Contribution of physiological and morphological adjustments to drought resistance in two Mediterranean tree species

Plant water potential (ψ), its components, and gas exchange data of two Mediterranean co-occurring woody species (Quercus ilex L. and Phillyrea latifolia L.) were measured in response to seasonal changes in water availability over two consecutive years. The relative contribution of physiological and morphological adjustments to drought resistance was assessed through Principal Component Analyses. There were large adjustments in stomatal conductance (∼36 % of accounted variance). Net photosynthetic rate and water use efficiency were closely tuned to water availability and accounted for ∼17 % of variance. The slope of the water potential vs. relative water content (dψ/dRWC₀) below zero pressure potential increased as a result of seasonal and ontogenic increases in apoplastic water fraction and accounted for ∼20 % variance. This tolerance mechanism was accompanied by an increased range of positive pressure potential, suggesting a functional role of sclerophylly in these Mediterranean evergreens. Similarly, changes in the slope of dψ/dRWC in the range of positive pressure potential (∼13 % of accounted variance) were associated to variations in cell wall elasticity and resulted in lower RWC at zero pressure potential. When considering the species studied separately, the results indicated the primary role of stomatal regulation in the drought resistance of Qilex, while increased apoplastic water fraction had a major contribution in the drought resistance of P. latifolia. This research was supported by Spanish CICYT grants CLI99-0479 and REN-2002-00633. L.S. acknowledges the financial support from Ministerio de Ciencia y Tecnologia (“Ramon y Cajal” program, Spain). An erratum to this article is available at .     

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.     

Tree growth,mortality, and above-ground biomass accumulation in a holm oak forest under a five-year experimental field drought

A holm oak forest was exposed to an experimental drought during 5 years to elucidate the growth responses of the dominant species Quercus ilex, Arbutus unedo and Phillyrea latifolia. Soil water availability was partially reduced, about 15% as predicted for this area for the next decades by GCM and ecophysiological models, by plastic strips intercepting rainfall and by ditch exclusion of water runoff. The stem diameter increment was highly correlated with annual rainfall in all species, and drought treatment strongly reduced the diameter increment of Q. ilex (41%) and specially of A. unedo (63%), the species showing higher growth rates. Stem mortality rates were highly correlated with previous stem density, but drought treatment increased mortality rates in all species. Q. ilex showed the highest mortality rates (9% and 18% in control and drought plots, respectively), and P. latifolia experienced the lowest mortality rates (1% and 3% in control and drought plots, respectively). Drought strongly reduced the increment of live aboveground biomass during these 5 years (83%). A. unedo and Q. ilex experienced a high reduction in biomass increment by drought, whereas P. latifolia biomass increment was insensitive to drought. The different sensitivity to drought of the dominant species of the holm oak forest may be very important determining their future development and distribution in a drier environment as expected in Mediterranean areas for the next decades. These drier conditions could thus have strong effects on structure (species composition) and functioning (carbon uptake and biomass accumulation) of these Mediterranean forests.     

Adaptive photosynthetic strategies of the Mediterranean maquis species according to their origin

In consideration of their origin the adaptive strategies of the evergreen species of the Mediterranean maquis were analysed. Rosmarinus officinalis L., Erica arborea L., and Erica multiflora L. had the lowest net photosynthetic rate (P_N) in the favourable period [7.8±0.6 mol(CO₂) m^–2s^–1, mean value], the highest P_N decrease (on an average 86 % of the maximum) but the highest recovery capacity (>70 % of the maximum) at the first rainfall in September. Cistus incanus L. and Arbutus unedo L. had the highest P_N during the favourable period [15.5±5.2 mol(CO₂) m^–2s^–1, mean value], 79 % decrease during drought, and a lower recovery capacity (on an average 54 %). Quercus ilex L., Phillyrea latifolia L., and Pistacia lentiscus L. had an intermediate P_N in the favourable period [9.2±1.3 mol(CO2) m^–2s^–1, mean value], a lower reduction during drought (on an average 63 %), and a range from 62 % (Q. ilex and P. latifolia) to 39 % (P. lentiscus) of recovery capacity. The Mediterranean species had higher decrease in P_N and stomatal conductance during drought and a higher recovery capacity than the pre-Mediterranean species. Among the pre-Mediterranean species, P. latifoliahad the best adaptation to long drought periods also by its higher leaf mass per area (LMA) which lowered leaf temperature thus decreasing transpiration rate during drought. Moreover, its leaf longevity determined a more stable leaf biomass during the year. Among the Mediteranean species, R. officinalis was the best adapted species to short drought periods by its ability to rapidly recover. Nevertheless, R. officinalis had the lowest tolerance to high temperatures by its P_N dropping below half its maximum value when leaf temperature was over 33.6°C. R. officinalismay be used as a bioindicator species of global change.This revised version was published online in March 2005 with corrections to the page numbers.     

Ecology of vegetative regeneration after coppicing in macchia stands in central Italy

Vegetative regeneration after coppicing of Quercus ilex, Phillyrea latifolia, and Fraxinus ornus in two macchia stands in the Maremma Nature Park (Grosseto, Italy) has been studied.Observations during the first three years after the cut showed that horizontal space occupation by shoots is rapid (95% of the available space being reoccupied within 3 yrs.) and stool mortality very low. We observed differences among species in resprouting, growth pattern and shoot differentiation. Q. ilex and P. latifolia had a longer budburst period than F. ornus. There were no considerable differences in height growth among the three species in the first year. During the second and third years the height increments decreased. F. ornus average height increments were higher than those of Q. ilex and P. latifolia. These two species produced proleptic flushes from the first year on, while F. ornus only showed spring flush. The examination of proleptic shoots and of summer and autumn rainfall seem to show that the latter control Q. ilex growth and shoot differentiation more than that of other species. Differentiation among shoots of the same stool occurs from the first year onwards and the initial vigour differences amongst shoots remain during the following years. Resprouting, space occupation and shoot differentiation have been discussed in relation to rainfall and temperature.The high stool survival and the strong link between stool vigour parameters before and after coppicing confirm a certain stability in successional dynamics following a single coppicing event.Abbreviations CA Crown Projection Area - DBH Diameter at Breast Height (1.30 m) - F. ornus Fraxinus ornus - G Basal area at Breast Height (1.30 m) - HD Height of the dominant shoot per stool - ND Number of the dominant shoots per stool - P. latifolia Phillyrea latifolia - Q. ilex Quercus ilex     

Establishment of co‐existing Mediterranean tree species under a varying soil moisture regime

We investigated the responses of two co‐existing Mediterranean trees with different regeneration strategies (Phillyrea latifolia seedlings and Quercus ilex sprouts) to experimental drought below the forest canopy. We considered different recruitment stages and used leaf isotopic discrimination to estimate water use efficiency (WUE) and nitrogen availability and use. Drought decreased the emergence and survival of seedlings and sprouts. Survival and growth of older saplings were not influenced by drought. Seedling emergence of P. latifolia was higher than Q. ilex sprout production, but Q. ilex sprouts had higher survival and growth rates. These differences disappeared in the sapling stage. Carbon isotopic discrimination suggested that Q. ilex sprouts had higher WUE than P. latifolia seedlings. Drought increased WUE of recruits, particularly in Q. ilex. Water use regulation increased with ageing, particularly in P. latifolia. Q. ilex had higher δ¹⁵N values than P. latifolia; these were also higher under drier soil conditions. Current year seedlings had higher δ¹⁵N than saplings, particularly in P. latifolia, suggesting they exploit superficial soil layers. These results suggest that sprouts obtain benefit from resources stored in parent plants. At earlier stages, they perform better than seedlings. This response is not coupled to adult vulnerability to drought for these species, revealing the difficulty of predicting species dynamics during climate change.     

Chlorophyll fluorescence responses to temperature and water availability in two co-dominant Mediterranean shrub and tree species in a long-term field experiment simulating climate change

A rain exclusion experiment simulating drought conditions expected in Mediterranean areas for the following decades (15% decrease in soil moisture) is being conducted since 1999 in a Mediterranean holm oak forest to study its response to the forecasted climatic changes for the coming decades. The maximum PSII quantum yield of primary photochemistry (Fv/Fm) was measured in Quercus ilex, and Phillyrea latifolia, the co-dominant species of the studied forest, from 1999 to 2009 in four plots: two of them were control plots and the other two plots received the rain exclusion treatment. In both species, the Fv/Fm values were highly dependent on air temperatures, and in a second term, in water availability. P. latifolia was the species with the larger decrease in Fv/Fm values induced by low air temperatures, while in hot seasons, the Fv/Fm values in P. latifolia were even higher than in Q. ilex. Rainfall exclusion decrease Fv/Fm values significantly only in few monitoring dates. The most drought resistant species P. latifolia was more affected by the experimental rainfall exclusion than Q. ilex that instead lost number of leaves per tree. There was a synergic effect of drought stress and winter cold in P. latifolia not observed in Q. ilex, but a more conservative strategy in P. latifolia maintaining leaves with a down-regulation of the linear photosynthetic electron transport. These results indicate that, although other physiological and reproductive strategies at whole plant level must be also taken into account, the warmer and drier environment expected for the following decades could favour the species more sensitive to cold and more resistant to drought, the shrub P. latifolia, in detriment of the tree Q. ilex as already observed in the field after severe heat-drought episodes.     

Rehydration effects on pressure-volume relationships in four temperate woody species: variability with site,time of season and drought conditions

Summary Seasonal pressure-volume (P-V) analyses were conducted on rehydrated and non-rehydrated leaves of Quercus rubra, Q. ilicifolia, Q. prinus, and Fraxinus americana in central Pennsylvania, U.S.A., to test the hypothesis that rehydration-induced shifts in P-V parameters occur in woody species from a non-arid region, and that the magnitude of these shifts increases with species drought tolerance and drought conditions. The species from a xeric ridge (Q. ilicifolia and Q. prinus) displayed increases of about 0.4–0.6 MPa in the osmotic potentials at full and zero turgor and a concurrent loss of symplastic solutes following 12 h and 24 h rehydration, particularly during a late-season drought. In contrast, the mesic, valley species (Q. rubra and F. americana) did not display significant shifts in osmotic parameters with rehydration at any time. In several instances, the relative water content at zero turgor (RWC₀) increased by about 6% (e.g., from 85% to 91%) and the bulk elastic modulus () decreased by about 4.0 MPa following rehydration and correction for the plateau effect; the magnitude of these shifts was greatest in the xeric species. However, when data were not corrected for the plateau effect, RWC₀ decreased by about 4% in some of the species/date combinations. Plateaus were also responsible for some of the decrease in with rehydration, but not for the shifts in osmotic potentials. The largest increases in osmotic potentials corresponded with decreases in tissue osmotic solute content. Rehydration-induced shifts in P-V parameters were responsible for masking or reducing most of the species and seasonal differences exhibited in nonrehydrated samples.     

Δ13C and tree-ring width reflect different drought responses in Quercus ilex and Pinus halepensis

Holm oak (Quercus ilex L.) and Aleppo pine (Pinus halepensis Mill) are representative of two different functional types of trees extensively found in the Mediterranean: evergreen sclerophyllous and drought-adapted conifers. The former is considered a partially drought-tolerant species, whereas the latter is a typically drought-avoiding, water-saving species. We postulated that contrasting strategies in response to water deficits in Q. ilex and P. halepensis would lead to a differential sensitivity to changes in water availability. To test this hypothesis, we compared the response of both species in growth rate (measured as radial increments) and intrinsic water use efficiency [WUE_i, as inferred from carbon isotope discrimination (¹³C) in wood samples] among sites from different provenance regions in NE Spain. We found significant differences in ¹³C and growth among provenance regions, partly explained by contrasting water availability. Wood ¹³C was positively related with precipitation and the ratio between precipitation and potential evapotranspiration (P / E). However, these relationships were stronger in P. halepensis (for P / E, r ²=0.67, P <0.001) than in Q. ilex (r ²=0.42, P <0.01). In addition, radial growth was positively related with precipitation and ¹³C in P. halepensis (r ²=0.32 and r ²=0.35, respectively, P <0.01), but not in Q. ilex. We concluded that P. halepensis was more sensitive than Q. ilex to water availability, showing faster increase in WUE_i in response to water stress. We also found that the effect of north/south aspect on ¹³C and growth was site-specific, and unrelated to climatic variables.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Comparative water relations of four Mediterranean oak species

The water relations and responses of two evergreen (Quercus ilex L. and Q. suber L.) and two deciduous (S. afares Pomel. and Q. faginea Will.) Quercus species were studied under experimental conditions. Two-year old seedlings grown in 30 l. pots were subjected to a drying period during which stomatal conductance, pre-dawn potential and minimum foliar potential were measured.The results shows that, for all species, the daily course of stomatal conductance agrees with the patterns proposed by Hinckley et al. (1978 & 1983). Concurrent with the species responses to short-term variation in water availability, it was found that pre-dawn leaf water potential controlled the maximum daily leaf conductance. There was a strong correlation between pre-dawn leaf potential and maximum daily conductance as described by the reciprocal function g_{srmax for}=(-0.47+2.61._p)^-1 the evergreen oaks and g_{srmax for}=(-1.94+7.39._p)^-1 for the deciduous species. These differences between the two groups may partialy explain their geograhic distributions, and suggest general questions concerning the mechanisms which optimize water-use efficiency in Mediterranean oak species.     

Seedling performance in sewage sludge-amended degraded Mediterranean woodlands

Biosolids have been widely used for land reclamation, but information on their use in restoration, i.e., on less degraded areas, is scarce. Biosolids may be used to restore forest ecosystems by fostering tree establishment in degraded shrublands. Detailed knowledge on the effects of biosolid application is needed to optimize such practice. We evaluated the effect of different rates (0, 7.5 and 14.5 kg dry weight per plant) and types of biosolid application on the performance of Pinus halepensis and Quercus ilex seedlings, and operational costs. Biosolids increased seedling mortality in both species, particularly when seedlings were planted in direct contact with them. Mortality mostly occurred during the first year, and was probably favored by soil shrinking and salinity. Foliar and needle nitrogen concentration increased with biosolid rate in the short term, but biosolids affected negatively (P. halepensis), or had no effect (Q. ilex) on phosphorus and potassium concentration. Biosolids had a positive effect on P. halepensis growth, and a negative effect on Q. ilex growth at the highest rate when seedlings were in contact with biosolids. Cost of this type of biosolid application approximately doubled plantation cost, but were similar or cheaper that landfill disposal of biosolids. The lowest application rate showed the best balance between seedling response and costs for P. halepensis, whereas biosolid use cannot be recommended for Q. ilex.     

Changes in leaf δC and δN for three Mediterranean tree species in relation to soil water availability

A rain exclusion experiment simulating drought conditions expected in Mediterranean areas for the following decades (15% decrease in soil moisture) was conducted in a Mediterranean holm oak forest to study the response of leaf δ¹³C, δ¹⁵N, and N concentrations to the predicted climatic changes for the coming decades. Plant material was sampled in 2000, 2003, 2004, and 2005 in eight plots: four of them were control plots and the other four plots received the rain exclusion treatment. Although there was a negative relationship between δ¹³C and soil moisture, for each species and year, the rain exclusion treatment did not have any significant effect on δ¹³C, and therefore on the intrinsic water use efficiency (iWUE) of the three dominant species: Phillyrea latifolia, Arbutus unedo, and Quercus ilex. On the other hand, rain exclusion clearly increased the δ¹⁵N values in the three species studied, probably indicating higher N losses at the soil level leading to a ¹⁵N enrichment of the available N. It suggested that rain exclusion exerted a greater effect on the nitrogen biogeochemical cycle than on the carbon assimilation process. δ¹⁵N values were inversely correlated with summer soil moisture in Q. ilex and A. unedo, but no relationship was observed in P. latifolia. This latter species showed the lowest iWUE values, but it was the only species with no decrease in annual basal increment in response to the rain exclusion treatment, and it also had the highest resistance to the hot and dry conditions projected for the Mediterranean basin in the coming decades. The different strategies to resist rain exclusion conditions of these species could induce changes in their competitive ability and future distribution. The losses of N from the ecosystem may further limit plant growth and ecosystem functioning.     

Growth at reduced turgor: irreversible and reversible cell-wall extension of maize coleoptiles and its implications for the theory of cell growth

The relationship between steady-state elongation rate (G) and turgor pressure (P; G/P curve) was investigated using isolated segments of maize (Zea mays L.) coleoptiles incubated in osmotic solutions of a water potential range of 0 to -10 bar (polyethylene glycol 6000 as osmoticum). Short-term elongation measurements revealed curvilinear G/P curves with a steep slope at high turgor and a shallow slope at low turgor. Owing to a decrease of osmotic pressure and turgor, there was a tendency for straightening of the G/P curves during long-term elongation. An elongation rate of zero was adjusted by lowering the turgor by 4.5 bar at a constant osmotic pressure of 6.7 bar. Auxin increased — whereas abscisic acid decreased — the slope of the G/P curve but these hormones had no effect on the threshold turgor of growth (Y = 2.2 bar). It is concluded that extensibility of the growing cell walls represented by the yielding coefficient of Lockhart's growth equation is turgor-dependent and therefore decreases to a very low value as the turgor approaches Y. When the turgor was kept at Y, a constant segment length was maintained over at least 6 h. However, separation of reversible (l_rev) and irreversible (l_irr) components of total (in vivo) length (l_tot = l_rev + l_irr) W measuring segment length before and after freezing/thawing revealed that l_irr increased continuously and l_rev decreased continuously at constant l_tot. After a step-down in turgor the segments grew in l_irr although they shrank in l_tot over the whole turgor range of 0irr irreversible length - l_rev reversible length - l_tot total length (= l_irr + l_rev) - _i osmotic pressure of cell sap - _i water potential of tissue - _o water potential of incubation medium - ABA abscisic acid - G growth rate - m yielding coefficient - P turgor pressure - PEG polyethylene glycol 6000 - Y yield threshold Supported by Deutsche Forschungsgemeinschaft (SFB 206). We thank R. Hertel for helpful comments.     

Correlation between loss of turgor and accumulation of abscisic acid in detached leaves

Mature leaves of Phaseolus vulgaris L. (red kidney bean), Xanthium strumarium L. (cocklebur), and Gossypium hirsutum L. (cotton) were used to study accumulation of abscisic acid (ABA) during water stress. The water status of individual, detached leaves was monitored while the leaves slowly wilted, and samples were cut from the leaves as they lost water. The leaf sections were incubated at their respecitive water contents to allow ABA to build up or not. At least 8 h were required for a new steady-state level of ABA to be established. The samples from any one leaf covered a range of known water potentials (), osmotic pressures (), and turgor pressures (p). The and p values were calculated from pressure-volume curves, using a pressure bomb to measure the water potentials. Decreasing water potential had little effect on ABA levels in leaves at high turgor. Sensitivity of the production of ABA to changes in progressively increased as turgor approached zero. At p=1 bar, ABA content averaged 4 times the level found in fully turgid samples. Below p=1 bar, ABA content increased sharply to as much as 40 times the level found in unstressed samples. ABA levels rose steeply at different water potentials for different leaves, according to the at which turgor became zero. These differences were caused by the different osmotic pressures of the leaves that were used; must cqual - for turgor to be zero. Leaves vary in , not only among species, but also between plants of one and the same species depending on the growing conditions. A difference of 6 bars (calculated at =0) was found between the osmotic pressures of leaves from two groups of G. hirsutum plants; one group had previously experienced periodic water stress, and the other group had never been stressed. When individual leaves were subsequently wilted, the leaves from stress-conditioned plants required a lower water potential in order to accumulate ABA than did leaves from previously unstressed plants. On the basis of these results we suggest that turgor is the critical parameter of plant water relations which controls ABA production in water-stressed leaves.Abbreviations ABA abscisic acid - me-ABA abscisic-acid methyl ester - leaf water potential - osmotic pressure - p volumeaveraged turgor - volumetric modulus of elasticity     

Responses to water stress of apoplastic water fraction and bulk modulus of elasticity in sunflower (Helianthus annuus L.) genotypes of contrasting capacity for osmotic adjustment

The responses to water stress of the bulk modulus of elasticity () and the apoplastic water fraction were examined using six sunflower cultivars of differing capacity for osmotic adjustment (OA). Water stress did not affect the partitioning of water between apoplastic (ca. 20%) and symplastic fractions in leaves which expanded during the exposure to stress in any genotype. Hence, no genotype-linked effects on either the buffering of cell water status during stress or on the estimates of bulk leaf osmotic potential could be expected. Genotypes differed in the degree of change in (estimated from pressure/volume [P/V] curves) and OA (estimated using both ln RWC/ ln o plots and P/V curves) induced by exposure to stress. In three genotypes increased significantly (p=0.05) as a consequence of stress, in another three change were small. OA was the only attribute of the three examined that could have contributed to turgor maintenance under stress. There was a strong negative association between leaf expansion and degree of OA across genotypes (r=–0.91) and a strong positive one between OA and (r=0.94). However all genotypes evidenced some degree of OA. These results are consistent with part of the genotype differences in OA being attributable to variations in leaf expansion during exposure to stress.     

Dampening effects of long‐term experimental drought on growth and mortality rates of a Holm oak forest

Forests respond to increasing intensities and frequencies of drought by reducing growth and with higher tree mortality rates. Little is known, however, about the long‐term consequences of generally drier conditions and more frequent extreme droughts. A Holm oak forest was exposed to experimental rainfall manipulation for 13 years to study the effect of increasing drought on growth and mortality of the dominant species Quercus ilex, Phillyrea latifolia, and Arbutus unedo. The drought treatment reduced stem growth of A. unedo (?66.5%) and Q. ilex (?17.5%), whereas P. latifolia remained unaffected. Higher stem mortality rates were noticeable in Q. ilex (+42.3%), but not in the other two species. Stem growth was a function of the drought index of early spring in the three species. Stem mortality rates depended on the drought index of winter and spring for Q. ilex and in spring and summer for P. latifolia, but showed no relation to climate in A. unedo. Following a long and intense drought (2005–2006), stem growth of Q. ilex and P. latifolia increased, whereas it decreased in A. unedo. Q. ilex also enhanced its survival after this period. Furthermore, the effect of drought treatment on stem growth in Q. ilex and A. unedo was attenuated as the study progressed. These results highlight the different vulnerabilities of Mediterranean species to more frequent and intense droughts, which may lead to partial species substitution and changes in forest structure and thus in carbon uptake. The response to drought, however, changed over time. Decreased intra‐ and interspecific competition after extreme events with high mortality, together with probable morphological and physiological acclimation to drought during the study period, may, at least in the short term, buffer forests against drier conditions. The long‐term effects of drought consequently deserve more attention, because the ecosystemic responses are unlikely to be stable over time.Nontechnical summaryIn this study, we evaluate the effect of long‐term (13 years) experimental drought on growth and mortality rates of three forest Mediterranean species, and their response to the different intensities and durations of natural drought. We provide evidence for species‐specific responses to drought, what may eventually lead to a partial community shift favoring the more drought‐resistant species. However, we also report a dampening of the treatment effect on the two drought‐sensitive species, which may indicate a potential adaptation to drier conditions at the ecosystem or population level. These results are thus relevant to account for the stabilizing processes that would alter the initial response of ecosystem to drought through changes in plant physiology, morphology, and demography compensation.     

Wall yield threshold and effective turgor in growing bean leaves

The rate of cell enlargement depends on cell-wall extensibility (m) and on the amount of turgor pressure (P) which exceeds the wall yield threshold (Y). The difference (P-Y) is the growth-effective turgor (P _e). Values of P, Y and P _ehave been measured in growing bean (Phaseolus vulgaris L.) leaves with an isopiestic psychrometer, using the stress-relaxation method to derive Y. When rapid leaf growth is initiated by light, P, Y and P _eall decrease. Thereafter, while the growth rate declines in maturing leaves, Y continues to decrease and P _eactually increases. These data confirm earlier results indicating that the changes in light-stimulated leaf growth rate are primarily controlled by changes in m, and not by changes in P _e. Seedlings incubated at 100% relative humidity have increased P, but this treatment does not increase growth rate. In some cases Y changes in parallel with P, so that P _eremains unchanged. These data point out the importance of determining P _e, rather than just P, when relating cell turgor to the growth rate.Abbreviations and symbols FC fusicoccin - m wall extensibility - P turgor pressure - P _e effective turgor - RH relative humidity - Y yield threshold - _w water potential - _s osmotic potential     

Species-specific,seasonal, inter-annual,and historically-accumulated changes in foliar terpene emission rates in <Emphasis Type="Italic">Phillyrea latifolia</Emphasis> and <Emphasis Type="Italic">Quercus ilex</Emphasis> submitted to rain exclusion in the Prades Mountains (Catalonia)

Mediterranean vegetation emits large amounts of terpenes. We aimed to study the effects of the decreases in soil water availability forecast for the next decades by global circulation models and ecophysiological models on the terpene emissions by two widely distributed Mediterranean woody species, Phillyrea latifolia L. and Quercus ilex L. We subjected holm oak forest plots to an experimental soil drought of ca. 20% decrease in soil moisture by partial rainfall exclusion and runoff exclusion. We measured the emission rates throughout the seasons for two years with contrasting precipitation and soil moisture (16.6% average in 2003 vs. 6.4% as average in 2005). Among the detected volatile terpenes, only α-pinene and limonene were present in detectable quantities in all of the studied periods. Total terpene emitted ranged from practically zero (spring 2003) to 3.6 and 58.3 μg/(g dry wt h) (winter 2005 and summer 2003 for P. latifolia and Q. ilex, respectively). A clear seasonality was found in the emission rates (they were the highest in summer in both species) and also in the qualitative composition of the emission mix. Maximum emissions of α-pinene occurred in spring and maximum emissions of limonene in winter. Neither the inter-annual differences in water availability nor the rain exclusion treatment significantly affected the emissions in P. latifolia, but Q. ilex showed by 17% lower emissions during the drier second year of study, 2005, but more than two- and threefold increases with the drought treatment in summer 2003 and in summer 2005, respectively, showing historical accumulated effects. These results, which show increased monoterpene emission under the moderate drought produced by the treatment and decreased emission under the severe second year drought, and a much higher sensitivity to drought in Q. ilex than in P. latifolia, are useful in understanding the behavior of plant volatiles under Mediterranean conditions and in modeling future emission under changing climate conditions. They show that the usage of current models could lead to under- and overestimations of the emission under summer dry conditions, because most current algorithms are based on light and temperature only.     

Acclimation to light in seedlings of <Emphasis Type="Italic">Quercus petraea</Emphasis> (Mattuschka) Liebl. and <Emphasis Type="Italic">Quercus pyrenaica</Emphasis> Willd. planted along a forest-edge gradient

Photosynthetic acclimation of two co-occurring deciduous oaks (Quercus petraea and Quercus pyrenaica) to a natural light gradient was studied during one growing season. In the spring of 2003, 90 seedlings per species were planted along a transect resulting from a dense Pinus sylvestris stand, an adjacent thinned area and a 10-m-wide firebreak (16.5–60.9% Global Site Factor (GSF)). In two dates of the following summer, we measured leaf gas exchange, carboxylation efficiency (CE), chlorophyll and nitrogen content, light–response curves of chlorophyll a fluorescence parameters, and leaf mass per area (LMA). Summer was mild, as evidenced by leaf predawn water potential (Ψ_pd), which reduced the interactive effect of water stress on the response of seedlings to light. Q. pyrenaica had higher LMA, CE, stomatal conductance (g _{s max}) and photosynthesis per unit area than Q. petraea at all growth irradiances. , LMA, g _{s max} and electron transport rate (ETR) all increased with light availability (GSF) in a similar fashion in both species. Light had also a clear effect on the organization of Photosystem II (PS II), as deduced by chlorophyll a fluorescence curves. Chlorophyll concentration (Chl_m) decreased with increasing light availability in Q. pyrenaica but it did not in Q. petraea. Seedlings of Q. petraea acclimated to higher irradiances showed a greater non-photochemical quenching (NPQ) than those of Q. pyrenaica. This suggests a higher susceptibility to high light in Q. petraea, which would be consistent with a better adaptation to shade, inferred from the lower LMA or the lower rate of photosynthesis.     

The effects of phosphorus nutrition and soil pH on the growth of some ancient woodland indicator plants and their interaction with competitor species

The forest under-storey herbs Anemone nemorosa, Lamiastrum galeobdolon and Veronica montana are generally considered indicator species of old, broadleaved woodland sites where the soil fertility is often low. In a glasshouse bioassay, however, all three species not only showed large positive growth responses to supplied P concentrations (0–10mgL ^–1) solutions, but also tolerated high P concentrations (20–40mgL ^–1), well above those normally found in their natural habitat. Plants responded by raising the concentrations of P in their shoot and root tissues and increasing their biomass, resulting in an increased P uptake. A shade-tolerant competitor species, Urtica dioica, also grew vigorously across the full range of P concentrations, restricting the growth of the woodland species. This emphasises the difficulty of establishing semi-natural woodland vegetation in the presence of competitor species, for example in situations where new woodlands are planted on fertile ex-agricultural soils containing large residual concentrations of P. The influence of soil pH on the growth and nutrient relations of A. nemorosa, L. galeobdolon, V. montana, Poa trivialisandU. dioicawas determined in a separate experiment using an ex-arable soil as the growing medium with pH levels adjusted from 7.4 to 5.8 and 4.3 respectively. Acidifying the soil enhanced growth, but reduced the concentrations of N, P and K in the leaves of all three woodland species, probably due to dilution of these minerals in the increased dry matter production. The competitor species (P. trivialis and U. dioica) responded in similar manner to the woodland indicator species. These results suggest that manipulating soil pH as a means of facilitating the establishment of woodland indicator species in new farm woods is unlikely, in the short term, to be effective where competitor species are present.