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Effects of Plant Size on Reproductive Output and Offspring Performance In Journal of Blackwell Science, Ltd Ecology 2002 Effects of plant size on reproductive output and offspring 90, 958–966 performance in the facultative biennial Digitalis purpurea NINA SLETVOLD Department of Biology, Division of Botany and Plant Physiology, University of Oslo, PO Box 1045 Blindern, N-0316 Oslo, Norway Summary 1 Monocarpy in facultative biennials can be favoured by selection when there is a more than proportional increase in fitness with size. The possible contributions of larger reproductive output and higher offspring quality to such an increase were investigated in the facultative biennial plant Digitalis purpurea. Straw mass (rosette leaves and flower- ing stem excluding fruits and seeds) was determined for 100 individuals of varying size and correlated with the number of flowers and seeds per flower to examine whether sex- allocation is size dependent. 2 To determine whether maternal size influences offspring growth, and whether this effect is more pronounced in the presence of competition, seeds from 32 maternal plants were grown alone, and with one, two or four competitors. 3 Total seed number increased proportionally with size. Seed quality, measured as seed size, germination percentage and speed also increased with maternal size. The combined fitness measures of total seed mass (seed number × seed size), number of seedlings (seed number × germination percentage) and total female fitness (seed number × germination percentage × mean offspring dry mass after 14 weeks) all increased disproportionally with size. 4 Sex-allocation was size dependent, with large plants emphasizing the female function. 5 Offspring dry mass after both 8 and 14 weeks of growth was strongly influenced by competition, whereas the effect of maternal size was significant only at the first harvest date. Competition did not accentuate the effects of maternal size. 6 The results support that monocarpy exists in D. purpurea because of increased offspring quality rather than a disproportionate advantage in seed output of large individuals. Key-words: delayed reproduction, maternal size-effects, monocarpy, offspring quality, seed size variation Journal of Ecology (2002) 90, 958–966 monocarpic species that can complete their life cycle Introduction within 2 years, but where a delay of reproduction until Individual plant size is usually one of the best predic- the third or later year is commonly observed (Werner tors of current and future reproductive output, and the 1975; van der Meijden & van der Waals-Kooi 1979; relative advantages or disadvantages of reproducing as Gross 1981; de Jong et al. 1986; Klinkhamer et al. a large individual have been broadly discussed (Stearns 1987a, b, 1991; de Jong & Klinkhamer 1988; Campbell 1992). The question of whether there exists an optimal 1997). A monocarpic perennial life-history could be size for reproduction has been of particular interest in of selective advantage when there is a more than pro- the study of facultative biennials (Kachi & Hirose portional fitness gain with size (Schaffer 1974; Schaffer 1985; de Jong et al. 1989; Kachi 1990). These are & Gadgil 1975). This gain could be caused by a more than proportional increase in reproductive output with plant size, or by a higher quality of the offspring pro- Correspondence: Nina Sletvold, Department of Biology, Division of Botany and Plant Physiology, University of Oslo, duced by larger individuals (Stearns 1992). © 2002 British PO Box 1045 Blindern, N-0316 Oslo, Norway (fax 47 22 85 46 Initiation of reproduction in facultative biennial spe- Ecological Society 64; e-mail [email protected]). cies depends on size rather than on age (Lacey 1986), 959 and the individual plant has to reach a certain thresh- reproductive output is proportional to size in most Plant size and old size before any reproduction can be initiated Cynoglossum officinale populations in most years reproduction (Wesselingh et al. 1993). Plants with identical thresh- (Klinkhamer & de Jong 1987; Klinkhamer et al. 1990). old sizes could differ in timing of reproduction, due to A more than proportional increase was found in differences in local habitat quality. Whenever growth is Verbascum thapsus (original data in Reinartz 1984; restricted, due to competition and/or low habitat qual- reanalyses based on population means by Klinkhamer ity, it will take longer to reach the threshold size. The et al. 1990), but a decrease in Carlina vulgaris proximate cause of population variation in timing of (Klinkhamer et al. 1992) shows that, in biennials too, reproduction is, however, often plant-to-plant vari- convincing support is not available. ation in the size requirement for vernalization and/or The evidence of a positive correlation between photoinduction (Wesselingh & de Jong 1995; maternal size and offspring quality is predominantly Wesselingh & Klinkhamer 1996; van Dijk et al. 1997), zoological (Stearns 1992). Many studies on plants and evolution of an optimal size for reproduction will have, however, demonstrated considerable maternal occur through selection on these traits. Delayed repro- influence on seed size and quality. This maternal influ- duction can be caused by selection for an increased ence on seed provisioning/offspring phenotype is medi- threshold size for reproduction, and this will be advant- ated through other mechanisms than nuclear gene ageous when the demographic fitness loss due to transmission, and can be separated into genetic and longer pre-reproductive growth is more than compen- environmental effects. Genetic maternal effects can be sated by the fitness gain achieved by reproducing as a caused by strictly maternal inheritance of mitochon- large individual (Harper 1967; Schaffer & Gadgil 1975; dria, chloroplasts and/or plastids (Roach & Wulff Hart 1977; Silvertown 1983). 1987; Platenkamp & Shaw 1993). Environmental A more than proportional increase in reproductive maternal effects may be caused by several characteri- output with size could arise through a disproportion- stics such as temperature, light, soil nutrient level and ate access to resources by large plants. Alternatively, water availability (Gutterman 1992), as well as levels of large plants may attract more pollinators, and this pollination (Quesada et al. 1993, 1996) or herbivory could result in higher levels of both pollen deposition (Crawley & Nachapong 1985; Sills & Nienhuis 1995). and removal, increasing both female and male repro- Environmental maternal effects on offspring fitness ductive output. Large individuals of monocarpic spe- often seem to be transient (Miao et al. 1991; Wulff & cies tend to emphasize the female function (de Jong & Bazzaz 1992; Schmid & Dolt 1994; Andalo et al. 1999), Klinkhamer 1989), indicating that increased size is but may still have great impact if they influence life more important for seed production than for pollen cycle stages that are crucial for fitness, e.g, probability removal. The higher quality of offspring produced by and speed of germination (Alexander & Wulff 1985; large plants could arise through the same processes as Platenkamp & Shaw 1993). Competition has also been the higher reproductive output. A more effective shown to increase the influence of environmental resource acquisition by a large mother plant could maternal effects (Stratton 1989). The evidence of a pos- result in better provided seeds, particularly so for itive correlation between seed size and germination and monocarpic species that invest all available resources in subsequent seedling growth is substantial (Gross 1984; a single reproductive event. Alternatively, if large Stanton 1984; Wulff 1986a,b; Aarssen & Burton plants receive more pollinator visits per flower, this 1990; Schmid & Dolt 1994; Weiner et al. 1997), but the could lead to higher outcrossing rates and improved relationship between maternal size and seed size and seed quality. The relationship between outcrossing rate quality has received little attention. Klinkhamer & de and plant size is, however, problematic, as larger plants Jong (1987) found no significant relation between may also receive more geitonogamous pollinations maternal size and seed mass or germination percent- resulting in higher selfing rates (Iwasa et al. 1995) and age in the facultative biennial species Cynoglossum fitness loss due to inbreeding depression (Darwin officinale. 1876). The reasons for the evolution of the facultative bien- In contrast to most zoological studies, reproductive nial life-history strategy (Klinkhamer et al. 1997) are output explanations have generally been emphasized in still largely unresolved, and there are few data on the studies of plants. Optimal allocation models have relationship between size, reproductive output and shown that, in the absence of a more than proportional offspring quality in biennial species. In this study, I increase in reproductive output with plant size, the examine the relationship between plant size and optimal life-history is either annual or polycarpic per- reproductive output in the facultative biennial ennial, but never monocarpic perennial (Pugliese 1988; species Digitalis purpurea L., to address the following Iwasa & Cohen 1989). The empirical evidence of any questions: such disproportionate increase in reproductive output • Is there a more than proportional increase in repro- with size is, however, mixed (Samson & Werk 1986). ductive output with size in this species? © 2002 British Ecological Society, Decreasing, constant and increasing relationships have • Is offspring quality (measured as seed size,
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