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1.  Root Foraging Increases Performance of the Clonal Plant Potentilla reptans in Heterogeneous Nutrient Environments 
PLoS ONE  2013;8(3):e58602.
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.
doi:10.1371/journal.pone.0058602
PMCID: PMC3589344  PMID: 23472211
2.  Optimal use of leaf nitrogen explains seasonal changes in leaf nitrogen content of an understorey evergreen shrub 
Annals of Botany  2011;108(3):529-536.
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.
doi:10.1093/aob/mcr167
PMCID: PMC3158686  PMID: 21757476
Acclimation; Aucuba japonica; daily carbon gain; nitrogen-use efficiency (NUE); optimization theory; understorey
3.  Relationships between tree size, crown shape, gender segregation and sex allocation in Pinus halepensis, a Mediterranean pine tree 
Annals of Botany  2011;108(1):197-206.
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.
doi:10.1093/aob/mcr104
PMCID: PMC3119615  PMID: 21586528
Allocation; biomass; branch vigour; crown shape; gender segregation; nitrogen; Pinus halepensis; phosphorus; sex allocation; tree size
4.  Effects of Trampling on Morphological and Mechanical Traits of Dryland Shrub Species Do Not Depend on Water Availability 
PLoS ONE  2013;8(1):e53021.
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.
doi:10.1371/journal.pone.0053021
PMCID: PMC3547011  PMID: 23341918

Results 1-4 (4)