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A plant type 2 metallothionein (MT) from cork tissue responds to oxidative stress.

Expression of plant metallothionein genes has been reported in a variety of senescing tissues, such as leaves and stems, ripening fruits, and wounded tissues, and has been proposed to function in both metal chaperoning and scavenging of reactive oxygen species. In this work, it is shown that MT is also associated with suberization, after identifying a gene actively transcribed in Quercus suber cork cells as a novel MT. This cDNA, isolated from a phellem cDNA library, encodes a MT that belongs to type 2 plant MTs (QsMT). Expression of the QsMT cDNA in E. coli grown in media supplemented with Zn, Cd, or Cu has yielded recombinant QsMT. Characterization of the respective metal aggregates agrees well with a copper-related biological role, consistent with the capacity of QsMT to restore copper tolerance to a MT-deficient, copper-sensitive yeast mutant. Furthermore, in situ hybridization results demonstrate that RNA expression of QsMT is mainly observed under conditions related to oxidative stress, either endogenous, as found in cork or in actively proliferating tissues, or exogenous, for example, in response to H(2)O(2) or paraquat treatments. The putative role of QsMT in oxidative stress, both as a free radical scavenger via its sulphydryl groups or as a copper chelator is discussed.

Amino Acid Sequence↗

Temperature response of photosynthesis and internal conductance to CO2: results from two independent approaches.

The internal conductance to CO(2) transfer from intercellular spaces to chloroplasts poses a major limitation to photosynthesis, but few studies have investigated its temperature response. The aim of this study was to determine the temperature response of photosynthesis and internal conductance between 10 degrees C and 35 degrees C in seedlings of a deciduous forest tree species, Quercus canariensis. Internal conductance was estimated via simultaneous measurements of gas exchange and chlorophyll fluorescence ("variable J method"). Two of the required parameters, the intercellular photocompensation point (C(i)*) and rate of mitochondrial respiration in the light (R(d)), were estimated by the Laisk method. These were used to calculate the chloroplastic photocompensation point (Gamma*) in a simultaneous equation with g(i). An independent estimate of internal conductance was obtained by a novel curve-fitting method based on the curvature of the initial Rubisco-limited portion of an A/C(i) curve. The temperature responses of the rate of Rubisco carboxylation (V(cmax)) and the RuBP limited rate of electron transport (J(max)) were determined from chloroplastic CO(2) concentrations. The rate of net photosynthesis peaked at 24 degrees C. C(i)* was similar to reports for other species with a C(i)* of 39 micromol mol(-1) at 25 degrees C and an activation energy of 34 kJ mol(-1). Gamma* was very similar to the published temperature response for Spinacia oleracea from 20 degrees C to 35 degrees C, but was slightly greater at 10 degrees C and 15 degrees C. J(max) peaked at 30 degrees C, whereas V(cmax) did not reach a maximum between 10 degrees C and 35 degrees C. Activation energies were 49 kJ mol(-1) for V(cmax) and 100 kJ mol(-1) for J(max). Both methods showed that internal conductance doubled from 10 degrees C to 20 degrees C, and then was nearly temperature-independent from 20 degrees C to 35 degrees C. Hence, the temperature response of internal conductance could not be fitted to an Arrhenius function. The best fit to estimated g(i) was obtained with a three-parameter log normal function (R(2)=0.98), with a maximum g(i) of 0.19 mol m(-2) s(-1) at 29 degrees C.

Carbon Dioxide↗

Interception loss, throughfall and stemflow chemistry in pine and oak forests in northeastern Mexico.

Interception loss, gross precipitation, throughfall and stemflow solution chemistry beneath pine (Pinus pseudostrobus Lindl.), oak (Quercus sp.) and pine-oak natural forest canopies in northeastern Mexico were measured. Coefficients of variation for throughfall were 12% in pine and oak canopies and 17% in the mixed pine-oak canopy. The variability of stemflow averaged 66, 126 and 73% for pine, oak and the mixed pine-oak canopies, respectively. Linear regression analysis of net versus gross precipitation for the three canopies showed highly significant correlations (r = 0.974-0.984). Total precipitation during the experimental period was 974 mm and estimated interception loss was 19.2, 13.6 and 23% for the pine, oak and pine-oak canopies, respectively. Stemflow did not occur following rainfall events of less than 4 mm and, in all canopies, stemflow represented a minimal proportion of gross precipitation (0.60, 0.50 and 0.03% for pine, oak and pine-oak, respectively). Throughfall pH in pine (6.2), oak (6.3) and pine-oak (6.3) canopies was significantly more acidic than gross precipitation (6.6). Stemflow pH ranged from 3.7 (pine) to 6.0 (oak). The pine-oak canopy registered the highest throughfall and stemflow electrical conductivities, 104 and 188 microS cm(-1), respectively. Net nutrient leaching of K, Mg, Na, Fe, Mn and Zn was significantly higher from the pine-oak canopy than from the pure pine and oak canopies. Mean depositions of Ca and Cu in throughfall behaved similarly among the three types of canopies. A greater proportion of Zn in gross precipitation was absorbed by the oak canopy than by the pine and pine-oak canopies. Enrichment factors beneath the pine-oak canopy relative to gross precipitation varied from 1.2 to 3.2 for macro-nutrients (Ca, K, Mg and Na) and from 1.4 to 3.1 for micro-nutrients (Cu, Fe, Mn and Zn). Stemflow depositions of Ca, K, Mg and Cu were higher in the pine-oak canopy, whereas stemflow depositions of Na, Fe, Mn and Zn were higher in the pine canopy.

Mexico↗

Within-tree variation in transpiration in isolated evergreen oak trees: evidence in support of the pipe model theory.

Within-tree variation in sap flow density (SFD) was measured in two isolated evergreen oak (Quercus ilex L.) trees growing in an oak savannah (dehesa) in southwest Spain. Sap flow was estimated by the constant heating method. Three sensors were installed in the trunk of each tree in three orientations: northeast (NE), northwest (NW) and south (S). Sap flow density was monitored continuously from May 18 to September 27, 1993. Daily values of SFD ranged between 500 and 4500 mm3 mm-2 day-1. There were significant differences in SFD between orientations; SFD was higher in the NE and NW orientations than in the S orientation. These differences were noted on both a daily and seasonal time scale, and were less pronounced on cloudy days and at the end of the drought period, when SFD was relatively low. Our results support the idea that branches of trees can be viewed as a collection of small independent plants.

Immunosuppressive Agents↗

Photosynthetic responses to water deficit in six Mediterranean sclerophyll species: possible factors explaining the declining distribution of Rhamnus ludovici-salvatoris, an endemic Balearic species.

We sought to explain the declining distribution in the Balearic Islands of the endemic shrub Rhamnus ludovici-salvatoris R. Chodat, by comparing its photosynthetic response to drought with that of several widely distributed, competing Mediterranean species (R. alaternus L., Quercus ilex L., Pistacia lentiscus L., Q. humilis Mill. and P. terebinthus L.). All of the study species, except for the two Rhamnus species, avoided desiccation by rapidly adjusting their stomatal conductance at the onset of drought, and maintaining constant leaf relative water content. The two Rhamnus species showed desiccation-tolerant behavior; i.e., as drought progressed, their predawn leaf relative water content decreased simultaneously with stomatal closure. All four desiccation-avoiding species showed a significant positive correlation between leaf thermal dissipation (estimated by the fluorescence parameter NPQ (non-photochemical quenching)) and the de-epoxidation state of the xanthophyll cycle (DPS). The two Rhamnus species exhibited maximum DPS regardless of treatment, but only R. alaternus increased NPQ in response to drought. Rhamnus ludovici-salvatoris had a high ratio of photorespiration to photosynthesis and a low intrinsic water-use efficiency; traits that are likely to be unfavorable for plant productivity under arid conditions. It also had the lowest DPS and thermal dissipation among the six species. We conclude that the photosynthetic traits of R. ludovici-salvatoris account for its limited ability to compete with other species in the Mediterranean region.

Carbon Dioxide↗

Contrasting distribution and seasonal dynamics of carbohydrate reserves in stem wood of adult ring-porous sessile oak and diffuse-porous beech trees.

We tested the hypothesis that broad-leaved forest species with contrasting wood anatomy and hydraulic system (ring-porous versus diffuse-porous) also differ in distribution and seasonal dynamics of carbohydrate reserves in stem wood. Total nonstructural carbohydrate (TNC) reserves (starch and sugars) were measured enzymatically in the 10 youngest stem xylem rings of adult oak (Quercus petraea (Matt.) Liebl.) and beech (Fagus sylvatica L.) trees during an annual cycle. Radial distribution of carbohydrates was investigated according to ring age. On all dates, oak trees had twofold higher TNC concentration than beech trees (41 versus 23 mg g(DM)(-1)), with starch accounting for the high TNC concentration in oak. Seasonal dynamics of TNC concentration were significantly (P < 0.05) more pronounced in oak (20-64 mg TNC g(DM)(-1)) than in beech (17-34 mg TNC g(DM)(-1)). A marked decrease in TNC concentration was observed in oak trees during bud burst and early wood growth, whereas seasonal fluctuations in TNC concentrations in beech trees were small. The radial distribution of TNC based on ring age differed between species: TNC was restricted to the sapwood rings in oak, whereas in beech, it was distributed throughout the wood from the outermost sapwood ring to the pith. Although the high TNC concentrations in the outermost rings accounted for most of the observed seasonal pattern, all of the 10 youngest xylem rings analyzed participated in the seasonal dynamics of TNC in beech trees. The innermost sapwood rings of oak trees had low TNC concentrations. Stem growth and accumulation of carbon reserves occurred concomitantly during the first part of the season, when there was no soil water deficit. When soil water content was depleted, stem growth ceased in both species, whereas TNC accumulation was negligibly affected and continued until leaf fall. The contrasting dynamics and distribution of carbohydrate reserves in oak and beech are discussed with reference to differences in phenology, early spring growth and hydraulic properties between ring-porous trees and diffuse-porous trees.

Carbohydrate Metabolism↗

Solute accumulation of chestnut oak and dogwood leaves in response to throughfall manipulation of an upland oak forest.

To determine the biochemical basis of osmotic adjustment, seasonal and treatment differences in foliar water- soluble organic solutes and inorganic ions were investigated for two hardwood species that exhibited osmotic adjustment in a Throughfall Displacement Experiment at the Walker Branch Watershed near Oak Ridge, Tennessee. Leaf samples of overstory and understory chestnut oak (Quercus prinus L.) and understory dogwood (Cornus florida L.) were collected during the 1994 (wet) and 1995 (dry) growing seasons from each of three treatments: dry (-33% throughfall), ambient (control) and wet (+33% throughfall). Foliar soluble carbohydrates and organic acids were measured by gas chromatography-mass spectrometry. During May 1994, when the demand for sucrose was greatest, dogwood accumulated small amounts of glucose, quinic acid and Mg2+, offsetting a decline in nitrate concentration. As the mild drought continued and tree growth slowed, there was a significant accumulation of sucrose in dogwood in the dry treatment in June, and a trend toward increased K+. In overstory chestnut oak in the dry treatment over the same period, there were significant accumulations of fructose, glucose and K+, and a trend toward increased quinic acid accumulation. Sucrose did not become a key osmotically active compound in chestnut oak until August 1994. In 1995, with the exception of understory chestnut oak, there was no accumulation of K+ in either species. During the severe drought of 1995, monosaccharides, particularly glucose and fructose, accounted for most of the osmotic adjustment in both species. Among solutes, glucose constituted the largest accumulation in dogwood in the dry treatment in August 1995, followed by fructose and sucrose. There was only a moderate increase in solutes in chestnut oak trees in 1995, with fructose and glucose constituting over 50% of the predicted solute accumulation in July. Both species accumulated a wider array of solutes during the dry year than during the wet year, but treatment differences in solute accumulation in chestnut oak were partially limited by drought. The greater dehydration tolerance of chestnut oak than dogwood was evident in the higher baseline concentrations of organic solutes and the greater array of solutes accumulated in response to drought.

Adaptation, Physiological↗

Rate of stomatal opening, shoot hydraulic conductance and photosynthetic characteristics in relation to leaf abscisic acid concentration in six temperate deciduous trees.

Correlations between leaf abscisic acid concentration ([ABA]), stomatal conductance (gs), rate of stomatal opening in response to an increase in leaf water potential (si), shoot hydraulic conductance (L) and photosynthetic characteristics were examined in saplings of six temperate deciduous tree species: Acer platanoides L., Padus avium Mill., Populus tremula L., Quercus robur L., Salix caprea L. and Tilia cordata Mill. Species-specific values of foliar [ABA] were negatively related to the mean values of gs, si, L and light- and CO2- saturated net photosynthesis (P(max)), thus providing strong correlative evidence of a scaling of foliar gas exchange and hydraulic characteristics with leaf endogenous [ABA]. In addition, we suggest that mean gs, si, L and Pmax for mature leaves may partly be determined by the species-specific [ABA] during leaf growth. The most drought-intolerant species had the lowest [ABA] and the highest gs, suggesting that interspecific differences in [ABA] may be linked to differences in species-specific water-use efficiency. Application of high concentrations of exogenous ABA led to large decreases in gs, si and P(max), further underscoring the direct role of ABA in regulating stomatal opening and photosynthetic rate. Exogenous ABA also decreased L, but the decreases were considerably smaller than the decreases in gs, si and Pmax. Thus, exogenous ABA predominantly affected the stomata directly, but modification of L by ABA may also be an important mechanism of ABA action. We conclude that interspecific variability in endogenous [ABA] during foliage growth and in mature leaves provides an important factor explaining observed differences in L, gs, si and Pmax among temperate deciduous tree species.

Abscisic Acid↗

Modeling effects of hydrological changes on the carbon and nitrogen balance of oak in floodplains.

We extended the applicability of the ecosystem model BIOME-BGC to floodplain ecosystems to study effects of hydrological changes on Quercus robur L. stands. The extended model assesses floodplain peculiarities, i.e., seasonal flooding and water infiltration from the groundwater table. Our interest was the tradeoff between (a). maintaining regional applicability with respect to available model input information, (b). incorporating the necessary mechanistic detail and (c). keeping the computational effort at an acceptable level. An evaluation based on observed transpiration, timber volume, soil carbon and soil nitrogen content showed that the extended model produced unbiased results. We also investigated the impact of hydrological changes on our oak stands as a result of the completion of an artificial canal network in 1971, which has stopped regular springtime flooding. A comparison of the 11 years before versus the 11 years after 1971 demonstrated that the hydrological changes affected mainly the annual variation across years in leaf area index (LAI) and soil carbon and nitrogen sequestration, leading to stagnation of carbon and nitrogen stocks, but to an increase in the variance across years. However, carbon sequestration to timber was unaffected and exhibited no significant change in cross-year variation. Finally, we investigated how drawdown of the water table, a general problem in the region, affects modeled ecosystem behavior. We found a further amplification of cross-year LAI fluctuations, but the variance in soil carbon and nitrogen stocks decreased. Volume increment was unaffected, suggesting a stabilization of the ecosystem two decades after implementation of water management measures.

Carbon↗

Sap flow of three co-occurring Mediterranean woody species under varying atmospheric and soil water conditions.

We studied the seasonal patterns of water use in three woody species co-occurring in a holm oak forest in northeastern Spain. The three species studied, Quercus ilex L., Phillyrea latifolia L. and Arbutus unedo L., constitute more than 99% of the total basal area of the forest. The study period included the dry seasons of 1999 and 2000. Water use was estimated with Granier-type sap flux sensors. Standard meteorological variables, soil water content and leaf water potentials were also monitored. All monitored individuals reduced leaf-related sap flow (Q(l)) during the summer, concurrent with an increase in soil moisture deficit (SMD). Despite similar maximum Q(l) between species, the decline in Q(l) with increasing SMD was species-dependent. The average reduction in Q(l) between early summer and the peak of the drought was 74% for A. unedo (n = 3), 58% for P. latifolia (n = 3) and 87% for Q. ilex (n = 1). The relationship between canopy stomatal conductance (G(s)) and vapor pressure deficit (D) changed during the course of the drought, with progressively lower G(s) for any given D. Summertime reductions of Q(l) and G(s) were associated with between-species differences in vulnerability to xylem embolism, and with the corresponding degree of native embolism (lowest in P. latifolia and highest in Q. ilex). Our results, combined with previous studies in the same area, outlined differences among the species studied in manner of responding to water shortage, with P. latifolia able to maintain water transport at much lower water potentials than the other two species. In an accompanying experiment, A. unedo responded to an experimental reduction in water availability by reducing Q(l) during the summer. This species also modified its water use between years according to the different seasonal patterns of precipitation. These results are discussed in relation to the possible impacts that climate change will have on Q. ilex-dominated forests.

Ericaceae↗

Stomatal responses to drought at a Mediterranean site: a comparative study of co-occurring woody species differing in leaf longevity.

We studied stomatal responses to decreasing predawn water potential (Psipd) and increasing leaf-to-air water vapor pressure difference (VPD) of co-occurring woody Mediterranean species with contrasting leaf habits and growth form. The species included two evergreen oaks (Quercus ilex subsp. ballota (Desf.) Samp. and Q. suber L.), two deciduous oaks (Q. faginea Lam. and Q. pyrenaica Willd.) and two deciduous shrubs (Pyrus bourgaeana Decne. and Crataegus monogyna Jacq.). Our main objective was to determine if stomatal sensitivity is related to differences in leaf life span and leaf habit. The deciduous shrubs had the least conservative water-use characteristics, with relatively high stomatal conductance and low stomatal sensitivity to soil and atmospheric drought. As a result, Psipd decreased greatly in both species during the growing season, resulting in early leaf abscission in the summer. The deciduous oaks showed intermediate water-use characteristics, having maximum stomatal conductances and CO2 assimilation rates similar to or even higher than those of the deciduous shrubs. However, they had greater stomatal sensitivity to soil drying and showed less negative Psipd values than the deciduous shrubs. The evergreen oaks, and especially the species with the greatest leaf longevity, Q. ilex, exhibited the most conservative water-use behavior, having lower maximum stomatal conductances and greater sensitivity to VPD than the deciduous species. As a result, Psipd decreased less during the growing season in the evergreens than in the deciduous species, which may contribute to greater leaf longevity by avoiding irreversible damage during the summer drought. However, the combination of low maximum CO2 assimilation rates and high stomatal sensitivity to drought must have a negative impact on the final carbon budget of leaves with a long life span.

Crataegus↗

Tracing carbon uptake from a natural CO2 spring into tree rings: an isotope approach.

We analyzed 14C, 13C and 18O isotope variations over a 50-year period in tree rings of Quercus ilex L. trees growing at a natural CO2 spring in a Mediterranean ecosystem. We compared trees from two sites, one with high and one with low exposure to CO2 from the spring. The spring CO2 is free of 14C. Thus, this carbon can be traced in the wood, and the amount originating from the spring calculated. The amount decreased over time, from about 40% in 1950 to 15% at present for the site near the spring, indicating a potential difficulty in the use of natural CO2 springs for elevated CO2 research. The reason for the decrease may be decreasing emission from the spring or changes in stand structure, e.g., growth of the canopy into regions with lower concentrations. We used the 14C-calculated CO2 concentration in the canopy to determine the 13C discrimination of the plants growing under elevated CO2 by calculating the effective canopy air 13C/12C isotopic composition. The trees near the spring showed a 2.5 per thousand larger 13C discrimination than the more distant trees at the beginning of the investigated period, i.e., for the young trees, but this difference gradually disappeared. Higher discrimination under elevated CO2 indicated reduced photosynthetic capacity or increased stomatal conductance. The latter assumption is unlikely as inferred from the 18O data, which were insensitive to CO2 concentration. In conclusion, we found evidence for a downward adjustment of photosynthesis under elevated CO2 in Q. ilex in this dry, nutrient-poor environment.

Carbon Dioxide↗

Thermal optima of photosynthetic functions and thermostability of photochemistry in cork oak seedlings.

Temperature effects on photosynthesis were studied in seedlings of evergreen Mediterranean cork oak (Quercus suber L.). Responses to changes in temperature and the temperature optima of maximal carboxylation rate (V(cmax)) and maximal light-driven electron flux (J(max)) were estimated from gas exchange measurements and a leaf-level photosynthesis model. The estimated temperature optima were approximately 34 and 33 degrees C for V(cmax) and J(max), respectively, which fall within the lower range of temperature optima previously observed in deciduous tree species. The thermostability of the photosynthetic apparatus was estimated according to the temperature at which basal chlorophyll a fluorescence begins to increase (T(c)). The T(c) was highly variable, increasing from 42 to 51 degrees C when ambient temperature rose from 10 to 40 degrees C, and increasing from 44 to 54 degrees C with decreasing soil water availability while net CO(2) assimilation rate dropped to almost zero. When a heat shock was imposed, an additional small increase in T(c) was observed in drought-stressed and control seedlings. Maximal T(c) values following heat shock were about 56 degrees C, which, to our knowledge, are the highest values that have been observed in tree species. In conclusion, the intrinsic temperature responses of cork oak did not differ from those of other species (similar T(c) under ambient temperature and water availability, and relatively low thermal optima for photosynthetic capacity in seedlings grown at cool temperatures). However, the large ability of cork oak to acclimate to drought and elevated temperature may be an important factor in the tolerance of this evergreen Mediterranean species to summer drought and high temperatures.

Dehydration↗

Photosynthate distribution patterns in cherrybark oak seedling sprouts.

We used 14C tracers to determine photosynthate distribution in cherrybark oak (Quercus pagoda Raf.) seedling sprouts following release from competing mid-story vegetation. Fall acquisition of labeled photosynthates by seedlings followed expected source-sink patterns, with root and basal stem tissues serving as the primary sinks. Four months after the seedlings had been labeled with 14C, they were clipped to induce sprouting. First-flush stem and leaf tissues of the resulting seedling sprouts were the primary sinks for labeled photosynthates stored in root tissues. Second-flush stem and leaf tissues, and first-flush stem and leaf tissues the following growing season, were not primary sinks for labeled photosynthates stored in root tissues despite the high radioactivity in root tissues. Root tissues appeared to deposit photosynthates in a layering process whereby the last photosynthates stored in new xylem were the first to be depleted during the initiation of a growth flush the following spring. There were more labeled photosynthates in roots of released seedling sprouts compared with non-released seedling sprouts, indicating increased vigor of released seedling sprouts in response to greater light availability. In contrast, stem and source leaf tissues of non-released seedling sprouts contained greater percentages of labeled photosynthates compared with released seedling sprouts, indicating either greater sink strength or poorly developed xylem and phloem pathways that created inefficiencies in distribution to root tissues. The 14C distribution coefficients confirmed the distribution patterns and provided additional information on the important sinks in released and non-released cherrybark oak seedling sprouts.

Photosynthesis↗

Reliance on stored water increases with tree size in three species in the Pacific Northwest.

In tall old forests, limitations to water transport may limit maximum tree height and reduce photosynthesis and carbon sequestration. We evaluated the degree to which tall trees could potentially compensate for hydraulic limitations to water transport by increased use of water stored in xylem. Using sap flux measurements in three tree species of the Pacific Northwest, we showed that reliance on stored water increases with tree size and estimated that use of stored water increases photosynthesis. For Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), water stored in xylem accounted for 20 to 25% of total daily water use in 60-m trees, whereas stored water comprised 7% of daily water use in 15-m trees. For Oregon white oak (Quercus garryana Dougl. ex Hook.), water stored in xylem accounted for 10 to 23% of total daily water use in 25-m trees, whereas stored water comprised 9 to 13% of daily water use in 10-m trees. For ponderosa pine (Pinus ponderosa Dougl. ex Laws.), water stored in xylem accounted for 4 to 20% of total daily water use in 36-m trees, whereas stored water comprised 2 to 4% of daily water use in 12-m trees. In 60-m Douglas-fir trees, we estimated that use of stored water supported 18% more photosynthesis on a daily basis than would occur if no stored water were used, whereas 15-m Douglas-fir trees gained 10% greater daily photosynthesis from use of stored water. We conclude that water storage plays a significant role in the water and carbon economy of tall trees and old forests.

Northwestern United States↗

Climatic signal of earlywood vessels of oak on a maritime site.

Earlywood vessel lumen areas were measured in 72 consecutive tree rings in wood cores from oak (Quercus robur L.) trees in a maritime woodland. This anatomical time series was statistically correlated with climate data for the same time span. There was a strong dependence of earlywood vessel lumen area on rainfall between February and April, which reflects the role of water availability in vessel ontogeny. By inversion, earlywood vessel lumen areas can be used as a proxy to reconstruct spring precipitation beyond the archived weather records. Such information may be of value in the context of climate change.

Climate↗

Winter photoinhibition in the field involves different processes in four co-occurring Mediterranean tree species.

Photoinhibition was examined in four co-occurring Mediterranean evergreen tree species during two consecutive winters. In response to low temperatures and saturating light, Juniperus phoenicea L., Pinus halepensis Mill., Quercus coccifera L. and Q. ilex ssp. ballota (Desf.) Samp. exhibited marked chronic photoinhibition, indicated by low predawn maximal photochemical efficiency of photosystem II (PSII) (Fv/Fm). Low Fv/Fm values were correlated with high concentrations of xanthophyll cycle components (VAZ) and with the maintenance of high concentrations of zeaxanthin overnight (DPSpd). In all species, however, chronic photoinhibition was enhanced as the winter progressed in the absence of changes in DPSpd, suggesting cumulative damage toward the end of winter. Photoinhibition differed among species: P. halepensis always displayed significantly higher Fv/Fm values; and Q. coccifera had the lowest Fv/Fm values, showing a high sensitivity to the combination of high light and low temperatures. Differences among species were not fully explained by differences in the xanthophyll pool or its de-epoxidation state. Chronic photoinhibition overlapped with a dynamic photoinhibition as shown by the low values of photochemical efficiency of the open reaction centers of PSII at midday. Winter photoprotective strategies differed among species and may involve photoprotective mechanisms in addition to those associated with xanthophylls. The observed species-specific differences matched results obtained for the same species in summer; however, comparison of the two seasons suggests that the higher VAZ concentration observed in winter has an additional structural photoprotective role.

Chlorophyll↗

Environmental controls on sap flow in a northern hardwood forest.

Our objective was to gain a detailed understanding of how photosynthetically active radiation (PAR), vapor pressure deficit (D) and soil water interact to control transpiration in the dominant canopy species of a mixed hardwood forest in northern Lower Michigan. An improved understanding of how these environmental factors affect whole-tree water use in unmanaged ecosystems is necessary in assessing the consequences of climate change on the terrestrial water cycle. We used continuously heated sap flow sensors to measure transpiration in mature trees of four species during two successive drought events. The measurements were scaled to the stand level for comparison with eddy covariance estimates of ecosystem water flux (Fw). Photosynthetically active radiation and D together explained 82% of the daytime hourly variation in plot-level transpiration, and low soil water content generally resulted in increased stomatal sensitivity to increasing D. There were also species-specific responses to drought. Quercus rubra L. showed low water use during both dry and wet conditions, and during periods of high D. Among the study species, Acer rubrum L. showed the greatest degree of stomatal closure in response to low soil water availability. Moderate increases in stomatal sensitivity to D during dry periods were observed in Populus grandidentata Michx. and Betula papyrifera Marsh. Sap flow scaled to the plot level and Fw demonstrated similar temporal patterns of water loss suggesting that the mechanisms controlling sap flow of an individual tree also control ecosystem evapotranspiration. However, the absolute magnitude of scaled sap flow estimates was consistently lower than Fw. We conclude that species-specific responses to PAR, D and soil water content are key elements to understanding current and future water fluxes in this ecosystem.

Acer↗