Seasonal dimorphism in wood anatomy of the Mediterranean Cistus incanus L. subsp. incanus

In Mediterranean-type ecosystems, seasonal dimorphism is an adaptive strategy aimed to save water by developing brachyblasts with xeromorphic summer leaves as opposed to dolichoblasts with more mesomorphic winter leaves. The aim of this study was to analyse the anatomical properties of 1-year-old twigs of Cistus incanus subsp. incanus, a seasonally dimorphic shrub, to highlight properties allowing its adaptation to the Mediterranean environment. A more specific purpose was to verify the occurrence of seasonal dimorphism in wood anatomy in order to understand: (a) whether and to what extent the traits of efficiency/safety of water transport are expressed in brachyblasts and dolichoblasts, and (b) the effects on the formation of growth ring boundaries in wood. Our overall analysis showed that anatomical features of branches in C. incanus are designed to: (a) protect from desiccation by developing thick cuticle and suberized epidermal and sub-epidermal layers; (b) defend the plant from predators by accumulating phenolics; and (c) regulate water transport through the development of specific wood anatomy, according to the season, thus optimising properties of efficiency/safety. Regarding the latter point, our results indicated that brachyblast wood is safer than dolichoblast wood insofar as it has narrower and more frequent vessels; measurement of other specific anatomical traits, such as vessel wall thickness, suggested that brachyblast wood has a higher resistance to implosion due to drought-induced embolism. Finally, peculiar anatomy of brachyblast and dolichoblast wood results in the formation of so-called false rings. Hence, wood rings in C. incanus should be considered to be “seasonal” rather than “annual”.     

Leaf Structural Peculiarities in Sarcopoterium spinosum, a Seasonally Dimorphic Subshrub

Extensive investigations on the anatomy of the two leaf typesin a seasonally dimorphic subshrub revealed interesting variationsbetween summer and winter leaves. Summer leaves of Sarcopoteriumspinosum possess a thick epidermis composed of tannin-containingcells and large amounts of mucilage secreted through the innerpericlinal walls towards the mesophyll. A thick cuticle is alsopresent on the surface of the leaf. In winter leaves the epidermalcells produce no mucilage while phenolics are accumulated ingranular form only. Besides these, some other variations betweensummer and winter leaves are also discussed in respect of theability of the plant to withstand the unfavourable Mediterraneanconditions. Seasonal dimorphism, leaf anatomy, Sarcopoterium spinosum     

Physiological,morphological and anatomical trait variations between winter and summer leaves of Cistus species

Morphological, anatomical and physiological summer and winter leaf traits of Cistus incanus subsp. incanus, C. salvifolius and C. monspeliensis growing at the Botanical garden of Rome were analyzed. With regard to differences between summer and winter leaves of the considered species, leaf thickness (L) was 21% higher in summer than in winter leaves (mean of the considered species) and this increase was mostly the result of the increased palisade parenchyma thickness over the spongy parenchyma one (24 and 16% higher in summer than in winter leaves, respectively). Leaf mass area (LMA) and leaf tissue density (LTD) were 38% and 17% higher in summer than in winter leaves, respectively (mean of the considered species). The photosynthetic rate (P_N), stomatal conductance (g_s) and chlorophyll content (Chl) of summer leaves were 54%, 17% and 14% lower, respectively, than in winter leaves. C. monspeliensis summer leaves had the highest LMA, LTD, adaxial cuticle thickness (14.6 ± 1.8 mg cm⁻², 1091 ± 94 mg cm⁻³, and 5.8 ± 1.7 μm, respectively) and the lowest mesophyll intercellular spaces (f_ias 38 ± 3%). Moreover, C. monspeliensis had the highest P_N in summer (2.6 ± 0.1 μmol m⁻² s⁻¹) and C. incanus the highest P_N and WUE (84% and 59% higher than the other species) in the favorable period, associated to a higher f_ias (42 ± 2%). C. salvifolius had the highest P_N (54% higher than the other species) in winter. The plasticity index could allow a better interpretation of the habitat preference of the considered species. The physiological plasticity (PI_p = 0.39, mean value of the considered species) was higher than the morphological (PI_m = 0.22, mean value) and anatomical (PI_a = 0.13, mean value) plasticity. Moreover, among the considered species, C. salvifolius and C. incanus are characterized by a larger PI_a (0.14, mean value) which seems to be correlated with their wider ecological distribution and the more favorable conditions of the environments where they naturally occur. The highest PI_m (0.29) of C. monspeliensis indicates that it can play a high adaptive role in highly stressed environments, like fire degraded Mediterranean areas in which it occurs.     

Leaf Anatomy,Inclination, and Gas Exchange Relationships in Evergreen Sclerophqldous and Drought Semideciduous Shrub Species

There are significant differences in leaf life-span among evergreen sclerophyllous species and drought semideciduous species growing in the Mediterranean maquis. Cistus incamus, which has a leaf life-span of four-eight months, was characterised by the highest net photosynthetic rates (P _N), while Quercus ilex and Phillyrea latifolia, which maintain their leaves two-three and two-four years, respectively, had a lower P _N. The longer leaf life-span of the two evergreen sclerophyllous species may be justified to cover the high production costs of leaf protective structures such as cuticle, hairs, and sclereids: cuticle and hairs screen radiation penetrating into the more sensitive tissues, and sclereids have a light-guiding function. Q. ilex and P. latifolia have the highest leaf mass/area ratio (LMA = 209 g m^-2) and a mesophyll leaf density (2065 cells per mm² of leaf cross section area) about two times higher than C. incanus. In the typical evergreen sclerophyllous species the steepest leaf inclination ( = 56°) reduces 42 % of radiation absorption, resulting in a reduced physiological stress at leaf level, particularly in summer. C. incanus, because of its low leaf life-span, requires a lower leaf investment in leaf protective structures. It exhibits a drastic reduction of winter leaves just before summer drought, replacing them with smaller folded leaves. The lower leaf inclination ( = 44°) and the lower LMA (119 g m^-2) of C. incanus complement photosynthetic performance. Water use efficiency (WUE) showed the same trend in Q. ilex, P. latifolia, and C. incanus, decreasing 60 % from spring to summer, due to the combined effects of decreased CO₂ uptake and increased transpirational water loss.     

Antioxidant activity and protective effect on DNA cleavage of extracts from Cistus incanus L. and Cistus monspeliensis L.

The genus Cistus includes many typical species of Mediterranean flora; Cistus species are used as antidiarrheics, as general remedies for treatment of various skin diseases in folk medicine and as anti-inflammatory agents. These species contain flavonoids that are considered to be chain-breaking antioxidants. In this work, we have investigated the effects of crude aqueous extracts from Cistus incanus and Cistus monspeliensis on DNA cleavage and their free-radical scavenging capacity. In addition, their effect on lipid peroxidation in rat liver microsomes was evaluated. These extracts showed a protective effect on DNA cleavage and a dose-dependent free-radical scavenging capacity; Cistus monspeliensis was more active than Cistus incanus; these results were confirmed by a significant inhibition of lipid peroxidation in rat liver microsomes. The experimental evidence, therefore, suggests that because of their antioxidant activity these extracts may offer excellent photoprotection for skin and may be useful in the treatment of human diseases where oxidative stress plays a key role. This revised version was published online in July 2006 with corrections to the Cover Date.     

Earlier summer drought affects leaf functioning of the Mediterranean species Cistus monspeliensis L.

The Mediterranean vegetation is characterized by a high diversity of growth forms, habits and phenology that enable it to endure under harsh environmental conditions. It is however unclear whether these adaptations may allow plant survival under more extreme conditions, as predicted by climatic models under the perspective of climate change. A manipulative experiment aiming at anticipating summer aridity has been run to analyse the effects of the experimental drought on spring-leaf functioning and characteristics of the leaf-dimorphic Mediterranean shrub Cistus monspeliensis L.Assimilation rates were reduced under anticipated summer aridity due to a decrease of stomatal conductance, but only before morphological adaptations to drought (an increase of leaf mass per area) occurred. These adaptations were anticipated under experimental dry conditions, and causes photosynthetic performances to recover compared to previous dates. When natural summer aridity occurred, the leaf mass per area also changed in the control. However, this causes no recovery of the photosynthetic performances, because of the decrease of stomatal conductance due to low soil water content and leaf water potential values. Moreover, under experimental drought, leaf shedding was anticipated to reduce water losses, causing an overall reduction of leaf lifespan.     

Morphological and physiological modifications of Cistus salvifolius L. winter leaves in response to the rise in winter temperatures

The global climate is predicted to change in the next century; for the Mediterranean Basin, an increase in air temperature more than 4°C and a higher frequency of extreme climatic events such as drought and heat waves are expected. In this work, the response of Cistus salvifolius L. to the rise in winter temperature has been studied. Plants acclimated to winter conditions [outdoor (OUT)] were moved into a greenhouse [indoor (IND)] at higher temperature and eco-physiological behaviour was analysed on leaves after 15 days from plant transferring (IND_15d) and on leaves developed IND. IND leaves were characterized by reduced thickness, higher specific leaf area, higher CO₂ mesophyll conductance and photosynthetic rate, and lower respiratory rate than leaves grown OUT upon current winter conditions. In IND_15d leaves, no improvement of photochemical activity was found. When IND leaves were subjected to a rapid increase in air temperature, the CO₂ fixation was not limited indicating a high thermotolerance of photosynthetic machinery. The results for IND leaves indicate the occurrence of a strategy that merging changes in leaf structure as well as in photosynthetic regulation allow C. salvifolius to maintain an elevated carbon gain in response to temperature increase.     

Quantitative Analysis of the Structure of Etiolated Barley Leaf Using Stereological Methods

Anatomical and morphological characteristics of the first leafblade of barley (Hordeum vulgare L. cv. Korál) were studiedin light-grown plants and compared with those grown in darkness.Etiolated leaves had a higher length/width ratio and a higherproportion of intercellular spaces than light-grown leaves.There were also differences in the distribution of stomata inthe leaf. However, the leaf area and volume, as well as otherstomatal and mesophyll characteristics did not differ significantlybetween the two variants. This supports the hypothesis thatthe first leaf anatomy depends primarily on internal factorsof plant development.     

Aboveground biomass allocation patterns within Mediterranean sub-shrubs: A quantitative analysis of seasonal dimorphism

The monthly patterns of aboveground biomass allocation were studied in the branches of six Mediterranean sub-shrubs with different leaf phenology. Four of them were seasonally dimorphic species, and the remaining two were a winter deciduous and a cushion plant with photosynthetic stems. By the analysis of these species we aimed to identify different aboveground biomass allocation patterns within seasonally dimorphic species and to understand the role of seasonal dimorphism as a strategy to avoid the main stresses of mediterranean climate: summer drought and winter cold. The biomass allocation to the different living and photosynthetic fractions of 3-year-old branches was studied monthly for a minimum of 13 months per species. Leaf area (LA, mm²) and leaf mass per area (LMA, mg cm⁻²) measurements were used to characterize the diverse types of leaves of each species. Standing dead and senescent tissues accounted for a great percentage of the branch biomass of seasonally dimorphic species both during summer and winter. Different patterns of photosynthetic biomass allocation were found within the seasonally dimorphic species analysed. These patterns ranged from the moderate photosynthetic biomass oscillation of Salvia lavandulifolia to the almost deciduousness of Lepidium subulatum, and they were achieved by keeping alive, drying out or shedding different types of branches and leaves throughout the year. The formation of stress tolerant leaves and the reduction in the amount of photosynthetic biomass responded both to the occurrence of summer drought and winter cold. These results demonstrate that seasonal dimorphism is a flexible ecological strategy, as it comprises very different leaf phenologies and enables plants to escape both summer drought and winter cold.     

Effects of relative humidity and temperature stress on pollen viability of Cistus incanus and Myrtus communis

Cistus incanus and Myrtus communis are two species commonly found in Mediterranean shrublands, flowering respectively in spring and summer. Pollen of single flowers of C. incanus is available for dispersal for 8-10 hours, during which time viability decreases from 91% to 61%. Field measurements of the variation in the percent pollen viability during the anthesis of single flowers of M. communis shows that, immediately after anthesis, pollen viability is very high; by the end of the first day it significantly declines and 36 hours after anthesis almost all grains are unviable. Experiments carried out under controlled environment showed that, in C. incanus, the greatest loss of viability occurs at 100% humidity and high temperatures, while under dry conditions pollen viability remains high throughout a wide temperature range. Similarly, pollen viability of M. communis remains high for several hours at high temperature and dry conditions, while it rapidly decreases at high humidity. In conclusion, the temperature experiments indicate that in C. incanus and M. communis, pollen viability does not appear to be drastically reduced if the relative humidity of the environment is low. Therefore, humidity is a far more important factor determining pollen viability loss in both species.     

Interspecific Differences of Leaf Gas Exchange and Water Relations of Three Evergreen Mediterranean Shrub Species

Leaf gas exchange and plant water relations of three co-occurring evergreen Mediterranean shrubs species, Quercus ilex L. and Phillyrea latifolia L. (typical evergreen sclerophyllous shrubs) and Cistus incanus L. (a drought semi-deciduous shrub), were investigated in order to evaluate possible differences in their adaptive strategies, in particular with respect to drought stress. C. incanus showed the highest annual rate of net photosynthetic rate (P _N) and stomatal conductance (g _s) decreasing by 67 and 69 %, respectively, in summer. P. latifolia and Q. ilex showed lower annual maximum P _N and g _s, although P _N was less lowered in summer (40 and 37 %, respectively). P. latifolia reached the lowest midday leaf water potential (₁) during the drought period (–3.54±0.36 MPa), 11 % lower than in C. incanus and 19 % lower than in Q. ilex. Leaf relative water content (RWC) showed the same trend as ₁. C. incanus showed the lowest RWC values during the drought period (60 %) while they were never below 76 % in P. latifolia and Q. ilex; moreover C. incanus showed the lowest recovery of ₁ at sunset. Hence the studied species are well adapted to the prevailing environment in Mediterranean climate areas, but they show different adaptive strategies that may be useful for their co-occurrence in the same habitat. However, Q. ilex and P. latifolia by their water use strategy seem to be less sensitive to drought stress than C. incanus.     

Temporal and Spatial Development of the Cells of the Expanding First Leaf of Arabidopsis thaliana (L.) Heynh

The three-dimensional quantitative leaf anatomy in developingyoung (9–22 d) first leaves of wild type Arabidopsis thalianacv. Landsberg erecta from mitosis through cell and leaf expansionto the cessation of lamina growth has been studied. The domainsof cell division, the relative proportion of the cell typespresent during development and the production of intercellularspace in the developing leaf have been determined by image analysisof entire leaves sectioned in three planes. Mitotic activityoccurs throughout the youngest leaves prior to unfolding andcell expansion is initiated firstly at the leaf tip with a persistentzone of mitotic cells at the leaf base resulting in a gradientof development along the leaf axis, which persists in the olderleaves. Major anatomical changes which occur during the developmentare, a rapid increase in mesophyll volume, an increase in thevein network, and expansion of the intercellular spaces. Thepattern of cell expansion results in a 10-fold variation inmesophyll cell size in mature leaves. In the youngest leavesthe plan area of mesophyll cells varies between 100 µm²and 400 µm² whereas in mature leaves mesophyll cells rangein plan area from 800 µm² to 9500 µm². The volumesof mesophyll tissue and airspace under unit leaf area increase3-fold and 35-fold, respectively, during leaf expansion. Thevolume proportions of tissue types mesophyll:airspace:epiderrnal:vascularin the mature leaf are 61:26:12:1, respectively. This studyprovides comparative information for future identification andanalysis of leaf development mutants of Arabidopsis thaliana. Key words: Arabidopsis, quantitative leaf anatomy, leaf expansion, image analysis     

Morphological,chorological and ecological plasticity of Cistus incanus in the southern Balkans

Abstract

Two subspecies of Cistus incanus L. occurring in the southern Balkans were studied: C. incanus L. ssp. incanus and C. incanus ssp. creticus (L.) Heywood. After studying the morphological differences, the communities dominated by these two subspecies were sampled according to the central European method. The localities of the relevés show the distribution pattern of the two subspecies. The typical subspecies can be found in the area of distribution of thermophilous deciduous forests of Carpinion orientalis, whereas the subspecies creticus grows in the area of the evergreen broad‐leaved forests of Quercetea ilicis. The communities were classified as Diantho–Cistetum incani Micevski et Matevski ex ?arni, Matevski et ?ilc ass. nova hoc loco and Calicotomo villosae–Cistetum cretici Oberdorfer 1954. Analyses of traits (life span, life form, proportion of certain families, bioindicator values and chorotypes) reveal that the harshest site conditions prevail within Diantho–Cistetum incani, which appears as an initial stage of the vegetation succession. Calicotome–Cistetum cretici is a subsequent stage of succession, since the extreme degradation of vegetation on these sites results in ephemeral communities dominated by Poa bulbosa assigned to Romulion.     

Leaf and stem physiological responses to summer and winter extremes of woody species across temperate ecosystems

Winter cold limits temperate plant performance, as does summer water stress in drought‐prone ecosystems. The relative impact of seasonal extremes on plant performance has received considerable attention for individual systems. An integrated study compiling the existing literature was needed to identify overall trends. First, we conducted a meta‐analysis of the impacts of summer and winter on ecophysiology for three woody plant functional types (winter deciduous angiosperms, evergreen angiosperms and conifers), including data for 210 records from 75 studies of ecosystems with and without summer drought across the temperate zone. Second, we tested predictions by conducting a case study in a drought‐prone Mediterranean ecosystem subject to winter freezing. As indicators of physiological response of leaves and xylem to seasonal stress, we focused on stomatal conductance (g_s), percent loss of stem xylem hydraulic conductivity (PLC) and photochemical efficiency of photosystem II (F_v/F_m). Our meta‐analysis showed that in ecosystems without summer drought, g_s was higher during summer than winter. By contrast, in drought‐prone ecosystems many species maintained open stomata during winter, with potential strong consequences for plant carbon gain over the year. Further, PLC tended to increase and F_v/F_m to decrease from summer to winter for most functional types and ecosystems due to low temperatures. Overall, deciduous angiosperms were most sensitive to climatic stress. Leaf gas exchange and stem xylem hydraulics showed a coordinated seasonal response at ecosystems without summer drought. In our Mediterranean site subjected to winter freezing the species showed similar responses to those typically found for ecosystems without summer drought. We conclude that winter stress is most extreme for systems without summer drought and systems with summer drought and winter freezing, and less extreme for drought‐prone systems without freezing. In all cases the evergreen species show less pronounced seasonal responses in both leaves and stems than deciduous species.     

Hydrogen peroxide is involved in the acclimation of the Mediterranean shrub, Cistus albidus L., to summer drought

This study evaluated the possible role of hydrogen peroxide(H₂O₂) in the acclimation of a Mediterranean shrub, Cistus albidusL., to summer drought growing under Mediterranean field conditions.For this purpose, changes in H₂O₂ concentrations and localizationthroughout a year were analysed. H₂O₂ changes in response toenvironmental conditions in parallel with changes in abscisicacid (ABA) and oxidative stress markers, together with ligninaccumulation, xylem and sclerenchyma differentiation, and leafarea were also investigated. During the summer drought, leafH₂O₂ concentrations increased 11-fold, reaching values of 10µmol g^–1 dry weight (DW). This increase occurredmainly in mesophyll cell walls, xylem vessels, and sclerenchymacells in the differentiation stage. An increase in ABA levelspreceded that of H₂O₂, but both peaked at the same time in conditionsof prolonged stress. C. albidus plants tolerated high concentrationsof H₂O₂ because of its localization in the apoplast of mesophyllcells, xylem vessels, and in differentiating sclerenchyma cells.The increase in ABA, and consequently of H₂O₂, in plants subjectedto drought stress might induce a 3.5-fold increase in ascorbicacid (AA), which maintained and even decreased its oxidativestatus, thus protecting plants from oxidative damage. Afterrecovery from drought following late-summer and autumn rainfall,a decrease in ABA, H₂O₂, and AA to their basal levels (60 pmolg^–1 DW, 1 µmol g^–1 DW, and 20 µmol g^–1DW) was observed. Key words: Abscisic acid, ascorbate, ascorbate oxidative status, Cistus albidus, hydrogen peroxide, leaf plasticity, lignin, Mediterranean shrubs, oxidative markers, summer drought Received 29 July 2008; Revised 15 September 2008 Accepted 8 October 2008     

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.     

Co-ordination between xylem anatomy,plant architecture and leaf functional traits in response to abiotic and biotic drivers in a nurse cushion plant

Background and AimsPlants in dry Mediterranean mountains experience a double climatic stress: at low elevations, high temperatures coincide with water shortage during summer, while at high elevations temperature decreases and water availability increases. Cushion plants often act as nurses by improving the microclimate underneath their canopies, hosting beneficiary species that may reciprocally modify their benefactors’ microenvironment. We assess how the nurse cushion plant Arenaria tetraquetra subsp. amabilis adjusts its hydraulic system to face these complex abiotic and biotic constraints.MethodsWe evaluated intra-specific variation and co-ordination of stem xylem anatomy, leaf functional traits and plant architecture in response to elevation, aspect and the presence of beneficiary species in four A. tetraquetra subsp. amabilis populations in the Sierra Nevada mountains, southern Spain.Key ResultsXylem anatomical and plant architectural traits were the most responsive to environmental conditions, showing the highest mutual co-ordination. Cushions were more compact and had smaller, more isolated conductive vessels in the southern than in the northern aspect, which allow minimization of the negative impacts of more intense drought. Only vessel size, leaf mass per area and terminal branch length varied with elevation. Nurse cushions co-ordinated plant architecture and xylem traits, having higher canopy compactness, fewer leaves per branch and fewer, more isolated vessels than non-nurse cushions, which reflects the negative effects of beneficiary plants on nurse water status. In non-nurse cushions, plant architecture co-ordinated with leaf traits instead. The interacting effects of aspect and elevation on xylem traits showed that stress due to frost at high elevation constrained xylem anatomy in the north, whereas stress due to drought had a parallel effect in the south.ConclusionsTrait co-ordination was weaker under more demanding environmental conditions, which agrees with the hypothesis that trait independence allows plants to better optimize different functions, probably entailing higher adjustment potential against future environmental changes.     

Effects of Drought on Water Relations, Mineral Uptake, Water-soluble Carbohydrate Accumulation and Survival of Two Contrasting Populations of Cocksfoot (Dactylis glomerata L.)

The role physiological responses in survival of prolonged soilmoisture deficit was investigated in vegetative plants of twoDactylis glomerata populations: KM2 (drought-resistant) andLutetia (susceptible). The plants were grown in 1 m-deep soilcolumns in a controlled environment. After 56 d of full irrigation,water was withheld for 80 d, by which time all soil moisturehad been consumed; the plants were then rewatered for a further19 d. As drought progressed, leaf extension decreased to zero,water status declined, and water-soluble carbohydrates (WSC)at first increased and then decreased. The most pronounced differencesbetween the two populations was that all KM2 tillers survivedthe drought, but 34% of Lutetia tillers died. In comparisonwith Lutetia, KM2 was characterized by (a) slower shoot growthrate, (b) greater root density at depth, (c) maintenance ofhigher lamina relative water content, (d) greater osmotic adjustmentin leaf bases, (e) higher concentration of WSC in tiller bases,(f) greater ability to export WSC out of dying leaves, (g) lowercontent of metal ions but improved maintenance of P status,and (h) lower proline:amino acid ratio. The contribution ofthese responses to tiller survival under severe drought in controlledenvironments is contrasted with performance and persistenceof swards in the field in the harsher Mediterranean environment.Copyright1995, 1999 Academic Press Dactylis glomerata L., cocksfoot, water-stress, osmotic asjustment, drought survival, root-growth, water-soluble carbohydrates, phosphate, proline, variety comparison     

Seasonal dimorphism and winter chilling stress in Thymus sibthorpii

Seasonal dimorphism (summer/winter) has been so far studied only in a few plants and has been focused on summer drought stress. However, Thymus sibthorpii in the study area appears to be affected by winter chilling stress and not by summer drought stress. Thus, the winter leaves were thicker and more compact compared to the summer leaves and they had more stomata and peltate hairs, more sclerenchymatous fibers, vacuoles with phenolics, and chloroplasts than the summer leaves. In addition, their chloroplasts possessed large grana and starch grains. In the summer leaves, cell vacuoles in mesophyll did not contain phenolics, and chloroplasts were devoid of starch grains and had large plastoglobuli. Physiological measurements revealed higher net photosynthetic rate and chlorophyll content in the winter leaves than in the summer leaves. Proline and soluble sugar content along with antioxidative enzyme (superoxide dismutase, peroxidase, ascorbate peroxidase, glutathione reductase) activities were increased in the winter leaves.     

Trichomes and photosynthetic pigment composition changes: responses of <Emphasis Type="Italic">Quercus ilex </Emphasis>subsp. <Emphasis Type="Italic">ballota</Emphasis> (Desf.) Samp. and<Emphasis Type="Italic"> Quercus coccifera</Emphasis> L. to Mediterranean stress conditions

Sun and shade leaves of two Mediterranean Quercus species, Quercus ilex subsp. ballota (Desf.) Samp. and Quercus coccifera L., were compared by measuring leaf optical properties, photosynthetic pigment composition and photosystem II efficiency. The presence of trichomes in the adaxial (upper) leaf surface of Q. ilex subsp. ballota seems to constitute an important morphological mechanism that allows this species to maintain a good photosystem II efficiency during the summer. Q. coccifera has almost no trichomes and seems instead to develop other physiological responses, including a smaller light-harvesting antenna size, higher concentrations of violaxanthin cycle pigments and a higher (zeaxanthin + antheraxanthin)/(violaxanthin + antheraxanthin + zeaxanthin) ratio. Q. coccifera was not able to maintain a good photosystem II efficiency up to the end of the summer. In Q. ilex subsp. ballota leaves, natural loss or mechanical removal of adaxial-face leaf trichomes induced short-term decreases in photosystem II efficiency. These changes were accompanied by de-epoxidation of violaxanthin cycle pigments, suggesting that the absence of trichomes would trigger physiological responses in this species. Our data have revealed different patterns of response of Q. ilex subsp. ballota and Q. coccifera facing the stress conditions prevailing in the Mediterranean area.