BackgroundAs proposed by Darwin, climbers have been assumed to allocate a smaller fraction of biomass to support organs in comparison with self-supporting plants. They have also been hypothesized to possess a set of traits associated with fast growth, resource uptake and high productivity.ScopeIn this review, these hypotheses are evaluated by assembling and synthesizing published and unpublished data sets from across the globe concerning resource allocation, growth rates and traits of leaves, stems and roots of climbers and self-supporting species.ConclusionsThe majority of studies offer little support for the smaller allocation of biomass to stems or greater relative growth rates in climbers; however, these results are based on small sized (<1 kg) plants. Simulations based on allometric biomass equations demonstrate, however, that larger lianas allocate a greater fraction of above-ground biomass to leaves (and therefore less biomass to stems) compared with similar sized trees. A survey of leaf traits of lianas revealed their lower average leaf mass per area (LMA), higher N and P concentration and a slightly higher mass-based photosynthetic rate, as well as a lower concentration of phenolic-based compounds than in woody self-supporting species, consistent with the specialization of lianas towards the fast metabolism/rapid turnover end of the global trait spectra. Liana stems have an efficient hydraulic design and unique mechanical features, while roots appear to penetrate deeper soil levels than in trees and are often able to generate hydraulic pressure. Much remains to be learned, however, about these and other functional specializations of their axial organs and the associated trade-offs. Developmental switches between self-supporting, searcher and climbing shoots within the same individual are a promising field of comparative studies on trait association in lianas. Finally, some of the vast trait variability within lianas may be reduced when species with different climbing mechanisms are considered separately, and when phylogenetic conservatism is accounted for.

I tested hypotheses for ecological roles of storage carbohydrates in perennating organs (roots and branches) of alpine Oxytropis sericea, a leguminous herb. In naturally growing plants, total nonstructural carbohydrates achieved their maximal concentration in the fall, declined during winter, and reached minimal levels immediately after growth initiation in the spring. Experimental manipulation of carbon sink-source relations through shading of leaves of reproductive plants revealed that the normally unused portion of these carbohydrates is largely available for withdrawal. In another experiment, plants subjected to carbohydrate depletion through shading suffered decreased leaf growth after winter dormancy and had a lower probability of flowering and decreased inflorescence biomass. The dependence of reproductive growth on stored carbohydrates, however, was limited to its initial stages, because accumulation of storage carbohydrates occurred simultaneously with inflorescence expansion, flowering, and fruiting. Moreover, the whole-plant photosynthetic rate, estimated from gas exchange measurements also peaked at the time of inflorescence growth. To address whether stored reserves allow compensatory regrowth following defoliation, plants were subjected to experimental removal of leaves and inflorescences. Defoliated O. sericea partly regrew the lost leaves but withdrawal of stored carbohydrates was limited. Similarly, in a second defoliation experiment where infructescences were left intact, the plants used little stored carbohydrate and only partly compensated for fruit growth. However, carbohydrate accumulation was negatively affected by defoliation. While the ecological importance of stored nonstructural carbohydrates cannot be attributed to any function in isolation, winter respiration, leaf regrowth after winter, and early reproductive growth in O. sericea all depend to a significant extent on stored reserves. Maintaining a large storage pool may protect these functions in years when carbon status is less favorable than during this study.

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Spectra of leaf traits in northern temperate forest canopies reflect major differences in leaf longevity between evergreen conifers and deciduous broadleaf angiosperms, as well as plastic modifications caused by within-crown shading. We investigated (1) whether long-lived conifer leaves exhibit similar intra-canopy plasticity as short-lived broadleaves, and (2) whether global interspecific relationships between photosynthesis, nitrogen, and leaf structure identified for sun leaves adequately describe leaves differentiated in response to light gradients. We studied structural and photosynthetic properties of intra-tree sun and shade foliage in adult trees of seven conifer and four broadleaf angiosperm species in a common garden in Poland. Shade leaves exhibited lower leaf mass-per-area (LMA) than sun leaves; however, the relative difference was smaller in conifers than in broadleaves. In broadleaves, LMA was correlated with lamina thickness and tissue density, while in conifers, it was correlated with thickness but not density. In broadleaves, but not in conifers, reduction of lamina thickness was correlated with a thinner palisade layer. The more conservative adjustment of conifer leaves could result from a combination of phylogenetic constraints, contrasting leaf anatomies and shoot geometries, but also from functional requirements of long-lived foliage. Mass-based nitrogen concentration (Nmass) was similar between sun and shade leaves, and was lower in conifers than in deciduous broadleaved species. Given this, the smaller LMA in shade corresponded with a lower area-based N concentration (Narea). In evergreen conifers, LMA and Narea were less powerful predictors of area-based photosynthetic rate (Amax(area)) in comparison with deciduous broadleaved angiosperms. Multiple regression for sun and shade leaves showed that, in each group, Amax(mass) was related to Nmass but not to LMA, whereas LMA became a significant codeterminant of Amax(mass) in analysis combining both groups. Thus, a fundamental mass-based relationship between photosynthesis, nitrogen, and leaf structure reported previously also exists in a dataset combining within-crown and across-functional type variation.Electronic supplementary materialThe online version of this article (doi:10.1007/s00442-012-2279-y) contains supplementary material, which is available to authorized users.

Aim: Scots pine is one of the dominant tree species in forest ecosystems of the temperate and boreal zones in Eurasia. Since the Tertiary, it has persisted also in the Mediterranean region, forming relict populations. In this study, we investigate rangewide genetic diversity of Scots pine, aiming to provide comprehensive information on the phylogeography and genetic resources of the species.Location: Europe and Asia. Methods: Thirteen paternally inherited chloroplast microsatellite (cpSSR) loci were used to investigate the genetic structure of 62 populations from Fennoscandia and 30 populations from the Mediterranean. We also surveyed variability of two maternally inherited mitochondrial DNA (mtDNA) regions and additionally included literature data from 156 populations (248 populations in total).Results: Scots pine retains an overall high genetic diversity in the chloroplast genome but, unexpectedly, the historically younger populations from Fennoscandia showed statistically higher mean intrapopulation diversity than the Mediterranean stands (0.807 and 0.750, respectively). The latter also contained a lower number of private cpSSR haplotypes.Population differentiation with respect to mtDNA was higher (G ST = 0.628) than at cpSSR (G ST = 0.015) indicating an efficient pollen-mediated gene flow among refugial populations of the species. One novel haplotype mtDNA was found, increasing the number of known mitotypes for the species from 4 to 5. The geographic distribution of mitotypes was structured into four groups corresponding to the main refugial areas of the species. Main conclusions: Genetic impoverishment of the Mediterranean populations may enhance their vulnerability to future environmental changes. The spatial distribution of two mitotypes in Fennoscandia, featuring predominance of mitotype a in Norway and Sweden and mitotype b in Finland, gives strong support for dual colonization of that region from south-eastern (Finland) and south-western (Sweden) directions. These results thus provide new insights into both Holocene expansion of Scots pine and the present distribution of species' genetic resources. | 541 DERING Et al.