On the role of flavonoids in the integrated mechanisms of response of Ligustrum vulgare and Phillyrea latifolia to high solar radiation

The role of flavonoids in mechanisms of acclimation to high solar radiation was analysed in Ligustrum vulgare and Phillyrea latifolia, two Mediterranean shrubs that have the same flavonoid composition but differ strikingly in their leaf morpho-anatomical traits. In plants exposed to 12 or 100% solar radiation, measurements were made for surface morphology and leaf anatomy; optical properties, photosynthetic pigments, and photosystem II efficiency; antioxidant enzymes, lipid peroxidation and phenylalanine ammonia lyase; synthesis of hydroxycinnamates and flavonoids; and the tissue-specific distribution of flavonoid aglycones and ortho-dihydroxylated B-ring flavonoid glycosides. A denser indumentum of glandular trichomes, coupled with both a thicker cuticle and a larger amount of cuticular flavonoids, allowed P. latifolia to prevent highly damaging solar wavelengths from reaching sensitive targets to a greater degree than L. vulgare. Antioxidant enzymes in P. latifolia were also more effective in countering light-induced oxidative load than those in L. vulgare. Consistently, light-induced accumulation of flavonoids in L. vulgare, particularly ortho-dihydroxylated flavonoids in the leaf mesophyll, greatly exceeded that in P. latifolia. We conclude that the accumulation of flavonoid glycosides associated with high solar radiation-induced oxidative stress and, hence, biosynthesis of flavonoids appear to be unrelated to 'tolerance' to high solar radiation in the species examined.     

Interaction effects of root-zone salinity and solar irradiance on the physiology and biochemistry of Olea europaea

Root-zone salinity stress and high solar irradiance concomitantly occurs in the Mediterranean basin, where Olea europaea is the dominating fruit-tree crop-species. Although the effect of each individual stressor on plant performance has been widely investigated, much less is known on the interaction effects of salinity stress and solar irradiance on the physiology and biochemistry of olive plants. Here we analyzed how changes in root-zone NaCl concentration and sunlight radiation affect relevant physiological and biochemical features in olive cv. Cipressino. Two-year-old plants were supplied with 0 or 125 mM NaCl and exposed to 15% (shade) or 100% sunlight (sun) over a 5-week period, starting from July 10th, 2005. Measurements were conducted of (i) gas exchange and plant growth, (ii) the concentrations of cations and chloride, (iii) the concentrations of soluble carbohydrates, violaxanthin-cycle pigments and polyphenols, and (iv) the protein oxidation and the lipid peroxidation in the leaves. Salt-induced reductions in gas exchange performance and plant growth were greater at the sun than at the shade site, mostly due to light-induced changes in leaf water relations and vapour pressure deficit (vpd), rather than in the concentration of potentially toxic ions. Light-induced increases in leaf Na⁺ and Cl^? concentrations were countered by parallel enhancements in the concentrations of K⁺ and Ca²⁺. Sun leaves had sharply greater concentrations of mannitol and xanthophylls, irrespective of root-zone salinity. The amount of “newly assimilate carbon” allocated to polyphenols, especially to flavonoids, increased in response to salinity stress and high sunlight. Remarkably, the protein oxidation was greater in shade than in sun leaves of well-watered plants, and increased more at the shade than at the sun site because of high salinity. We suggest that heat-stress (on average maximum T exceeded 33 °C for 50% of the experimental period), which acted in concert with salinity stress and sunlight irradiance in determining plant responses in our experiment, was responsible for leaf oxidative damage in plants growing under contrasting solar radiation. Indeed, sun leaves of salt-stressed plants were equipped with an extraordinary-rich arsenal of antioxidant compounds, distributed in different cell compartments, i.e., mannitol, zeaxanthin and flavonoids, which likely countered effectively the oxidative damage driven by heat-stress, a clear example of cross-tolerance.     

Responses to changes in Ca2+ supply in two Mediterranean evergreens, Phillyrea latifolia and Pistacia lentiscus, during salinity stress and subsequent relief

BACKGROUND AND AIMS: Changes in root-zone Ca(2+) concentration affect a plant's performance under high salinity, an issue poorly investigated for Mediterranean xerophytes, which may suffer from transient root-zone salinity stress in calcareous soils. It was hypothesized that high-Ca(2+) supply may affect differentially the response to salinity stress of species differing in their strategy of Na(+) allocation at organ level. Phillyrea latifolia and Pistacia lentiscus, which have been reported to greatly differ for Na(+) uptake and transport rates to the leaves, were studied. Methods In plants exposed to 0 mM or 200 mM NaCl and supplied with 2.0 mM or 8.0 mM Ca(2+), under 100 % solar irradiance, measurements were conducted of (a) gas exchange, PSII photochemistry and plant growth; (b) water and ionic relations; (c) the activity of superoxide dismutase and the lipid peroxidation; and (d) the concentration of individual polyphenols. Gas exchange and plant growth were also estimated during a period of relief from salinity stress. Key Results The performance of Pistacia lentiscus decreased to a significantly smaller degree than that of Phillyrea latifolia because of high salinity. Ameliorative effects of high-Ca(2+) supply were more evident in Phillyrea latifolia than in Pistacia lentiscus. High-Ca(2+) reduced steeply the Na(+) transport to the leaves in salt-treated Phillyrea latifolia, and allowed a faster recovery of gas exchange and growth rates as compared with low-Ca(2+) plants, during the period of relief from salinity. Salt-induced biochemical adjustments, mostly devoted to counter salt-induced oxidative damage, were greater in Phillyrea latifolia than in Pistacia lentiscus. CONCLUSIONS: An increased Ca(2+) : Na(+) ratio may be of greater benefit for Phillyrea latifolia than for Pistacia lentiscus, as in the former, adaptive mechanisms to high root-zone salinity are primarily devoted to restrict the accumulation of potentially toxic ions in sensitive shoot organs.     

The biosynthesis of flavonoids is enhanced similarly by UV radiation and root zone salinity in L. vulgare leaves

Flavonoids have recently been suggested to have the potential to serve as antioxidants other than effective UV attenuators in photoprotection. Here, we tested the hypothesis that flavonoids accumulate in response to “excess light” in the presence or in the absence of UV radiation. In a UV exclusion experiment, we grew Ligustrum vulgare plants outdoors under 30% or 100% sunlight irradiance, by cutting-off the whole UV waveband. These plants were also exposed to UV irradiance or supplied with 125 mM NaCl at the root zone. Leaves of plants under 100% sunlight irradiance suffered from excess light, which was exacerbated greatly by root zone salinity stress. Salinity stress repressed the activities of antioxidant enzymes, particularly in full sunlight, and led to severe leaf oxidative damage. Dihydroxy B-ring-substituted flavonoids, namely quercetin 3-O- and luteolin 7-O-glycosides, accumulated steeply in response to sunlight irradiance in the absence of UV radiation. UV radiation and root zone NaCl increased, to a similar degree, the concentration of these flavonoids, which have a great potential to scavenge various forms of reactive oxygen. Treatment-induced changes in leaf phenylpropanoid concentration affected antioxidant activities to a greater extent than the UV-screening capacities of leaf extracts. Early responses to an abrupt increase in sunlight irradiance included a steep increase in the concentrations of quercetin derivatives and cyanidin 3-O-glucoside, with the latter negligibly absorbing in the UV-spectral region. In contrast, effective UV attenuators, such as hydroxycinnamates and monohydroxy B-ring flavonoids, were unresponsive to the light treatments. Overall, these findings lead to the hypothesis that flavonoids may have an important antioxidant function in photoprotection. This hypothesis is further corroborated by the large distribution of quercetin and luteolin derivatives in the vacuoles of mesophyll, not only in the corresponding compartments of epidermal cells, but also in full sunlight-treated leaves in the absence of UV radiation. Future experiments aimed at evaluating the relative contribution of flavonoids within the complex antioxidant defense systems operating in the leaf are needed to help conclusively address the relevance of their antioxidant functions in photoprotection.     

Mediterranean sea cliff plants: morphological and physiological responses to environmental conditions

Aims The plants of Mediterranean sea cliff ecosystems are resistant to several environmental challenges. In this study, six species typical of the coastal rocky cliffs have been analyzed in order to evaluate their diverse morphological and physiological responses to their environment across the seasons, and to examine the strategy of the ecological group to which each species belongs. Since these species are widespread across the Mediterranean region, our aim was also to highlight their ecophysiological features in habitats where the direct influence of the sea is stronger.Methods The selected species are characteristic of the sea cliffs of Elba island (Tyrrhenian sea, Italy): the halophyte Crithmum maritimum, the semideciduous Helichrysum italicum and Lavandula stoechas and the sclerophylls Myrtus communis, Quercus ilex and Rhamnus alaternus. Four morphological traits—canopy height, leaf area, specific leaf area and leaf dry matter content—and two physiological traits—leaf water potential (LWP) and photosynthetic efficiency (PE), measured before the dawn and at midday—were analyzed. Water potential was measured by a pressure chamber and photosynthetic efficiency was determined by the analysis of chlorophyll fluorescence. Plant performance was also evaluated by calculating chronic (PI chr) and dynamic photoinhibition (PI dyn).Important findings Crithmum maritimum showed high resistance to the recurrent dry periods, because of the high water storage capacity of its leaves and its PE declined markedly only in July, under the harshest climatic conditions. Semideciduous taxa utilize primarily an avoidance strategy, which aims at reducing the overall leaf surface, while sclerophylls mostly show a tolerance strategy towards the prevailing stressors, as demonstrated by LWP and PE, that are lower in the sclerophylls than in the semideciduous taxa during summer, due to osmoregulation and photoinhibition, respectively. Furthermore, variability of physiological parameters was higher in the sclerophylls than in the semideciduous taxa, because the former had to withstand wider oscillations of their LWP and PE. The sclerophyllous taxa underwent a slight loss of PE also in winter, likely owing to the combined action of low temperature and high irradiance. In Mediterranean sea cliff ecosystems, the stressful combination of high irradiance, high temperatures and low rainfall typical of the summer season may have been intensified by the shallow soil which displays a poor water storage capacity. On the other hand, winter stress, caused by high solar radiation and low temperatures, does not seem to seriously affect the performance of the studied species.     

Cavitation,stomatal conductance,and leaf dieback in seedlings of two co-occurring Mediterranean shrubs during an intense drought

Seedling shrubs in the Mediterranean semi-arid climate are subjected to intense droughts during summer. Thus, seedlings often surpass their limits of tolerance to water stress, resulting in the loss of hydraulic conductivity due to xylem cavitation. The response in terms of stomatal conductance, vulnerability to cavitation, leaf dieback, and survival were analysed in two co-occurring seedlings of mastic tree (Pistacia lentiscus L.) and kermes oak (Quercus coccifera L.) during an intense drought period. Both species reacted to drought with steep decreases in stomatal conductance before the critical water potential brought about the onset of cavitation events. Q. coccifera showed wider safety margins for avoiding runaway embolism than P. lentiscus and these differences could be related to the particular drought strategy displayed by each species: water saver or water spender. The limits for survival, resprout capacity and leaf dieback were also analysed in terms of loss of conductivity. By contrast with previous studies, the species showing higher seedling survival in the presence of drought also showed higher susceptibility to cavitation and operated with a lower safety margin for cavitation. Both species showed a leaf specific conductivity (LSC) threshold below which leaf biomass had to be regulated to avoid runaway embolism. However, each species displayed a different type of response: P. lentiscus conserved total leaf area up to 100% loss of LSC, whereas Q. coccifera continuously adjusted leaf biomass throughout the drought period in order to maintain the LSC very close to the maximum values recorded without loss of conductivity. Both species maintained the capacity for survival until the loss of conductivity was very nearly 100%.     

Floral and vegetative morphometrics of three Platonia insignis Mart. (Clusiaceae) populations,a native tree from the Brazilian Amazon

Platonia insignis Mart. (Clusiaceae), the bacurizeiro, is a native tree species from the Brazilian Amazon forests. Three populations of P. insignis have been observed in the north-east region of the state of Maranhão that differ in flower color: the red population that produces dark pink flowers, the pink population that produces light pink flowers, and the white population with yellowish-white flowers. From multivariate statistical analysis, we aimed at characterizing such populations using morpho-anatomical leaf and flower morphology parameters. A total of 40 P. insignis individuals have been sampled in the cities of São Luís and Chapadinha. The morphological traits varied more than the anatomical traits. Area, fresh mass, and dry mass were the leaf parameters that show more variations. Platonia insignis have hypostomatic or amphihypostomatic leaves. The length of the gynoecium+the length of the nectary, the total length and the length of gynoecium were the principal components considering flower analysis. The three populations did not show significant differences nor did they group using Ward's method. Individuals from the Chapadinha and São Luís red population have been separated according to leaf and flower morphological traits, and the morphological difference between individuals may represent early stages of geographical speciation.     

Non-random leaf orientation in Lactuca serriola L.

Abstract Leaf shapes and leaf orientation of Lactuca serriola serriola and serriola integrifolia were studied. Leaf shapes in L. serriola serriola differed greatly from those of L.serriola integrifolia, but leaf surface areas were similar. In exposed habitats, leaf orientation of cauline leaves of both forms was non-random, with leaves almost vertical and tending to orient with their lamina normal to the east and west. In the shade, cauline leaves oriented randomly. An experiment demonstrated that the orientation of leaves did not change significantly once they were fully expanded. The leaf orientation in L. serriola affected the diurnal distribution of solar irradiance intercepted by a leaf. Peak solar radiation fluxes are incident on the rosette leaves at midday, but on the cauline leaves the peak solar radiation flux occurs early in the morning and again late in the afternoon. The significance of this unusual leaf orientation is discussed in relation to water loss and carbon gain.     

Variation in leaf longevity of Pistacia lentiscus and its relationship to sex and drought stress inferred from leaf δ13C

1. Leaf formation, loss, retention, longevity and biomass on male branches of the evergreen mediterranean shrub Pistacia lentiscus , L. correlated strongly with water-use efficiency inferred from leaf δ¹³C across a gradient of precipitation on the island of Mallorca, Spain.
2. The correlations suggest that the leaf phenology is under control of drought-induced constraints on the carbon balance.
3. In fruiting female branches, the correlations between the inferred water-use efficiency and number of formed and retained leaves, leaf biomass and leaf longevity were non-significant. Leaf formation was strongly reduced by fruiting and the females compensated the reduced photosynthetic capacity by retaining older leaves for a longer time than male plants.
4. It is suggested that leaf longevity in females is under strong control of resource allocation to fruit formation which is 'overlaid' on the drought-induced carbon stress, which led to the observed longer leaf longevity in females than in males.     

祁连山高寒草地甘肃臭草叶性状与坡向间的关系北大核心CSCD

植物功能性状可反映植物对环境的适应。在祁连山高寒退化草地,利用Arc GIS建立研究区域的数字高程模型,提取坡向数据,采用标准化主轴估计方法(standardized major axis estimation,SMA),研究了不同坡向甘肃臭草(Melica przewalskyi)叶性状间的关系。结果表明:1)甘肃臭草叶面积和叶体积在北坡最大,东坡、西坡、南坡逐渐减小,叶干质量无显著变化;2)叶面积与叶干质量在北坡、东坡和西坡呈等速生长关系,在南坡呈异速生长关系,且叶干质量的增长速度大于叶面积的增长速度;3)叶体积与叶干质量在4个坡向上呈异速生长关系,且叶干质量的增长速度均小于叶体积的增长速度。甘肃臭草叶性状关系随坡向的变化反映了该物种在异质生境中具有较强的叶片形态可塑性,从而有利于其适应和占据高寒退化生境。     

Leaf Physiological Traits and their Importance for Species Success in a Mediterranean Grassland

We related leaf physiological traits of four grassland species (Poa pratensis, Lolium perenne, Festuca valida, and Taraxacum officinale), dominant in a Mediterranean grassland, to their origin and success at community level. From early May to mid-June 1999, four leaf samplings were done. Species originating from poor environments (P. pratensis, F. valida) had low carbon isotope discrimination (), specific leaf area (SLA), leaf water and mineral contents, and net photosynthetic rate on mass basis (P _mass) but high chlorophyll content. The reverse traits were evident for the fast-growing species (L. perenne, T. officinale). Under the resource-limiting conditions (soil nitrogen and water) of the Mediterranean grassland, the physiological traits of P. pratensis and F. valida showed to be more adapted to these conditions leading to high species abundance and dominance.     

Photo- and antioxidative protection during summer leaf senescence in Pistacia lentiscus L. grown under Mediterranean field conditions

Summer leaf senescence in Pistacia lentiscus L. plants serves to remobilize nutrients from the oldest leaves to the youngest ones, and therefore contributes to plant survival during the adverse climatic conditions typical of Mediterranean summers, i.e. water deficit superimposed on high solar radiation and high temperatures. To evaluate the extent of photo- and antioxidative protection during leaf senescence of this species, changes in carotenoids, including xanthophyll cycle pigments, and in the levels of ascorbate and alpha-tocopherol were measured prior to and during summer leaf senescence in 3-year-old plants grown under Mediterranean field conditions. Although a chlorophyll loss of approx. 20% was observed during the first stages of leaf senescence, no damage to the photosynthetic apparatus occurred as indicated by constant maximum efficiencies of photosystem II photochemistry. During this period the de-epoxidation state of the xanthophyll cycle, and lutein, neoxanthin and ascorbate levels were kept constant. At the same time beta-carotene and alpha-tocopherol levels increased by approx. 9 and 70%, respectively, presumably conferring photo- and antioxidative protection to the photosynthetic apparatus. By contrast, during the later stages of leaf senescence, characterized by severe chlorophyll loss, carotenoids were moderately degraded (neoxanthin by approx. 20%, and both lutein and beta-carotene by approx. 35%), ascorbate decreased by approx. 80% and alpha-tocopherol was not detected in senescing leaves. This study demonstrates that mechanisms of photo- and antioxidative protection may play a major role in maintaining chloroplast function during the first stages of leaf senescence, while antioxidant defences are lost during the latest stages of senescence.     

Exploring Biodiversity in the Bacterial Community of the Mediterranean Phyllosphere and its Relationship with Airborne Bacteria

We studied the structure and diversity of the phyllosphere bacterial community of a Mediterranean ecosystem, in summer, the most stressful season in this environment. To this aim, we selected nine dominant perennial species, namely Arbutus unedo, Cistus incanus, Lavandula stoechas, Myrtus communis, Phillyrea latifolia, Pistacia lentiscus, Quercus coccifera (woody), Calamintha nepeta, and Melissa officinalis (herbaceous). We also examined the extent to which airborne bacteria resemble the epiphytic ones. Genotype composition of the leaf and airborne bacteria was analysed by using denaturing gradient gel electrophoresis profiling of a 16S rDNA gene fragment; 75 bands were cloned and sequenced corresponding to 28 taxa. Of these, two were found both in the air and the phyllosphere, eight only in the air, and the remaining 18 only in the phyllosphere. Only four taxa were found on leaves of all nine plant species. Cluster analysis showed highest similarity for the five evergreen sclerophyllous species. Aromatic plants were not grouped all together: the representatives of Lamiaceae, bearing both glandular and non-glandular trichomes, formed a separate group, whereas the aromatic and evergreen sclerophyllous M. communis was grouped with the other species of the same habit. The epiphytic communities that were the richest in bacterial taxa were those of C. nepeta and M. officinalis (Lamiaceae). Our results highlight the remarkable presence of lactic acid bacteria in the phyllosphere under the harsh conditions of the Mediterranean summer, the profound dissimilarity in the structure of bacterial communities in phyllosphere and air, and the remarkable differences of leaf microbial communities on neighbouring plants subjected to similar microbial inocula; they also point to the importance of the leaf glandular trichome in determining colonization patterns.     

散孔材与环孔材树种枝干、叶水力学特性的比较研究

为揭示散孔材与环孔材树种树木水分生理特性的差异,选取了常见的3种散孔材落叶树种(毛白杨、法国梧桐和樱花)和3种环孔材落叶树种(刺槐、合欢和白蜡),研究了其枝干与叶水力学性质的差异及其协调性。结果表明:3种环孔材树种枝干横截面积基础上的最大比导水率(Ks-max)大于3种散孔材树种,但其木质部对空穴化的脆弱性(P50branch)高于散孔材树种,6种树木枝干的水分传输能力和抵抗空穴化能力之间存在一种相互制约的权衡关系。3种散孔材与3种环孔材树种的叶最大水力导度(Kl-max)和水力脆弱性(P50leaf)并无显著差异;对于3种散孔材树种,叶的水力脆弱性要高于枝干,但对3种环孔材树种而言,枝干的水力脆弱性要高于叶。6种树木枝干和叶的水力学性质(Kmax、P50)之间并无相关关系。这些结果表明:散孔材与环孔材树种的枝干水力学特性有明显差异,但叶水力学特性无差异;枝干与叶水力学性质之间是相互独立的。     

C:N:P Stoichiometry and Leaf Traits of Halophytes in an Arid Saline Environment,Northwest China

Salinization is an important and increasingly prevalent issue which has broad and profound effects on plant survival and distribution pattern. To understand the patterns and potential drivers of leaf traits in saline environments, we determined the soil properties, leaf morphological traits (specific leaf area, SLA, and leaf dry matter content, LDMC), leaf chemical traits (leaf carbon, C, nitrogen, N, and phosphorus, P, stoichiometry) based on 142 observations collected from 23 sites in an arid saline environment, which is a vulnerable ecosystem in northwest China. We also explored the relationships among leaf traits, the responses of leaf traits, and plant functional groups (herb, woody, and succulent woody) to various saline environments. The arid desert halophytes were characterized by lower leaf C and SLA levels, higher N, but stable P and N:P. The leaf morphological traits were correlated significantly with the C, N, and P contents across all observations, but they differed within each functional group. Succulent woody plants had the lowest leaf C and highest leaf N levels among the three functional groups. The growth of halophytes might be more limited by N rather than P in the study area. GLM analysis demonstrated that the soil available nutrients and plant functional groups, but not salinity, were potential drivers of leaf C:N:P stoichiometry in halophytes, whereas species differences accounted for the largest contributions to leaf morphological variations. Our study provides baseline information to facilitate the management and restoration of arid saline desert ecosystem.     

Phenotypic plasticity in response to light in the coffee tree

Phenotypic plasticity to light availability was examined at the leaf level in field-grown coffee trees (Coffea arabica). This species has been traditionally considered as shade-demanding, although it performs well without shade and even out-yields shaded coffee. Specifically, we focused our attention on the morpho-anatomical plasticity, the balance between light capture and excess light energy dissipation, as well as on physiological traits associated with carbon gain. A wide natural light gradient, i.e., a diurnal intercepted photon irradiance differing by a factor of 25 between the deepest shade leaves and the more exposed leaves in the canopy, was explored. Responses of most traits to light were non-linear, revealing the classic leaf sun vs. leaf shade dichotomy (e.g., compared with sun leaves, shade leaves had a lower stomatal density, a thinner palisade mesophyll, a higher specific leaf area, an improved light capture, a lower respiration rate, a lower light compensating point and a limited capacity for photoprotection). The light-saturated rates of net photosynthesis were higher in sunlit than in shade leaves, although sun leaves were not efficient enough to use the extra light supply. However, sun leaves showed well-developed photoprotection mechanisms in comparison to shade leaves, which proved sufficient for avoiding photoinhibition. Specifically, a higher non-photochemical quenching coefficient was found in parallel to increases in: (i) zeaxanthin pools, (ii) de-epoxidation state of the xanthophyll cycle, and (iii) activities of some antioxidant enzymes. Intracanopy plasticity depended on the suite of traits considered, and was high for some physiological traits associated with photoprotection and maintenance of a positive carbon balance under low light, but low for most morpho-anatomical features. Our data largely explain the successful cultivation of the coffee tree in both exposed and shade environments, although with a poor resource-use efficiency in high light.     

Twig morphology and anatomy of Mediterranean trees and shrubs related to drought tolerance

De Micco V. and Aronne G. 2008. Twig morphology and anatomy of Mediterranean trees and shrubs related to drought tolerance. Bot. Helv. 118: 139 – 148. Woody species populating Mediterranean ecosystems have evolved phenological, physiological and morphological adaptations to summer drought, but their degree of drought tolerance varies, so that different species dominate along gradients from xeric to more mesic environments. In this study, we analysed morpho-anatomical properties of leaves and twigs of eight Mediterranean woody species. To test whether there is a consistent pattern of adaptation to drought, we hypothesised that the rank order of species along the xeric-mesic gradient would be reflected by increasing or decreasing trends in twig properties or in relationships between leaf area and branch size. One-year-old twigs were sampled from plants growing in a common site on the Tresino promontory in southern Italy. Measurements included leaf number and area, branch length and diameter, as well as the relative amounts of pith, xylem and cortex tissues at the basis of branches. All traits varied significantly among species, but none of them correlated with the rank order of species along the xeric-mesic gradient. In addition, the relationship between leaf area and branch size (length or diameter) was not consistently related to the drought tolerance of the species. Inconsistencies could partly be explained by variations in twig anatomy. These results support the view that not single traits, but combinations of morpho-anatomical features influence the overall capability of species to withstand dry conditions. Submitted 12 May 2008; Accepted 24 September 2008 Subject editor: Sabine Güsewell     

Leaf physiological traits in relation to resprouter ability in the Mediterranean Basin

In Mediterranean ecosystems, fire is a strong selective agent among plants, and the different post-fire regeneration strategies (e.g. resprouting and non-resprouting) have implications for other plant traits. Because young plants of non-resprouters need to grow quickly and mature well before the next fire, we predict that they should possess leaf traits related to increased efficiency in growth and resource acquisition compared with resprouter species. To test this hypothesis, we measured specific leaf area, leaf nitrogen and carbon concentrations and leaf physiological traits, including gas exchange parameters and chlorophyll fluorescence, in 19 Mediterranean species cultivated in a common garden. Both cross-species and phylogenetically informed analyses suggest that non-resprouters have better physiological performance at the leaf level (i.e. higher photosynthetic capacity) than resprouters. All these results suggest that non-resprouter species are able to take greater advantage for vegetative growth and carbon fixation than resprouters during periods when water is readily available. The contrasted physiological differences between resprouters and non-resprouters reinforce the idea that these two syndromes are functionally different (i.e. they are functional types).     

Relationships between carbon isotope discrimination, dry matter production, and harvest index in durum wheat

Carbon isotope discrimination has been proposed as a criterion for the indirect selection to improve transpiration efficiency and grain yield in bread wheat and barley. Less attention has been devoted to durumwheat (Triticum durumDesf.) despite its economic importance in the Mediterranean basin. The Δ genetic variation and its relationships to dry matter production and harvest index in durum wheat were investigated in this study. For this purpose, field experiments were conducted on 144 durum wheat accessions under Mediterranean conditions (South of France) during three consecutive years with contrasting climatic conditions. Grain yield, above-ground biomass, harvest index, and carbon isotope discrimination of flag leaf and kernel were measured. Differences between years, noted for both leaf and kernel carbon isotope discrimination, were probably related to the variation in water availability from year to year. A large genotypic variation was also noticed for both leaf and kernel carbon isotope discrimination. The two traits were found to be positively correlated with grain yield within and across years, which confirms the interest in carbon isotope discrimination for selection for grain yield improvement under Mediterranean conditions. Both kernel and leaf carbon isotope discrimination correlated better with harvest index than with grain yield, suggesting that carbon isotope discrimination could reflect the efficiency of carbon partitioning to the kernel. The lack of correlation between leaf carbon isotope discrimination and both harvest index and grain yield in favourable water conditions (1996) was probably due to the difference in water availability between the period until flag leaves sampling (favourable conditions) and the strong water stress which accompanied the grain filling. Kernel carbon isotope discrimination correlated better with both harvest index and grain yield than did leaf carbon isotope discrimination. Moreover, a higher broad-sense heritability was obtained for kernel carbon isotope discrimination than for leaf carbon isotope discrimination. As a result, kernel carbon isotope discrimination appeared to be a better predictive criterion for efficiency of the carbon partitioning to the kernel (harvest index), and hence for grain yield, than did flag leaf carbon isotope discrimination.     

Construction costs and physico-chemical properties of the assimilatory organs of Nepenthes species in Northern Borneo

BACKGROUND AND AIMS: Species of the Nepenthaceae family are under-represented in studies of leaf traits and the consequent view of mineral nutrition and limitation in carnivorous plants. This study is aimed to complement existing data on leaf traits of carnivorous plants. METHODS: Physico-chemical properties, including construction costs (CC), of the assimilatory organs (leaf and pitcher) of a guild of lowland Nepenthes species inhabiting heath and/or peat swamp forests of Brunei, Northern Borneo were determined. KEY RESULTS: Stoichiometry analyses indicate that Nepenthes species are nitrogen limited. Most traits vary appreciably across species, but greater variations exist between the assimilatory organs. Organ mass per unit area, dry matter tissue concentration (density), nitrogen (N), phosphorus (P), carbon, heat of combustion (H(c)) and CC values were higher in the leaf relative to the pitcher, while organ thickness, potassium (K) and ash showed the opposite trend. Cross-species correlations indicate that joint rather than individual consideration of the leaf and the pitcher give better predictive relationships between variables, signalling tight coupling and functional interdependence of the two assimilatory organs. Across species, mass-based CC did not vary with N or P, but increases significantly with tissue density, carbon and H(c), and decreases with K and ash contents. Area-based CC gave the same trends (though weaker in strength) in addition to a significant positive correlation with tissue mass per unit area. CONCLUSIONS: The lower CC value for the pitcher is in agreement with the concept of low marginal cost for carnivory relative to conventional autotrophy. The poor explanatory power of N, P or N : P ratio with CC suggests that factors other than production of expensive photosynthetic machinery (which calls for a high N input), including concentrations of lignin, wax/lipids or osmoregulatory ions like K(+), may give a better explanation of the CC variation across Nepenthes species.