and D’Antonio, 1996; Garnier, 1998). (Bazzaz, 1996; Prach et al., 1997; Vile et al., 2006 for the But the way in which internal nitrogen is distributed successional sere examined in the present study). among vegetative and reproductive organs is poorly Resource allocation patterns are traditionally studied understood. The studied species were therefore grown at as ratios between organ and total plant biomasses two nitrogen levels (limiting vs. non-limiting), to analyse (Reekie and Bazzaz, 1987a). However, biomass (and the impacts of nitrogen availability on the nitrogen nutrient) ratios usually vary with plant size; therefore concentrations of the different plant parts and on the the partitioning approach potentially confounds alloca- patterns of biomass and nitrogen allocation. tion patterns with plant size (Samson and Werk, 1986; The questions addressed in this study are as follows: Klinkhamer et al., 1990; Jasienski and Bazzaz, 1999; (1) Do species strategies vary according to successional Weiner, 2004). To examine life history tradeoffs as status when grown under common experimental condi- reflected by allocation patterns, several authors hence tions? Expectations are that species from the early argued that allometric functions are more appropriate successional stage would allocate, allometrically as well tools than biomass (or nutrient) ratios (Samson and as proportionally, more biomass to reproduction, and Werk, 1986; Klinkhamer et al., 1992; Sugiyama and produce smaller, less provisioned, but more numerous Bazzaz, 1998). The first aim of this study was to examine seeds than species from the late successional stage. (2) how allocation patterns differed among 18 plant species Does nitrogen availability alter species allocation characteristics of different stages of Mediterranean old- patterns and seed traits? Expectations are that high field successions, using this allometric approach. nitrogen supply would increase the biomasses and Most current knowledge on resource allocation nitrogen concentrations of vegetative and reproductive patterns in species characteristic of different succes- organs, subsequently altering resource allocation pat- sional stages is derived from studies conducted in situ terns between vegetative and reproductive organs, and (e.g. Newell and Tramer, 1978; Gleeson and Tilman, ultimately modifying the investment in reproduction. 1990; but see Jongejans et al., 2006 for a common garden experiment mimicking a successional gradient). In such studies, it is not possible to disentangle the effects of species identity from those of environmental Material and methods factors. In the Mediterranean successional seres of concern here, substantial differences in environmental Study site conditions were found as succession proceeds: light penetration at ground level decreased two-fold (Kaza- The experiment was conducted from October 2003 to kou and Navas, 2004) while total soil nitrogen September 2005 in the experimental garden of the concentration increased three-fold (Garnier et al., Centre d’Ecologie Fonctionnelle et Evolutive (CEFE, 2004) between recently abandoned fields and fields CNRS) in Montpellier, France (431590 N, 31510 E, 60 m abandoned 40 years earlier. Since light and nitrogen above sea level). The climate is Mediterranean sub- may exert strong controls on plant allocation patterns humid (Daget, 1977) with cool to cold winters, marked (see Poorter and Nagel, 2000 for a review), the detection summer drought and unpredictability of precipitation in of intrinsic differences among species from different time and amount. At the beginning of the experiment in successional stages requires that all species be grown October 2003, the soil pH was 7.82, which was close to under identical conditions. We therefore grew the 18 the pH values of the old-field succession from which the species in an experimental garden to test whether species originated (cf. Garnier et al., 2004). ARTICLE IN PRESS C. Fortunel et al. / Perspectives in Plant Ecology, Evolution and Systematics 11 (2009) 267–283 269 Table 1. List of the 18 species characteristic of three stages of a Mediterranean old-field succession that were grown in an experimental garden (Flora: Euro+Med Plantbase, www.emplantbase.org). Genus Species Family Taxonomic group Life history Successional stage Arenaria serpyllifolia Caryophyllaceae
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