Costs of reproduction due to resource allocation trade-offs have long been recognized as key forces in life history evolution, but little is known about their functional or genetic basis. Arabidopsis lyrata, a perennial relative of the annual model plant A. thaliana with a wide climatic distribution, has populations that are strongly diverged in resource allocation. In this study, we evaluated the genetic and functional basis for variation in resource allocation in a reciprocal transplant experiment, using four A. lyrata populations and F 2 progeny from a cross between North Carolina (NC) and Norway parents, which had the most divergent resource allocation patterns. Local alleles at quantitative trait loci (QTL) at a North Carolina field site increased reproductive output while reducing vegetative growth. These QTL had little overlap with flowering date QTL. Structural equation models incorporating QTL genotypes and traits indicated that resource allocation differences result primarily from QTL effects on early vegetative growth patterns, with cascading effects on later vegetative and reproductive development. At a Norway field site, North Carolina alleles at some of the same QTL regions reduced survival and reproductive output components, but these effects were not associated with resource allocation trade-offs in the Norway environment. Our results indicate that resource allocation in perennial plants may involve important adaptive mechanisms largely independent of flowering time. Moreover, the contributions of resource allocation QTL to local adaptation appear to result from their effects on developmental timing and its interaction with environmental constraints, and not from simple models of reproductive costs.
DIFFERENTIAL allocation of resources to current reproduction vs. continued growth and maintenance is a central feature of life history differences among organisms (Stearns 1992;Roff and Fairbairn 2007). Reproduction is widely recognized as costly, leading to trade-offs between investment in reproduction vs. growth and maintenance and ultimately survival (Bell 1980;Reznick 1985;Lovett Doust 1989). Optimal resource allocation strategies are likely to vary by environment due to differential effects of allocation on growth rates, survival, and fecundity, thus leading to selection for different allocation strategies in different environments (Williams 1966;Charnov and Schaffer 1973;Bell 1980;Reznick 1985;Biere 1995;Johnson 2007).Among iteroparous organisms, which must allocate some resources to somatic maintenance in order to survive to reproduce multiple times, a wide range of proportional investments in growth and maintenance vs. reproduction is possible (van Noordwijk and De Jong 1986). Genetic variation in traits subject to trade-offs results in what has been termed "structured pleiotropy," in which genetic covariances between traits result from functional constraints imposed by the limiting resources (De Jong 1990;Stearns et al. 1991; see Figure 1B). However, differences in resource acquisi...