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Influence of weather on cork-ring width.

Ring-width series of cork from Quercus suber L. trees growing at two sites in Extremadura (southwestern Spain) were analyzed in relation to monthly precipitation and temperature, and to climatic indices combining both variables. Ring width of cork showed strong positive correlations with precipitation, especially during the fall and winter. Moderately low temperatures were favorable for cork growth, except in winter and during the onset of phellogen activity. We conclude that drought or temperature, or both, can limit cork growth during the annual drought period.

Rain↗

Energy dissipation in drought-avoiding and drought-tolerant tree species at midday during the Mediterranean summer.

Photosynthetic performance was monitored during two consecutive summers in four co-occurring evergreen Mediterranean tree species growing on a south-facing rocky slope. In response to midday water stress, the drought-avoiding species Pinus halepensis Mill. exhibited marked stomatal closure (g(s)) but no changes in stem water potential (Psi(s)), whereas the drought-tolerant species Quercus coccifera L., Q. ilex ssp. ballota (Desf.) Samp. and Juniperus phoenicea L. displayed declines in midday g(s) and Psi(s). The higher resistance to CO(2) influx in needles of P. halepensis compared with the other species did not result in either a proportional increase in non-radiative dissipation of excess energy or photo-inactivation of photosystem II (PSII). No significant differences were found among species either in the de-epoxidation state of the xanthophyll cycle (DPS) or in the pool of its components on a total chlorophyll basis (VAZ). Despite contrasting midday assimilation rates, the three drought-tolerant species all exhibited a pronounced drop in photochemical efficiency at midday that was characterized by a decrease in the excitation capture efficiency of the open PSII centers. Although photoinhibition was not fully reversed before dawn, it apparently did not result in cumulative photo-damage. Thus, the drought-avoiding and drought-tolerant species employed different mechanisms for coping with excess light during the midday depression in photosynthesis that involved contrasting midday photochemical efficiencies of PSII and different degrees of dynamic photoinhibition as a photo-protective mechanism. These behaviors may be related to the different mechanisms employed by drought-avoiding and drought-tolerant species to withstand water deficit.

Journal Article↗

Seasonal and yearly variations in light use and nitrogen use by seedlings of four deciduous broad-leaved tree species invading larch plantations.

Several deciduous broad-leaved tree species, differing in leaf phenology, invade larch (Larix kaempferii (Lamb.) Carrière) plantations in Japan. The understory light environment of larch forests changes drastically between the leafy and leafless periods. To determine how the invading seedlings exploit the changing light environment, and if phenological differences reflect the light- and nitrogen-use traits of the seedlings, we measured leaf phenology, seasonal changes in light-saturated photosynthetic rate (P(sat)), leaf nitrogen (N) content (N(area)), chlorophyll/nitrogen ratio (Chl/N), specific leaf area (SLA) and N remobilization rate (NRMR) over 3 years. The mid-successional or gap-phase species, Magnolia hypoleuca Siebold & Zucc., had a short leafy period and high P(sat) and NRMR. In contrast, two late-successional tree species, Prunus ssiori Friedr. Schmidt, which undergoes leaf flush before larch, and Carpinus cordata Blume, which maintains green leaves until frost, both had low P(sat) and NRMR but exploited the opportunity for growth during the period when the larch canopy trees were leafless. Quercus mongolica Fisch. ex Ledeb. var. crispula (Blume) Ohashi, a mid-late-successional species that underwent leaf flush at the same time as the overstory larch, had values of photosynthetic parameters between those of the gap-phase and late-successional species. Among species, M. hypoleuca and Q. mongolica had higher photosynthetic rates and photosynthetic N-use efficiencies. In all species, the relationship between N(area) and P(sat) showed species-specific yearly fluctuations; however, there was no yearly fluctuation in the relationship between N(area) and P(sat) at CO2 saturation. Yearly fluctuations in the N(area)-P(sat) relationship appeared to be induced by changes in SLA and N-use characteristics, which in turn are affected by climatic variations.

Chlorophyll↗

Soil respiration in a mixed temperate forest and its contribution to total ecosystem respiration.

Soil respiration (SR) was measured with an infrared gas analyzer in nine plots representative of the heterogeneous vegetation in a mixed coniferous-deciduous forest in the Belgian Campine region. Selected plots included the two most representative overstory species (Pinus sylvestris L. and Quercus robur L.) in combination with the most representative understory species of the forest. A model that includes temperature and water as the main controlling variables was fitted to the data. We found large spatial variability in SR among plots, with typically lower fluxes under the coniferous overstory than under the deciduous overstory (means of 4.8 +/- 0.4 and 8.8 +/- 0.5 Mg C ha(-1) year(-1), respectively). Total annual soil carbon (C) emissions were estimated by weighting fluxes from different types of vegetation according to their relative contribution to the footprint area of the eddy covariance flux measurement. The relative contribution of the two main tree species to the footprint-weighted total SR varied among seasons with the more abundant coniferous overstory contributing the most to total SR during most of the year. Nonetheless, during summer, the contribution of deciduous plots to total SR was disproportionally high because of the more pronounced seasonality of belowground metabolic activity. Net ecosystem carbon dioxide exchange was measured by eddy covariance, and we estimated total ecosystem respiration (TER) with footprint-constrained nighttime fluxes. Mean total annual SR and TER were 6.1 +/- 0.11 and 9.1 +/- 1.15 Mg C ha(-1) year(-1), respectively. The 95% confidence interval of the ratio of annual SR:TER ranged from 0.58 to 0.76, with a mean of 0.67. The contribution of SR to TER tended to vary seasonally, with minimum contributions during summer (less than 50% of TER) and maximum contributions during winter (about 94% of TER).

Carbon Dioxide↗

A semi-automated data processing system for root growth analysis: application to a growing oak seedling.

A data acquisition and processing system for analyzing root structure and growth is presented. Two-dimensional drawings are made of root systems growing against the inclined, transparent wall of a rhizotron. For analysis, a root is treated as a number of straight segments between points determined by the position of root apices at successive times of observation and the insertion points of lateral roots. The coordinates of the points are digitized by means of a graphics tablet and the description of the root system completed with information about root branching pattern and root order. From the recorded data, various root characteristics and growth parameters can be computed with a program available from the authors. Use of the data processing system in studies of root architecture is discussed and its application in the analysis of root growth of a pedunculate oak seedling (Quercus robur L.) reported.

Journal Article↗

Comparative water relations of three successional hardwood species in central Wisconsin.

Water relations of co-occurring understory saplings of Quercus ellipsoidalis E.J. Hill, an early successional, xeric species, Populus tremuloides Michx., an early successional, mesic species, and Acer rubrum L., a late successional species that occurs on both wet and dry sites, were evaluated on four dates during the 1986 growing season. The understory was characterized by high soil water content, low irradiance and low vapor pressure deficit throughout the growing season. Stomatal conductance and calculated transpiration flux were lowest for A. rubrum and highest for P. tremuloides and Q. ellipsoidalis. Except early in the growing season, leaf water potentials were lower in P. tremuloides than in the other species. Populus tremuloides had the highest bulk modulus of elasticity, Q. ellipsoidalis the lowest. Over the growing season, Populus tremuloides and Q. ellipsoidalis, but not A. rubrum, exhibited a decrease in osmotic potential at both full and zero turgor. Of the three species, Populus tremuloides exhibited the sharpest decrease in leaf water potential and turgor pressure with decreasing relative water content.

Journal Article↗

Auxin and ethylene regulation of diameter growth in trees.

Recent studies on the phytohormonal regulation of seasonal cell-division activity in the cambium, primary-wall radial expansion of cambial derivatives, differentiation of xylem cells, and growth of the cortex in forest trees of the north temperate zone are reviewed. Indol-3-ylacetic acid (IAA, auxin) has been characterized by combined gas chromatography-mass spectrometry (GC-MS) in the cambial region of Abies balsamea, Pinus densiflora, Pinus sylvestris and Quercus robur. All of the evidence supports the hypothesis that developing leaves and extending shoots are primary sources of IAA. The rate of ethylene emanation varies among conifer species when adjoining phloem and cambial tissues are incubated in vitro. The cambium from young cuttings of Abies balsamea produces more ethylene than that from older cuttings. Ethylene production by seven-year-old Abies balsamea cambium is substantially increased in vitro when the tissue is provided with exogenous 1-aminocyclopropane-1-carboxylic acid and IAA. In response to elevated ethylene concentrations, cortex growth is accelerated in both hardwood and conifer seedlings. Ethrel (2-chloroethylphosphonic acid) increases ray size and ray-cell number and promotes traumatic resin-canal development in xylem. In Ulmus americana, endogenous ethylene concentrations are inversely correlated with cambial activity. Ethylene decreases vessel diameter in Acer negundo, Acer platanoides and Ulmus americana. Several studies suggest that ethylene has a role in regulating reaction-wood formation in both conifers and hardwoods.

Journal Article↗

Solar equivalent leaf area: an efficient biometrical parameter of individual leaves, trees and stands.

The solar equivalent leaf area (A(s)), a simply and easily determined biometrical parameter of leaves, trees and stands, was derived theoretically. The parameter is defined as projected leaf area weighted for the time integral of irradiance at a given location in the canopy relative to that of fully irradiated leaves at the top of the canopy. The efficiency of A(s) as a basis for estimating stand-area transpiration of a mature oak (Quercus robur L.) forest from measurements of transpiration by individual trees was compared with that of other stand and tree characteristics. Stand transpiration estimates based on A(s) were more precise and less prone to systematic error than estimates based on basal area, timber volume, projected tree crown area, projected leaf area, or leaf dry mass. Solar equivalent leaf area reflects both the amount and the physiological properties of leaves and can be used as a measure of tree size and functional capacity. It can be calculated from ordinary forest inventory data on trees and stands, adjusted according to simple phyllometric data. It appears to have wide application in ecological and forestry studies for relating the physiological characteristics of individual leaves to those of entire trees or stands.

Journal Article↗

Influence of explant source on in vitro axillary shoot formation in oak seedlings.

In vitro shoot development was slower for apical shoot explants of young oak (Quercus robur L.) seedlings growing on Woody Plant Medium containing activated charcoal than for nodal shoot explants. The rate of in vitro shoot development was slowest in explants taken from seedlings that were undergoing rapid shoot elongation and most rapid in explants taken from seedlings that had stopped elongating and had fully expanded leaves. Maximum rooting was achieved on half-strength Woody Plant Medium containing activated charcoal. Rooting ability was not influenced by explant source.

Journal Article↗

Interactive effects of resource availabilities and defoliation on photosynthesis, growth, and mortality of red oak seedlings.

Responses of forest trees to defoliation by insects such as gypsy moth vary greatly from site to site and from individual to individual. To determine whether some of this variation could be explained by variation in other stress factors, red oak (Quercus rubra L.) seedlings were exposed to low and high light, water, mineral nutrient, and defoliation treatments, in a complete factorial design in a greenhouse. Significant interactions were observed among factors for photosynthesis, growth, and mortality, indicating that the response to defoliation was influenced by other stresses. Defoliation increased the photosynthetic capacity per unit leaf area of seedlings grown in the low-water, but not in the high-water, regime. In response to defoliation, growth of seedlings in a low-mineral-nutrient, or low-light, regime was depressed less than that of seedlings grown in a high-mineral-nutrient, or high-light, regime. However, defoliation resulted in a similar percent reduction in biomass in all seedlings in both the high and the low light, water, and mineral nutrient treatments. Defoliation-induced mortality of shaded plants was twice that of plants grown in full sun.

Journal Article↗

Distribution and metabolism of current photosynthate by single-flush northern red oak seedlings.

Carbon distribution and metabolism by northern red oak seedlings (Quercus rubra L.) were followed for 72 h after a 30-min photosynthetic exposure to (14)CO(2). Approximately 50% of the assimilated carbon was lost during this time, presumably through respiration. Most of the (14)C recovered in the plant remained in the source leaf. Most of the exported (14)C was found in stems and especially roots. Position of the source leaf did not affect distribution of translocated (14)C; however, upper source leaves retained slightly more (14)C than lower source leaves. Most (14)C in all tissues was found initially in sugars. Labeled carbon in this fraction decreased rapidly and increased in other chemical fractions, particularly residue (structural carbohydrates and lignin). More (14)C was incorporated into residue than into any other chemical fraction, indicating continued growth by each of the tissues during the lag stage of seedling development. Labeled carbon increased in proteins for 6 to 12 h after treatment, then remained essentially constant for the remainder of the transport period, indicating both rapid incorporation of the pulsed (14)C into, and slow turnover of, the labeled protein components. In source leaves, (14)C incorporated into starch in the light was lost during the dark period, a typical diurnal storage pattern. In stems, (14)C in starch increased for 12 h, then decreased during the dark period, and then increased during the rest of the transport period. This pattern indicates that stems have both diurnal and long-term storage pools of starch. In roots, (14)C in starch increased rapidly for 12 h, then remained constant for the rest of the transport period, indicating primarily long-term storage in this tissue. Peak (14)C in amino acids in taproot preceded that in stem tissue, indicating recycling of (14)C from the roots to the shoot in amino acids.

Journal Article↗

Radiocesium contamination in a submediterranean semi-natural ecosystem following the Chernobyl accident: measurements and models.

Radiocesium dynamics in a Quercus conferta Kit ecosystem in Northern Greece have been extensively studied over the years 1993-1995. Radiocesium distribution in the different parts of the ecosystem was measured. A total 137Cs inventory of 243+/-66 MBq ha(-1) due to the Chernobyl accident was measured in all parts of the ecosystem. Almost 90% of this inventory is still in the upper layers of the soil and the forest floor. In particular 13.4% is in the forest floor, 52.6% in the Ah horizon, and 23.4% in the upper 5 cm of the soil. Only 2.2% of this inventory is in the above ground biomass. The mean total 137Cs deposited on the forest floor from the above ground biomass is 0.18 MBq ha(-1) y(-1). Cesium leaching from the forest floor is negligible. The radiocesium distribution in soil is fixed and in equilibrium, at least since 1993. Most of radiocesium is not available for migration. Cesium migration in soil was modeled by a) an "equivalent diffusion" model with different initial conditions and b) a "compartment" model derived from a diffusion-advection model. A compartment model for the contamination of living biomass is proposed. The total absorbed dose rate in air as well as the contribution due to 137Cs from the Chernobyl accident was determined inside the forest, by in-situ gamma spectrometry.

Biomass↗

Contribution of 137Cs to the total absorbed gamma dose rate in air in a Greek forest ecosystem: measurements and Monte Carlo computations.

The absorbed gamma dose rate in air 1 m above soil due to natural gamma emitters and 137Cs from the Chernobyl accident was determined inside a Quercus conferta Kit ecosystem in Northern Greece by combination of Monte Carlo simulations with the MCNP code and in-situ gamma spectrometry measurements. The total absorbed gamma dose rate in air is about 64 nGy h(-1), where 40% of this value is due to 137Cs and 60% to natural gamma emitters. The Monte Carlo simulations indicated that the gamma absorbed dose rate in air due to 137Cs is mainly due (70%) to unscattered radiation and to a lesser extent (30%) to the scattered radiation. The results obtained with the Monte Carlo simulations for the unscattered radiation were in very good agreement with the experimental values deduced by in-situ gamma spectrometry measurements. From the combination of the Monte Carlo simulations and in-situ gamma spectrometry measurements a conversion factor C = 1 nGy h(-1)/kBq m(-2) was deduced for 137Cs. This factor must be used with caution and only for forest sites similar to the one used for this work.

Air Pollution, Radioactive↗

Warmer springs disrupt the synchrony of oak and winter moth phenology.

Spring temperatures have increased over the past 25 years, to which a wide variety of organisms have responded. The outstanding question is whether these responses match the temperature-induced shift of the selection pressures acting on these organisms. Organisms have evolved response mechanisms that are only adaptive given the existing relationship between the cues organisms use and the selection pressures acting on them. Global warming may disrupt ecosystem interactions because it alters these relationships and micro-evolution may be slow in tracking these changes. In particular, such shifts have serious consequences for ecosystem functioning for the tight multitrophic interactions involved in the timing of reproduction and growth. We determined the response of winter moth (Operophtera brumata) egg hatching and oak (Quercus robur) bud burst to temperature, a system with strong selection on synchronization. We show that there has been poor synchrony in recent warm springs, which is due to an increase in spring temperatures without a decrease in the incidence of freezing spells in winter. This is a clear warning that such changes in temperature patterns may affect ecosystem interactions more strongly than changes in mean temperature.

Animals↗

Dimethylallyl diphosphate and geranyl diphosphate pools of plant species characterized by different isoprenoid emissions.

Dimethylallyl diphosphate (DMADP) and geranyl diphosphate (GDP) are the last precursors of isoprene and monoterpenes emitted by leaves, respectively. DMADP and GDP pools were measured in leaves of plants emitting isoprene (Populus alba), monoterpenes (Quercus ilex and Mentha piperita), or nonemitting isoprenoids (Prunus persica). Detectable pools were found in all plant species, but P. persica showed the lowest pool size, which indicates a limitation of the whole pathway leading to isoprenoid biosynthesis in nonemitting species. The pools of DMADP and GDP of nonemitting, isoprene-emitting, and monoterpene-emitting species were partially labeled (generally 40%-60% of total carbon-incorporated (13)C) within the same time by which volatile isoprenoids are fully labeled (15 min). This indicates the coexistence of two pools for both precursors, the rapidly labeled pool presumably occurring in chloroplasts and thereby synthesized by the methylerythritol phosphate pathway and the nonlabeled pool presumably located in the cytosol and synthesized by the mevalonic pathway. In M. piperita storing monoterpenes in specialized leaf structures, the GDP pool remained totally unlabeled, indicating either that monoterpenes are totally formed by the mevalonic pathway or that labeling occurs slowly in comparison to the large pool of stored monoterpenes in this plant. The pools of DMADP and GDP increased during the season (from May to July) but decreased when the leaf was darkened or exposed to very high temperature. In the dark, the pool of DMADP of the isoprene-emitting species decreased faster than the pool of GDP. However, after 6 h of darkness, both pools were depleted to about 10% of the pool size in illuminated leaves. This indicates that both the chloroplastic and the cytosolic pools of precursors are depleted in the dark. When comparing measurements over the season and at different temperatures, an inverse correlation was observed between isoprene emission by P. alba and the DMADP pool size and between monoterpene emission by Q. ilex and the GDP pool size. This suggests that the pool size does not limit the emission of isoprenoids. Rather, it indicates that the flux of volatile isoprenoids effectively controls the size of their pools of precursors.

Diphosphates↗

Responses of Antioxidative Systems to Drought Stress in Pendunculate Oak and Maritime Pine as Modulated by Elevated CO2.

The aim of the present study was to investigate the effects of an enhanced CO2 concentration alone or in combination with drought stress on antioxidative systems of a deciduous (oak; Quercus robur) and an evergreen (pine; Pinus pinaster) tree species. The seedlings were grown for one season in a greenhouse in tunnels supplied with 350 or 700 [mu]L L-1 CO2. The experiment was repeated in a second year. Antioxidants, protective enzymes, soluble protein, and pigments showed considerable fluctuations in different years. Elevated CO2 caused significant reductions in the activities of superoxide dismutases in both oak and pine. The activities of ascorbate peroxidase and catalase were also reduced in most cases. The activities of dehydroascorbate reductase, monodehydroascorbate radical reductase, glutathione reductase, and guaiacol peroxidase were affected little or not at all by elevated CO2. When the trees were subjected to drought stress by withholding water, the activities of antioxidative enzymes decreased in leaves of pine and oak grown at ambient CO2 and increased in plants grown at elevated CO2 concentrations. The present results suggest that growth in elevated CO2 might reduce oxidative stress to which leaf tissues are normally exposed and enhance metabolic flexibility to encounter increased stress by increases in antioxidative capacity.

Journal Article↗

Freezing Characteristics of Rigid Plant Tissues (Development of Cell Tension during Extracellular Freezing).

The freezing characteristics and development of cell tension during extracellular freezing were examined in supercooling stem tissues of riverbank grapes (Vitis riparia) and cold-hardened leaves of live oak (Quercus virginiana) and mountain cranberry (Vaccinium vitis-idaea). Dormant stem xylem and pith tissues of river-bank grapes were resistant to freeze-induced dehydration above the homogeneous nucleation temperature, and they developed cell tension reaching a maximum of 27 MPa. Similarly, extracellular freezing induced cell tension in the leaves of live oak and mountain cranberry. Maximum cell tension in the leaves of live oak was 16.8 MPa and 8.3 MPa in the leaves of mountain cranberry. Following peak tensions in the leaves, a decline in the pressure was observed with progressive freezing. The results suggest that resistance to cell deformation during extracellular freezing due to cell-wall rigidity can lead to reduced cell dehydration and increased cell tension. A relationship to predict freezing behavior in plant tissues based on cell rigidity is presented. Based on cell-water relations and ice nucleation rates, cell-wall rigidity has been shown to effect the freezing characteristics of plant tissues, including freeze-induced dehydration, supercooling, and homogeneous nucleation temperatures.

Journal Article↗

Isoprene Increases Thermotolerance of Isoprene-Emitting Species.

Isoprene-emitting plants lose a large portion of their assimilated C as isoprene. Because isoprene synthesis can be regulated, it has been assumed that isoprene benefits the plant. Since the rate of isoprene emission from leaves is highly responsive to temperature, we hypothesized that isoprene benefits plants by increasing their thermotolerance. We used three methods to measure isopreneinduced thermotolerance in leaves. Each technique assayed thermotolerance under conditions that suppressed endogenous isoprene synthesis. When measured by chlorophyll fluorescence, thermotolerance of kudzu (Pueraria lobata [Willd.] Ohwi.) leaves increased as much as 4[deg]C in very low light. With higher light, isoprene increased thermotolerance of kudzu leaves by as much as 10[deg]C. When measured as the temperature at which photosynthesis declined to zero, thermotolerance increased with added isoprene by 2.5[deg]C. All three measures of thermotolerance were dose dependent. Both fluorescence techniques also showed isoprene-induced thermotolerance in white oak (Quercus alba L.). Thermotolerance was not observed in bean (Phaseolus vulgaris var Linden), a species that does not emit isoprene. None of the experiments was designed to determine the mechanism of thermotolerance, but we theorize that isoprene functions by enhancing hydrophobic interactions in membranes.

Journal Article↗