Comparative transcriptome profiling of potato cultivars infected by late blight pathogen Phytophthora infestans: Diversity of quantitative and qualitative responses.

The Estonia potato cultivar Ando has shown elevated field resistance to Phytophthora infestans, even after being widely grown for over 40 years. A comprehensive transcriptional analysis was performed using RNA-seq from plant leaf tissues to gain insight into the mechanisms activated for the defense after infection. Pathogen infection in Ando resulted in about 5927 differentially expressed genes (DEGs) compared to 1161 DEGs in the susceptible cultivar Arielle. The expression levels of genes related to plant disease resistance such as serine/threonine kinase activity, signal transduction, plant-pathogen interaction, endocytosis, autophagy, mitogen-activated protein kinase (MAPK), and others were significantly enriched in the upregulated DEGs in Ando, whereas in the susceptible cultivar, only the pathway related to phenylpropanoid biosynthesis was enriched in the upregulated DEGs. However, in response to infection, photosynthesis was deregulated in Ando. Multi-signaling pathways of the salicylic-jasmonic-ethylene biosynthesis pathway were also activated in response to Phytophthora infestans infection.

Variation in Leaf Volatile Emissions in Potato (Solanum tuberosum) Cultivars with Different Late Blight Resistance.

Volatile organic compounds (VOCs) play key roles in plant abiotic and biotic stress resistance, but even for widespread crops, there is limited information on variations in the magnitude and composition of constitutive VOC emissions among cultivars with varying stress resistance. The foliage VOC emissions from nine local and commercial potato cultivars (Alouette, Sarme, Kuras, Ando, Anti, Jõgeva Kollane, Teele, 1681-11, and Reet) with medium to late maturities and varying Phytophthora infestans (the causative agent of late blight disease) resistance backgrounds were analyzed to gain an insight into the genetic diversity of constitutive VOC emissions and to test the hypothesis that cultivars more resistant to Phytophthora infestans have greater VOC emissions and different VOC fingerprints. Forty-six VOCs were identified in the emission blends of potato leaves. The majority of the VOCs were sesquiterpenes (50% of the total number of compounds and 0.5-36.9% of the total emissions) and monoterpenes (30.4% of the total number of compounds and 57.8-92.5% of the total VOC emissions). Qualitative differences in leaf volatiles, mainly in sesquiterpenes, were related to the potato genotype background. Among the volatile groups, the monoterpenes α-pinene, ß-pinene, Δ3-carene, limonene, and p-cymene, the sesquiterpenes (E)-ß-caryophyllene and α-copaene, and green leaf volatile hexanal were the major volatiles in all cultivars. A higher share of VOCs known to have antimicrobial activities was observed. Interestingly, the cultivars were grouped into high and low resistance categories based on the VOC profiles, and the total terpenoid and total constitutive VOC emission scale positively with resistance. To support and expedite advances in breeding for resistance to diseases such as late blight disease, the plant research community must develop a fast and precise approach to measure disease resistance. We conclude that the blend of emitted volatiles is a fast, non-invasive, and promising indicator to identify cultivars resistant to potato late blight disease.

Mesophyll conductance to CO2 and Rubisco as targets for improving intrinsic water use efficiency in C3 plants.

Water limitation is a major global constraint for plant productivity that is likely to be exacerbated by climate change. Hence, improving plant water use efficiency (WUE) has become a major goal for the near future. At the leaf level, WUE is the ratio between photosynthesis and transpiration. Maintaining high photosynthesis under water stress, while improving WUE requires either increasing mesophyll conductance (gm ) and/or improving the biochemical capacity for CO2 assimilation-in which Rubisco properties play a key role, especially in C3 plants at current atmospheric CO2 . The goals of the present analysis are: (1) to summarize the evidence that improving gm and/or Rubisco can result in increased WUE; (2) to review the degree of success of early attempts to genetically manipulate gm or Rubisco; (3) to analyse how gm , gsw and the Rubisco's maximum velocity (Vcmax ) co-vary across different plant species in well-watered and drought-stressed conditions; (4) to examine how these variations cause differences in WUE and what is the overall extent of variation in individual determinants of WUE; and finally, (5) to use simulation analysis to provide a theoretical framework for the possible control of WUE by gm and Rubisco catalytic constants vis-à-vis gsw under water limitations.

Temperature responses of the Rubisco maximum carboxylase activity across domains of life: phylogenetic signals, trade-offs, and importance for carbon gain.

Temperature response of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) catalytic properties directly determines the CO2 assimilation capacity of photosynthetic organisms as well as their survival in environments with different thermal conditions. Despite unquestionable importance of Rubisco, the comprehensive analysis summarizing temperature responses of Rubisco traits across lineages of carbon-fixing organisms is lacking. Here, we present a review of the temperature responses of Rubisco carboxylase specific activity (c(cat)(c)) within and across domains of life. In particular, we consider the variability of temperature responses, and their ecological, physiological, and evolutionary controls. We observed over two-fold differences in the energy of activation (ΔH(a)) among different groups of photosynthetic organisms, and found significant differences between C3 plants from cool habitats, C3 plants from warm habitats and C4 plants. According to phylogenetically independent contrast analysis, ΔH(a) was not related to the species optimum growth temperature (T growth), but was positively correlated with Rubisco specificity factor (S(c/o)) across all organisms. However, when only land plants were analyzed, ΔH(a) was positively correlated with both T(growth) and S(c/o), indicating different trends for these traits in plants versus unicellular aquatic organisms, such as algae and bacteria. The optimum temperature (T(opt)) for k(cat)(c) correlated with S(c/o) for land plants and for all organisms pooled, but the effect of T growth on T(opt) was driven by species phylogeny. The overall phylogenetic signal was significant for all analyzed parameters, stressing the importance of considering the evolutionary framework and accounting for shared ancestry when deciphering relationships between Rubisco kinetic parameters. We argue that these findings have important implications for improving global photosynthesis models.

Rubisco catalytic properties optimized for present and future climatic conditions.

Because of its catalytic inefficiencies, Rubisco is the most obvious target for improvement to enhance the photosynthetic capacity of plants. Two hypotheses are tested in the present work: (1) existing Rubiscos have optimal kinetic properties to maximize photosynthetic carbon assimilation in existing higher plants; (2) current knowledge allows proposal of changes to kinetic properties to make Rubiscos more suited to changed conditions in chloroplasts that are likely to occur with climate change. The catalytic mechanism of Rubisco results in higher catalytic rates of carboxylation being associated with decreased affinity for CO2, so that selection for different environments involves a trade-off between these two properties. The simulations performed in this study confirm that the optimality of Rubisco kinetics depends on the species and the environmental conditions. In particular, environmental drivers affecting the CO2 availability for carboxylation (Cc) or directly shifting the photosynthetic limitations between Rubisco and RuBP regeneration determine to what extend Rubisco kinetics are optimally suited to maximize CO2 assimilation rate. In general, modeled values for optimal kinetic reflect the predominant environmental conditions currently encountered by the species in the field. Under future climatic conditions, photosynthetic CO2 assimilation will be limited by RuBP-regeneration, especially in the absence of water stress, the largest rise in [CO2] and the lowest increases in temperature. Under these conditions, the model predicts that optimal Rubisco should have high Sc/o and low kcat(c).

Isoprene function in two contrasting poplars under salt and sunflecks.

In the present study, biogenic volatile organic compound (BVOC) emissions and photosynthetic gas exchange of salt-sensitive (Populus x canescens (Aiton) Sm.) and salt-tolerant (Populus euphratica Oliv.) isoprene-emitting and non-isoprene-emitting poplars were examined under controlled high-salinity and high-temperature and -light episode ('sunfleck') treatments. Combined treatment with salt and sunflecks led to an increased isoprene emission capacity in both poplar species, although the photosynthetic performance of P. × canescens was reduced. Indeed, different allocations of isoprene precursors between the cytosol and the chloroplast in the two species were uncovered by means of (13)CO2 labeling. Populus × canescens leaves, moreover, increased their use of 'alternative' carbon (C) sources in comparison with recently fixed C for isoprene biosynthesis under salinity. Our studies show, however, that isoprene itself does not have a function in poplar survival under salt stress: the non-isoprene-emitting leaves showed only a slightly decreased photosynthetic performance compared with wild type under salt treatment. Lipid composition analysis revealed differences in the double bond index between the isoprene-emitting and non-isoprene-emitting poplars. Four clear metabolomics patterns were recognized, reflecting systemic changes in flavonoids, sterols and C fixation metabolites due to the lack/presence of isoprene and the absence/presence of salt stress. The studies were complemented by long-term temperature stress experiments, which revealed the thermotolerance role of isoprene as the non-isoprene-emitting leaves collapsed under high temperature, releasing a burst of BVOCs. Engineered plants with a low isoprene emission potential might therefore not be capable of resisting high-temperature episodes.

Photosynthetic acclimation to light in woody and herbaceous species: a comparison of leaf structure, pigment content and chlorophyll fluorescence characteristics measured in the field.

Acclimation of foliage photosynthetic properties occurs with varying time kinetics, but structural, chemical and physiological factors controlling the kinetics of acclimation are poorly understood, especially in field environments. We measured chlorophyll fluorescence characteristics, leaf total carotenoid (Car), chlorophyll (Chl) and nitrogen (N) content and leaf dry mass per area (LMA) along vertical light gradients in natural canopies of the herb species, Inula salicina and Centaurea jacea, and tree species, Populus tremula and Tilia cordata, in the middle of the growing season. Presence of stress was assessed on the basis of night measurements of chlorophyll fluorescence. Our aim was to compare the light acclimation of leaf traits, which respond to light availability at long (LMA and N), medium (Chl a/b ratio, Car/Chl ratio) and short time scales (fluorescence characteristics). We found that light acclimation of nitrogen content per unit leaf area (N(area)), chlorophyll content per unit dry mass (Chl(mass)) and Chl/N ratio were related to modifications in LMA. The maximum PSII quantum yield (F(v) /F(m)) increased with increasing growth irradiance in I. salicina and P. tremula but decreased in T. cordata. Leaf growth irradiance, N content and plant species explained the majority of variability in chlorophyll fluorescence characteristics, up to 90% for steady-state fluorescence yield, while the contribution of leaf total carotenoid content was generally not significant. Chlorophyll fluorescence characteristics did not differ strongly between growth forms, but differed among species within a given growth form. These data highlight that foliage acclimation to light is driven by interactions between traits with varying time kinetics.

Manipulation of VOC emissions with methyl jasmonate and carrageenan in the evergreen conifer Pinus sylvestris and evergreen broadleaf Quercus ilex.

Plant defence can be induced by exposing plants to the plant hormone jasmonic acid (JA) or its volatile ester, methyl jasmonate (MeJA). Carrageenans (Carr) - sulphated D-galactans extracted from red algae - can also induce plant defences. In this study, the effects of exogenous MeJA and Carr application (concentration 300 and 12.7 µmol, respectively) on volatile emissions from two widespread evergreen woody species, Pinus sylvestris (nine Turkish and one Finnish provenance) and Quercus ilex (Italian provenance) were investigated. We collected headspace samples from seedlings and analysed the quality and quantity of volatile compounds emitted by treated and control plants. In total, 19 monoterpenes, 10 sesquiterpenes, 10 green leaf volatiles (GLVs) and two aromatic compounds were emitted by P. sylvestris from all the provenances studied. Foliar MeJA application clearly affected the volatile profiles of trees from all the provenances. Effects of Carr were genotype specific. In Q. ilex, emissions of sesquiterpenes, GLVs and the homoterpene (E)-DMNT were all induced by MeJA application. However, emissions of most constitutively emitted monoterpenes were significantly reduced. Carr application also led to a significant reduction in monoterpene emissions, but without corresponding increases in other emissions. Our results indicate that exogenously applied MeJA and Carr can both significantly modify the volatile profiles of P. sylvestris and Q. ilex, but also that there are important provenance- and species-specific differences in the overall degree of elicitation and compositions of elicited compounds.

Higher allocation to low cost chemical defenses in invasive species of Hawaii.

The capacity to produce carbon-based secondary compounds (CBSC), such as phenolics (including tannins) and terpenes as defensive compounds against herbivores or against neighboring competing plants can be involved in the competition between alien and native plant species. Since the Hawaiian Islands are especially vulnerable to invasions by alien species, we compared total phenolic (TP), total tannin (Tta), and total terpene (TT) leaf contents of alien and native plants on Oahu Island (Hawaii). We analyzed 35 native and 38 alien woody plant species randomly chosen among representative current Hawaiian flora. None of these CBSC exhibited phylogenetic fingerprinting. Alien species had similar leaf TP and leaf Tta contents, and 135% higher leaf TT contents compared with native species. Alien plants had 80% higher leaf TT:N leaf content ratio than native plants. The results suggest that apart from greater growth rate and greater nutrient use, alien success in Oahu also may be linked to greater contents of low cost chemical defenses, such as terpenes, as expected in faster-growing species in resource rich regions. The higher TT contents in aliens may counterbalance their lower investment in leaf structural defenses and their higher leaf nutritional quality. The higher TT provides higher effectiveness in deterring the generalist herbivores of the introduced range, where specialist herbivores are absent. In addition, higher TT contents may favor aliens conferring higher protection against abiotic and biotic stressors. The higher terpene accumulation was independent of the alien species origin, which indicates that being alien either selects for higher terpene contents post-invasion, or that species with high terpene contents are pre-adapted to invasiveness. Although less likely, an originally lower terpene accumulation in Hawaiian than in continental plants that avoids the increased attraction of specialist enemies associated to terpenes may not be discarded.

Acclimation of photosynthetic characteristics of the moss Pleurozium schreberi to among-habitat and within-canopy light gradients.

Light availability varies strongly among moss habitats and within the moss canopy, and vertical variation in light within the canopy further interacts with the age gradient. The interacting controls by habitat and canopy light gradient and senescence have not been studied extensively. We measured light profiles, chlorophyll (Chl), carotenoid (Car) and nitrogen (N) concentrations, and photosynthetic electron transport capacity (J(max)) along habitat and canopy light gradients in the widespread, temperate moss Pleurozium schreberi to separate sources of variation in moss chemical and physiological traits. We hypothesised that this species, like typical feather mosses with both apical and lateral growth, exhibits greater plasticity in the canopy than between habitats due to deeper within-canopy light gradients. For the among-habitat light gradient, Chl, Chl/N and Chl/Car ratio increased with decreasing light availability, indicating enhanced light harvesting in lower light and higher capacity for photoprotection in higher light. N and J(max) were independent of habitat light availability. Within the upper canopy, until 50-60% above-canopy light, changes in moss chemistry and photosynthetic characteristics were analogous to patterns observed for the between-habitat light gradient. In contrast, deeper canopy layers reflected senescence of moss shoots, with pigment and nitrogen concentrations and photosynthetic capacity decreasing with light availability. Thus, variation in chemical and physiological traits within the moss canopy is a balance between acclimation and senescence. This study demonstrates extensive light-dependent variation in moss photosynthetic traits, but also that between-habitat and within-canopy light gradient affects moss physiology and chemistry differently.

Modeling volatile isoprenoid emissions--a story with split ends.

Accurate prediction of plant-generated volatile isoprenoid fluxes is necessary for reliable estimation of atmospheric ozone and aerosol formation potentials. In recent years, significant progress has been made in understanding the environmental and physiological controls on isoprenoid emission and in scaling these emissions to canopy and landscape levels. We summarize recent developments and compare different approaches for simulating volatile isoprenoid emission and scaling up to whole forest canopies with complex architecture. We show that the current developments in modeling volatile isoprenoid emissions are "split-ended" with simultaneous but separated efforts in fine-tuning the empirical emission algorithms and in constructing process-based models. In modeling volatile isoprenoid emissions, simplified leaf-level emission algorithms (Guenther algorithms) are highly successful, particularly after scaling these models up to whole regions, where the influences of different ecosystem types, ontogenetic stages, and variations in environmental conditions on emission rates and dynamics partly cancel out. However, recent experimental evidence indicates important environmental effects yet unconsidered and emphasize, the importance of a highly dynamic plant acclimation in space and time. This suggests that current parameterizations are unlikely to hold in a globally changing and dynamic environment. Therefore, long-term predictions using empirical algorithms are not necessarily reliable. We show that process-based models have large potential to capture the influence of changing environmental conditions, in particular if the leaf models are linked with physiologically based whole-plant models. This combination is also promising in considering the possible feedback impacts of emissions on plant physiological status such as mitigation of thermal and oxidative stresses by volatile isoprenoids. It might be further worth while to incorporate main features of these approaches in regional empirically-based emission estimations thereby merging the "split ends".

Monoterpene emissions from ornamental trees in urban areas: a case study of Barcelona, Spain.

Research on biogenic volatile organic compound (BVOC) emissions has mainly focused on native species in natural ecosystems. However, much of the ozone and aerosol formation occurs in city atmospheres due to BVOC emissions by local urban vegetation. Plant composition of urban habitats is often dominated by non-native ornamental plant species, for which only limited data on BVOC emissions are available. To gain insight into the influence of ornamental vegetation on the urban atmospheric reactivity in Barcelona, Spain, we studied volatile isoprenoid emissions in 11 widespread ornamental tree species (three conifers and nine angiosperms). We found significant monoterpene emissions in all studied species, with normalized emission potentials (T=30 degrees C, photosynthetic photon flux density (PPFD)=1000 micromol x m(-2) x s(-1)) ranging between 0.2 to 110 microg x g(-1) (dry weight) h(-1). Depending on species, the emissions were dominated by alpha- and beta-pinene, myrcene, alpha- and beta-phellandrene, carene, limonene and eucalyptol. These data demonstrate that ornamental plants may significantly contribute to the BVOC load in urban atmospheres and also underscore the importance of broadleaf angiosperms as significant monoterpene emitters.

Foliar limonene uptake scales positively with leaf lipid content: "non-emitting" species absorb and release monoterpenes.

Monoterpenes synthesized and released by emitting vegetation can be taken up by neighboring non-emitting plants, but the uptake capacity of non-emitting species has not been studied extensively. We investigated the foliar uptake potential of the hydrophobic monoterpene limonene in 13 species of contrasting leaf structure and lipid content to determine the structural and chemical controls of monoterpene uptake. Leaf dry mass per unit area (M(A,D)) varied 6.5-fold, dry to fresh mass ratio (D(F)) 2.7-fold, lipid content per dry mass (L(M)) 2.5-fold and per unit area (L(A)) 4.6-fold across the studied species. Average foliar limonene uptake rate (U(A)) from air at saturating limonene partial pressures varied from 0.9 to 6 nmol m(-2) s(-1), and limonene leaf to air partition coefficient (K(FA), ratio of limonene content per dry mass to limonene partial pressure) from 0.7 to 6.8 micromol kg(-1) Pa(-1). U(A) and K(FA) scaled positively with leaf lipid content, and were independent of D(F), indicating that variation in leaf lipid content was the primary determinant of species differences in monoterpene uptake rate and K(FA). Mass-based limonene uptake rates further suggested that thinner leaves with greater surface area per unit dry mass have higher uptake rates. In addition, limonene lipid to air partition coefficient (K(LA)=K(FA)/L(M)) varied 19-fold, indicating large differences in limonene uptake capacity at common leaf lipid content. We suggest that the significant uptake of hydrophobic monoterpenes when monoterpene ambient air concentration is high and release when the concentration is low should be included in large-scale monoterpene emission models.

Simultaneous growth and emission measurements demonstrate an interactive control of methanol release by leaf expansion and stomata.

Emission from plants is a major source of atmospheric methanol. Growing tissues contribute most to plant-generated methanol in the atmosphere, but there is still controversy over biological and physico-chemical controls of methanol emission. Methanol as a water-soluble compound is thought to be strongly controlled by gas-phase diffusion (stomatal conductance), but growth rate can follow a different diurnal rhythm from that of stomatal conductance, and the extent to which the emission control is shared between diffusion and growth is unclear. Growth and methanol emissions from Gossypium hirsutum, Populus deltoides, and Fagus sylvatica were measured simultaneously. Methanol emission from growing leaves was several-fold higher than that from adult leaves. A pronounced diurnal rhythm of methanol emission was observed; however, this diurnal rhythm was not predominantly determined by the diurnal rhythm of leaf growth. Large methanol emission peaks in the morning when the stomata opened were observed in all species and were explained by release of methanol that had accumulated in the intercellular air space and leaf liquid pool at night in leaves with closed stomata. Cumulative daily methanol emissions were strongly correlated with the total daily leaf growth, but the diurnal rhythm of methanol emission was modified by growth rate and stomatal conductance in a complex manner. While in G. hirsutum and in F. sylvatica maxima in methanol emission and growth coincided, maximum growth rates of P. deltoides were observed at night, while maximum methanol emissions occurred in the morning. This interspecific variation was explained by differences in the share of emission control by growth processes, by stomatal conductance, and methanol solubilization in tissue water.

Photosynthetic acclimation to simultaneous and interacting environmental stresses along natural light gradients: optimality and constraints.

There is a strong natural light gradient from the top to the bottom in plant canopies and along gap-understorey continua. Leaf structure and photosynthetic capacities change close to proportionally along these gradients, leading to maximisation of whole canopy photosynthesis. However, other environmental factors also vary within the light gradients in a correlative manner. Specifically, the leaves exposed to higher irradiance suffer from more severe heat, water, and photoinhibition stresses. Research in tree canopies and across gap-understorey gradients demonstrates that plants have a large potential to acclimate to interacting environmental limitations. The optimum temperature for photosynthetic electron transport increases with increasing growth irradiance in the canopy, improving the resistance of photosynthetic apparatus to heat stress. Stomatal constraints on photosynthesis are also larger at higher irradiance because the leaves at greater evaporative demands regulate water use more efficiently. Furthermore, upper canopy leaves are more rigid and have lower leaf osmotic potentials to improve water extraction from drying soil. The current review highlights that such an array of complex interactions significantly modifies the potential and realized whole canopy photosynthetic productivity, but also that the interactive effects cannot be simply predicted as composites of additive partial environmental stresses. We hypothesize that plant photosynthetic capacities deviate from the theoretical optimum values because of the interacting stresses in plant canopies and evolutionary trade-offs between leaf- and canopy-level plastic adjustments in light capture and use.

Development of leaf photosynthetic parameters in Betula pendula Roth leaves: correlations with photosystem I density.

The global modelling of photosynthesis is based on exact knowledge of the leaf photosynthetic machinery. The capacities of partial reactions of leaf photosynthesis develop at different rates, but it is not clear how the development of photoreactions and the Calvin cycle are co-ordinated. We investigated the development of foliar photosynthesis in the temperate deciduous tree Betula pendula Roth. using a unique integrated optical/gas exchange methodology that allows simultaneous estimation of photosystem I and II (PS I and PS II) densities per leaf area, interphotosystem electron transport activities, and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) kinetic properties. We combined these measurements with in vitro determinations of Rubisco, soluble protein and chlorophyll contents. We observed a strong increase in leaf photosynthetic capacity in developing leaves per leaf area, as well as per dry mass, that was paralleled by accumulation of leaf Rubisco. Enhanced mesophyll conductance was the outcome of increased carboxylation capacity and increased CO(2) diffusion conductance. However, Rubisco was only partly activated in the leaves, according to in vivo measurements of Rubisco kinetics. The amount of active Rubisco increased in proportion with development of PS I, probably through a direct link between Rubisco activase and PS I electron transport. Since the kinetics for post-illumination P700 re-reduction did not change, the synthesis of cytochrome b(6)f complex was also proportional to PS I. The synthesis of PS II began later and continued for several days after reaching the full PS I activity, but leaf chlorophyll was shared equally between the photosystems. Due to this, the antenna of PS II was very large and not optimally organized, leading to greater losses of excitation and lower quantum yields in young leaves. We conclude that co-ordinated development of leaf photosynthesis is regulated at the level of PS I with subordinated changes in PS II content and Rubisco activation.

Leaf-level phenotypic variability and plasticity of invasive Rhododendron ponticum and non-invasive Ilex aquifolium co-occurring at two contrasting European sites.

To understand the role of leaf-level plasticity and variability in species invasiveness, foliar characteristics were studied in relation to seasonal average integrated quantum flux density (Qint) in the understorey evergreen species Rhododendron ponticum and Ilex aquifolium at two sites. A native relict population of R. ponticum was sampled in southern Spain (Mediterranean climate), while an invasive alien population was investigated in Belgium (temperate maritime climate). Ilex aquifolium was native at both sites. Both species exhibited a significant plastic response to Qint in leaf dry mass per unit area, thickness, photosynthetic potentials, and chlorophyll contents at the two sites. However, R. ponticum exhibited a higher photosynthetic nitrogen use efficiency and larger investment of nitrogen in chlorophyll than I. aquifolium. Since leaf nitrogen (N) contents per unit dry mass were lower in R. ponticum, this species formed a larger foliar area with equal photosynthetic potential and light-harvesting efficiency compared with I. aquifolium. The foliage of R. ponticum was mechanically more resistant with larger density in the Belgian site than in the Spanish site. Mean leaf-level phenotypic plasticity was larger in the Belgian population of R. ponticum than in the Spanish population of this species and the two populations of I. aquifolium. We suggest that large fractional investments of foliar N in photosynthetic function coupled with a relatively large mean, leaf-level phenotypic plasticity may provide the primary explanation for the invasive nature and superior performance of R. ponticum at the Belgian site. With alleviation of water limitations from Mediterranean to temperate maritime climates, the invasiveness of R. ponticum may also be enhanced by the increased foliage mechanical resistance observed in the alien populations.

Site fertility and the morphological and photosynthetic acclimation of Pinus sylvestris needles to light.

Morphological and photosynthetic acclimation of current-year needles to canopy gradients in light availability (seasonal mean integrated quantum flux density, Q(int)) was studied in the temperate conifer, Pinus sylvestris L., at two sites of contrasting nutrient availability. The nutrient-rich site supported a monospecific P. sylvestris stand on an old-field. The trees were approximately 30 years old and 19-21 m tall. Mean foliar N and P contents (+/- SD) were 1.53 +/- 0.11% and 0.196 +/- 0.017%, respectively. The nutrient-poor site was located on a raised bog supporting a sparse stand of 50- to 100-year-old trees, with a height of 1-2 m, and mean needle N and P contents of 0.86 +/- 0.12% and 0.074 +/- 0.010%, respectively. At both sites, needle thickness (T) and width (W) increased with increasing Qint, and leaf dry mass per unit leaf area (MA) was also greater at higher irradiance. The light effects on MA-the product of needle density (D) and volume to total area ratio (V/AT)-resulted primarily from large increases in V/AT with Qint rather than from modifications of D, which was relatively insensitive to light. Although needle morphology versus light relationships were qualitatively similar at both sites, needles were shorter, and the slopes of W, T, MA and V/AT versus light relationships were lower, at the nutrient-poor than at the nutrient-rich site, indicating that the plasticity of foliar morphological characteristics was affected by nutrient availability. As a result of lower plasticity, needles at the nutrient-poor site were narrower, thinner, and had lower MA at high irradiance than needles at the nutrient-rich site. The maximum carboxylase activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (Vcmax) and the maximum photosynthetic electron transport rate (Jmax) scaled positively with foliar N and P contents. The correlations were generally stronger with P than with N, suggesting that needle photosynthetic capacity was more heavily limited by the availability of P than of N. The Jmax/Vcmax ratio was positively related to the foliar P/N ratio, indicating that Jmax was more strongly suppressed than Vcmax under conditions of low P availability. Phosphorus and N deficiency also limited the plasticity of foliar photosynthetic characteristics. There was a moderate increase in needle photosynthetic capacity of up to 1.6-fold from the bottom to the top of the canopy at the nutrient-rich site, but net assimilation rates were essentially independent of canopy position at the nutrient-poor site. Stomatal constraints on photosynthesis were similar between the sites, indicating that photosynthetic acclimation was curtailed at the biochemical level. We conclude that the foliar capacity for morphological and physiological acclimation to high light significantly decreases with decreasing nutrient availability in P. sylvestris, and that both N and P availability are potentially important determinants of foliar carbon gain capacities.

Responses of foliar photosynthetic electron transport, pigment stoichiometry, and stomatal conductance to interacting environmental factors in a mixed species forest canopy.

We studied limitations caused by variations in leaf temperature and soil water availability on photosynthetic electron transport rates calculated from foliar chlorophyll fluorescence analysis (U) in a natural deciduous forest canopy composed of shade-intolerant Populus tremula L. and shade-tolerant Tilia cordata Mill. In both species, there was a positive linear relationship between light-saturated U (Umax) per unit leaf area and mean seasonal integrated daily quantum flux density (Ss, mol per square m per day). Acclimation of leaf dry mass per area and nitrogen per area to growth irradiance largely accounted for this positive scaling. However, the slopes of the Umax versus Ss relationships were greater on days when leaf temperature was high than on days when leaf temperature was low. Overall, Umax varied 2.5-fold across a temperature range of 20-30 degrees C. Maximum stomatal conductance (Gmax) also scaled positively with Ss. Although Gmax observed during daily time courses, and stomatal conductances during Umax measurements declined in response to seasonally decreasing soil water contents, was insensitive to prolonged water stress, and was not strongly correlated with stomatal conductances during its estimation. These results suggest that photorespiration was an important electron sink when intercellular CO2 concentration was low because of closed stomata. Given that xanthophyll cycle pool size (VAZ, sum of violaxanthin, antheraxanthin, and zeaxanthin) may play an important role in dissipation of excess excitation energy, the response of VAZ to fluctuating light and temperature provided another possible explanation for the stable Umax. Xanthophyll cycle carotenoids per total leaf chlorophyll (VAZ/Chl) scaled positively with integrated light and negatively with daily minimum air temperature, whereas the correlation between VAZ/Chl and irradiance was best with integrated light averaged over 3 days preceding foliar sampling. We conclude that the potential capacity for electron transport is determined by long-term acclimation of U to certain canopy light conditions, and that the rapid adjustment of the capacity for excitation energy dissipation plays a significant part in the stabilization of this potential capacity. Sustained high capacity of photosynthetic electron transport during stress periods provides an explanation for the instantaneous response of U to short-term weather fluctuations, but also indicates that U restricts potential carbon gain under conditions of water limitation less than does stomatal conductance.

Variability in Leaf Morphology and Chemical Composition as a Function of Canopy Light Environment in Coexisting Deciduous Trees.

Morphology, chemical composition, and photosynthetic capacity of leaf laminas were investigated in Populus tremula L. and Tilia cordata Mill. along a canopy light gradient. Variables determining the thickness of boundary layer for heat and water exchange at a given wind speed-effective leaf width (Ww) and length (Wd)-scaled positively with daily integrated quantum flux density averaged over the season (Qint, mol m-2 d-1) in T. cordata, but Wd decreased and Ww was constant with increasing Qint in P. tremula, bringing about a moderately improved capacity for convective cooling at greater irradiances in the latter species. Foliar stable carbon isotope discrimination (Delta) decreased with increasing Qint, demonstrating that, possibly because of more severe foliar water stress, leaves operated at a lower intercellular CO2 concentration in the upper canopy. Further analysis of foliar characteristics provided additional evidence of the interaction between water stress and Qint. Leaf dry matter content and both components of lamina dry mass per area (MA)-lamina thickness and density (dry mass per unit volume, rhoB)-increased with increasing Qint in both species. The rhoB and lamina dry matter content were also positively related to lamina carbon concentration, variability in which along the canopy was related to changes in carbon-rich lignin concentration. Since both increases in lamina density and lignin concentration improve leaf tolerance of low-water potentials, these foliar modifications were interpreted as indicative of acclimation to enhanced water limitations in high light. For the whole material, foliar nitrogen concentrations decreased with increasing rhoB, suggesting that an improvement of foliar mechanical strength may result in declining foliar assimilative potential. However, foliar photosynthetic electron transport capacity per unit area increased with increasing rhoB, possibly because increases in rhoB with light are not only attributable to greater cell wall lignification but also to denser packing of leaf cells, in particular, in fractional increases in palisade tissues with Qint. Because of a positive scaling of leaf thickness and density with total tree height, MA was greater in taller trees of T. cordata, foliage of which also had lower Delta and was likely to function with less open stomata. In summary, we conclude that leaf water stress, which scales with both Qint and total tree height, is a major factor altering foliage structure and assimilative capacity.