Optimal use of leaf nitrogen explains seasonal changes in leaf nitrogen content of an understorey evergreen shrub.

BACKGROUND AND AIMS: Understorey evergreen species commonly have a higher leaf nitrogen content in winter than in summer. Tested here is a hypothesis that such changes in leaf nitrogen content maximize nitrogen-use efficiency, defined as the daily carbon gain per unit nitrogen, under given temperature and irradiance levels. METHODS: The evergreen shrub Aucuba japonica growing naturally at three sites with different irradiance regimes in Japan was studied. Leaf photosynthetic characteristics, Rubisco and leaf nitrogen with measurements of temperature and irradiance monthly at each site were determined. Daily carbon gain was determined as a function of leaf nitrogen content to calculate the optimal leaf nitrogen content that maximized daily nitrogen-use efficiency. KEY RESULTS: As is known, the optimal leaf nitrogen content increased with increasing irradiance. The optimal leaf nitrogen content also increased with decreasing temperature because the photosynthetic capacity per Rubisco decreased. Across sites and months, the optimal leaf nitrogen content was close to the actual leaf nitrogen content and explained the variation in actual leaf nitrogen by 64 %. Sensitivity analysis showed that the effect of temperature on optimal nitrogen content was similar in magnitude to that of irradiance. CONCLUSIONS: Understorey evergreen species regulate leaf nitrogen content so as to maximize nitrogen-use efficiency in daily carbon gain under changing irradiance and temperature conditions.

Relationships between tree size, crown shape, gender segregation and sex allocation in Pinus halepensis, a Mediterranean pine tree.

BACKGROUND AND AIMS: Sex allocation has been studied mainly in small herbaceous plants but much less in monoecious wind-pollinated trees. The aim of this study was to explore changes in gender segregation and sex allocation by Pinus halepensis, a Mediterranean lowland pine tree, within tree crowns and between trees differing in their size or crown shape. METHODS: The production of new male and female cones and sex allocation of biomass, nitrogen and phosphorus were studied. The relationship between branch location, its reproductive status and proxies of branch vigour was also studied. KEY RESULTS: Small trees produced only female cones, but, as trees grew, they produced both male and female cones. Female cones were produced mainly in the upper part of the crown, and male cones in its middle and lower parts. Lateral branch density was correlated with the number of male but not female cones; lateral branches were more dense in large than in small trees and even denser in hemispherical trees. Apical branches grew faster, were thicker and their phosphorus concentration was higher than in lateral shoots. Nitrogen concentration was higher in cone-bearing apical branches than in apical vegetative branches and in lateral branches with or without cones. Allocation to male relative to female function increased with tree size as predicted by sex allocation theory. CONCLUSIONS: The adaptive values of sex allocation and gender segregation patterns in P. halepensis, in relation to its unique life history, are demonstrated and discussed. Small trees produce only female cones that have a higher probability of being pollinated than the probability of male cones pollinating; the female-first strategy enhances population spread. Hemispherical old trees are loaded with serotinous cones that supply enough seeds for post-fire germination; thus, allocation to males is more beneficial than to females.

The leaf anatomy of a broad-leaved evergreen allows an increase in leaf nitrogen content in winter.

In temperate regions, evergreen species are exposed to large seasonal changes in air temperature and irradiance. They change photosynthetic characteristics of leaves responding to such environmental changes. Recent studies have suggested that photosynthetic acclimation is strongly constrained by leaf anatomy such as leaf thickness, mesophyll and chloroplast surface facing the intercellular space, and the chloroplast volume. We studied how these parameters of leaf anatomy are related with photosynthetic seasonal acclimation. We evaluated differential effects of winter and summer irradiance on leaf anatomy and photosynthesis. Using a broad-leaved evergreen Aucuba japonica, we performed a transfer experiment in which irradiance regimes were changed at the beginning of autumn and of spring. We found that a vacant space on mesophyll surface in summer enabled chloroplast volume to increase in winter. The leaf nitrogen and Rubisco content were higher in winter than in summer. They were correlated significantly with chloroplast volume and with chloroplast surface area facing the intercellular space. Thus, summer leaves were thicker than needed to accommodate mesophyll surface chloroplasts at this time of year but this allowed for increases in mesophyll surface chloroplasts in the winter. It appears that summer leaf anatomical characteristics help facilitate photosynthetic acclimation to winter conditions. Photosynthetic capacity and photosynthetic nitrogen use efficiency were lower in winter than in summer but it appears that these reductions were partially compensated by higher Rubisco contents and mesophyll surface chloroplast area in winter foliage.

Author Correction: Identifying long-term stable refugia for relict plant species in East Asia.

In the original version of this Article, an incorrect sample size was provided for the number of relict species (443 instead of 442) and the number of relict forests (423 instead of 422). These errors have been corrected in both the PDF and HTML versions of the Article.

Identifying long-term stable refugia for relict plant species in East Asia.

Today East Asia harbors many "relict" plant species whose ranges were much larger during the Paleogene-Neogene and earlier. The ecological and climatic conditions suitable for these relict species have not been identified. Here, we map the abundance and distribution patterns of relict species, showing high abundance in the humid subtropical/warm-temperate forest regions. We further use Ecological Niche Modeling to show that these patterns align with maps of climate refugia, and we predict species' chances of persistence given the future climatic changes expected for East Asia. By 2070, potentially suitable areas with high richness of relict species will decrease, although the areas as a whole will probably expand. We identify areas in southwestern China and northern Vietnam as long-term climatically stable refugia likely to preserve ancient lineages, highlighting areas that could be prioritized for conservation of such species.

Testing sustainability by prospective and retrospective demographic analyses: evaluation for palm leaf harvest.

Harvesting nontimber forest products (NTFPs) is a major economic activity in tropical forests. As many NTFPs are overexploited, sustainability analyses are required to set harvest guidelines. Here we introduce and apply a new approach to evaluating sustainability, which combines prospective (elasticity) and retrospective (Life Table Response Experiments [LTRE]) demographic analyses of matrix population models. We relate the elasticity of vital rates (representing their importance for population growth rate, lamda) to their contribution to harvest-induced change in lamda ("LTRE contribution"). When high-elasticity vital rates have a low LTRE contribution, exploitation is potentially sustainable as negative effects for population growth are buffered. If the reverse is found, there is little scope for sustainability because crucial vital rates are affected. Our approach is less sensitive to chance fluctuations than the commonly used sustainability criterion of lamda = 1.0, as it does not depend on the absolute value of lamda. We applied this analysis to Geonoma deversa, a clustered forest understory palm. We studied three experimentally defoliated and control populations in a Bolivian rainforest during two years. Cutting all leaves of large ramets did not change mortality but strongly affected growth and reproduction. In spite of severe changes in some vital rates, population growth rate was not significantly reduced after defoliation. A literature review revealed that six other understory palms species responded very similarly to defoliation. The combination of LTRE contributions and elasticity analyses showed that low-elasticity vital rates were mainly responsible for the defoliation-induced change in lamda for Geonoma deversa. For two other understory palms (Astrocaryum mexicanum and Chamaedorea radicalis) new demographic analyses yielded very similar results. For Geonoma, the LTRE contribution-elasticity relation strongly changed when we mimicked harvest damage. Adding 5% mortality to defoliated palms caused stronger change in lamda, mainly due to changes in a high-elasticity vital rate (survival). Therefore, harvest practices that involve stem killing are clearly unsustainable. Our results show that commercial leaf cutting in Geonoma deversa is potentially sustainable, and that this is likely the case for understory palms in general. Our approach to evaluating harvest sustainability can be applied to other NTFPs.

Mobile dune fixation by a fast-growing clonal plant: a full life-cycle analysis.

Desertification is a global environmental problem, and arid dunes with sparse vegetation are especially vulnerable to desertification. One way to combat desertification is to increase vegetation cover by planting plant species that can realize fast population expansion, even in harsh environments. To evaluate the success of planted species and provide guidance for selecting proper species to stabilize active dunes, demographic studies in natural habitats are essential. We studied the life history traits and population dynamics of a dominant clonal shrub Hedysarum laeve in Inner-Mongolia, northern China. Vital rates of 19057 ramets were recorded during three annual censuses (2007-2009) and used to parameterize Integral Projection Models to analyse population dynamics. The life history of H. laeve was characterized by high ramet turnover and population recruitment entirely depended on clonal propagation. Stochastic population growth rate was 1.32, suggesting that the populations were experiencing rapid expansion. Elasticity analysis revealed that clonal propagation was the key contributor to population growth. The capacity of high clonal propagation and rapid population expansion in mobile dunes makes H. laeve a suitable species to combat desertification. Species with similar life-history traits to H. laeve are likely to offer good opportunities for stabilizing active dunes in arid inland ecosystems.

Root foraging increases performance of the clonal plant Potentilla reptans in heterogeneous nutrient environments.

BACKGROUND: Plastic root-foraging responses have been widely recognized as an important strategy for plants to explore heterogeneously distributed resources. However, the benefits and costs of root foraging have received little attention. METHODOLOGY/PRINCIPAL FINDINGS: In a greenhouse experiment, we grew pairs of connected ramets of 22 genotypes of the stoloniferous plant Potentilla reptans in paired pots, between which the contrast in nutrient availability was set as null, medium and high, but with the total nutrient amount kept the same. We calculated root-foraging intensity of each individual ramet pair as the difference in root mass between paired ramets divided by the total root mass. For each genotype, we then calculated root-foraging ability as the slope of the regression of root-foraging intensity against patch contrast. For all genotypes, root-foraging intensity increased with patch contrast and the total biomass and number of offspring ramets were lowest at high patch contrast. Among genotypes, root-foraging intensity was positively related to production of offspring ramets and biomass in the high patch-contrast treatment, which indicates an evolutionary benefit of root foraging in heterogeneous environments. However, we found no significant evidence that the ability of plastic foraging imposes costs under homogeneous conditions (i.e. when foraging is not needed). CONCLUSIONS/SIGNIFICANCE: Our results show that plants of P. reptans adjust their root-foraging intensity according to patch contrast. Moreover, the results show that the root foraging has an evolutionary advantage in heterogeneous environments, while costs of having the ability of plastic root foraging were absent or very small.

Effects of trampling on morphological and mechanical traits of dryland shrub species do not depend on water availability.

In semiarid drylands water shortage and trampling by large herbivores are two factors limiting plant growth and distribution. Trampling can strongly affect plant performance, but little is known about responses of morphological and mechanical traits of woody plants to trampling and their possible interaction with water availability. Seedlings of four shrubs (Caragana intermedia, Cynanchum komarovi, Hedysarum laeve and Hippophae rhamnoides) common in the semiarid Mu Us Sandland were grown at 4% and 10% soil water content and exposed to either simulated trampling or not. Growth, morphological and mechanical traits were measured. Trampling decreased vertical height and increased basal diameter and stem resistance to bending and rupture (as indicated by the increased minimum bend and break force) in all species. Increasing water availability increased biomass, stem length, basal diameter, leaf thickness and rigidity of stems in all species except C. komarovii. However, there were no interactive effects of trampling and water content on any of these traits among species except for minimum bend force and the ratio between stem resistance to rupture and bending. Overall shrub species have a high degree of trampling resistance by morphological and mechanical modifications, and the effects of trampling do not depend on water availability. However, the increasing water availability can also affect trade-off between stem strength and flexibility caused by trampling, which differs among species. Water plays an important role not only in growth but also in trampling adaptation in drylands.

Height convergence in response to neighbour growth: genotypic differences in the stoloniferous plant Potentilla reptans.

Using a new experimental set up, the way in which height growth of stoloniferous plants is adjusted to that of their neighbours, as well as differences between genotypes in their ability to keep up with neighbour height growth were tested. Five Potentilla reptans genotypes inherently differing in petiole length were subjected to three experimental light gradients, involving light intensity and red : far-red ratio. Each plant was placed in a vertically adjustable cylinder of green foil, and the treatments differed in the speed of cylinder height increase and final height. Total weight of plants decreased from the 'Slow' to the 'Fast' treatment, while petiole length increased. Leaves reaching the top of the cylinder stopped petiole elongation, resulting in similar final heights for all genotypes in the 'Slow' treatment. In the 'Fast' treatment only the fastest-growing genotype maintained its position in the top of the cylinder and genotypes differed strongly in final height within the cylinders. Plants adjust their height growth to that of the surrounding vegetation, leading to height convergence in short light gradients that slowly increase. These adjustments and genotypic differences in ability to keep up with fast-growing neighbours can influence the outcome of competition for light.

Symbiotic nitrogen fixation in a tropical rainforest: 15N natural abundance measurements supported by experimental isotopic enrichment.

* Leguminous trees are very common in the tropical rainforests of Guyana. Here, species-specific differences in N(2) fixation capability among nodulating legumes growing on different soils and a possible limitation of N(2) fixation by a relatively high nitrogen (N) and low phosphorus (P) availability in the forest were investigated. * Leaves of 17 nodulating species and 17 non-nodulating reference trees were sampled and their delta(15)N values measured. Estimates of N(2) fixation rates were calculated using the (15)N natural abundance method. Pot experiments were conducted on the effect of N and P availability on N(2) fixation using the (15)N-enriched isotope dilution method. * Nine species showed estimates of > 33% leaf N derived from N(2) fixation, while the others had low or undetectable N(2) fixation rates. High N and low P availability reduced N(2) fixation substantially. * The results suggest that a high N and low P availability in the forest limit N(2) fixation. At the forest ecosystem level, N(2) fixation was estimated at c. 6% of total N uptake by the tree community. We conclude that symbiotic N(2) fixation plays an important role in maintaining high amounts of soil available N in undisturbed forest.

Tuber size variation and organ preformation constrain growth responses of a spring geophyte.

Functional responses to environmental variation do not only depend on the genetic potential of a species to express different trait values, but can also be limited by characteristics, such as the timing of organ (pre-) formation, aboveground longevity or the presence of a storage organ. In this experiment we tested to what degree variation in tuber size and organ preformation constrain the responsiveness to environmental quality and whether responsiveness is modified by the availability of stored resources by exposing the spring geophyte Bunium bulbocastanum to different light and nutrient regimes. Growth and biomass partitioning were affected by initial tuber size and resource availability. On average, tuber weight amounted to 60%, but never less than 30% of the total plant biomass. Initial tuber size, considered an estimate of the total carbon pool available at the onset of treatments, affected plant growth and reproduction throughout the experiment but had little effect on the responsiveness of plants to the treatments. The responsiveness was partly constrained by organ preformation: in the second year variation of leaf number was considerably larger than in the first year of the treatments. The results indicate that a spring geophyte with organ preformation has only limited possibilities to respond to short-term fluctuations of the environment, as all leaves and the inflorescence are preformed in the previous growing season and resources stored in tubers are predominantly used for survival during dormancy and are not invested into plastic adjustments to environmental quality. Such spring geophytes have only limited possibilities to buffer environmental variation. This explains their restriction to habitats characterized by predictable changes of the environmental conditions.