Seed size, number and strategies in annual plants: a comparative functional analysis and synthesis.

BACKGROUND AND AIMS: Plants depend fundamentally on establishment from seed. However, protocols in trait-based ecology currently estimate seed size but not seed number. This can be rectified. For annuals, seed number should simply be a positive function of vegetative biomass and a negative function of seed size. METHODS: Using published values of comparative seed number as the 'gold standard' and a large functional database, comparative seed yield and number per plant and per m2 were predicted by multiple regression. Subsequently, ecological variation in each was explored for English and Spanish habitats, newly calculated C-S-R strategies and changed abundance in the British flora. KEY RESULTS: As predicted, comparative seed mass yield per plant was consistently a positive function of plant size and competitive ability, and largely independent of seed size. Regressions estimating comparative seed number included, additionally, seed size as a negative function. Relationships differed numerically between regions, habitats and C-S-R strategies. Moreover, some species differed in life history over their geographical range. Comparative seed yield per m2 was positively correlated with FAO crop yield, and increasing British annuals produced numerous seeds. Nevertheless, predicted values must be viewed as comparative rather than absolute: they varied according to the 'gold standard' predictor used. Moreover, regressions estimating comparative seed yield per m2 achieved low precision. CONCLUSIONS: For the first time, estimates of comparative seed yield and number for >800 annuals and their predictor equations have been produced and the ecological importance of these regenerative traits has been illustrated. 'Regenerative trait-based ecology' remains in its infancy, with work needed on determinate vs. indeterminate flowering ('bet-hedging'), C-S-R methodologies, phylogeny, comparative seed yield per m2 and changing life history. Nevertheless, this has been a positive start and readers are invited to use estimates for >800 annuals, in the Supplementary data, to help advance 'regenerative trait-based ecology' to the next level.

Trade-offs between seed and leaf size (seed-phytomer-leaf theory): functional glue linking regenerative with life history strategies and taxonomy with ecology?

Background and Aims: While the 'worldwide leaf economics spectrum' (Wright IJ, Reich PB, Westoby M, et al. 2004. The worldwide leaf economics spectrum. Nature : 821-827) defines mineral nutrient relationships in plants, no unifying functional consensus links size attributes. Here, the focus is upon leaf size, a much-studied plant trait that scales positively with habitat quality and components of plant size. The objective is to show that this wide range of relationships is explicable in terms of a seed-phytomer-leaf (SPL) theoretical model defining leaf size in terms of trade-offs involving the size, growth rate and number of the building blocks (phytomers) of which the young shoot is constructed. Methods: Functional data for 2400+ species and English and Spanish vegetation surveys were used to explore interrelationships between leaf area, leaf width, canopy height, seed mass and leaf dry matter content (LDMC). Key Results: Leaf area was a consistent function of canopy height, LDMC and seed mass. Additionally, size traits are partially uncoupled. First, broad laminas help confer competitive exclusion while morphologically large leaves can, through dissection, be functionally small. Secondly, leaf size scales positively with plant size but many of the largest-leaved species are of medium height with basally supported leaves. Thirdly, photosynthetic stems may represent a functionally viable alternative to 'small seeds + large leaves' in disturbed, fertile habitats and 'large seeds + small leaves' in infertile ones. Conclusions: Although key elements defining the juvenile growth phase remain unmeasured, our results broadly support SPL theory in that phytometer and leaf size are a product of the size of the initial shoot meristem (≅ seed mass) and the duration and quality of juvenile growth. These allometrically constrained traits combine to confer ecological specialization on individual species. Equally, they appear conservatively expressed within major taxa. Thus, 'evolutionary canalization' sensu Stebbins (Stebbins GL. 1974. Flowering plants: evolution above the species level . Cambridge, MA: Belknap Press) is perhaps associated with both seed and leaf development, and major taxa appear routinely specialized with respect to ecologically important size-related traits.

Does retained-seed priming drive the evolution of serotiny in drylands? An assessment using the cactus Mammillaria hernandezii.

PREMISE OF THE STUDY: Serotinous plants retain their seeds for a long time. In deserts, retained seeds undergo hydration-dehydration cycles and thus may become primed. Priming enhances germination and seedling vigor. We test the hypothesis that serotiny evolves because it provides a site protected from predators in which seeds can become primed. Rainfall-cued dispersal of primed seeds may enhance this effect. METHODS: We tested this hypothesis with Mammillaria hernandezii through protein-content analyses; field and laboratory germination experiments with primed, unprimed, and retained seeds; and fitness estimations from demographic models. KEY RESULTS: Hydration-dehydration cycles induced priming, enhancing germination. Artificial priming and retention in the parent plant for 1 yr induced similar changes in seed protein patterns, suggesting that priming occurs naturally while seeds are retained. Under field conditions, germination of seeds retained for 1 yr more than doubled that of seeds of the same cohort that were not primed or that remained buried for 1 yr. The first seeds to germinate died rapidly. Serotinous plants whose seeds underwent priming had higher fitness than those whose seeds were in the soil seed bank or that did not experience priming. CONCLUSIONS: Priming in soil seed banks may be costly because of high predation, so seed protection during priming is sufficient to promote the evolution of serotiny. Bet hedging contributes to this process. Rapid germination of primed seeds that respond to brief rainfall events is disadvantageous because such rainfall is insufficient for seedling survival. Serotinous species counteract this cost by cueing dispersal with heavy precipitation.