Phytophthora disease of Quercus ilex in south‐western Spain

Oak decline that was affecting three holm oak sites in the province of Huelva (south‐western Spain) was studied during 1998–1999. The syndromes of dieback and sudden death have been observed and, in both cases, foliar symptoms were associated with root rot. Characterization of the fungal isolates from necrotic roots led us to identify Phytophthora cinnamomi A2 as consistently associated with the disease. The optimum growth temperatures of these isolates were very high (30°C). Inoculation tests under controlled conditions demonstrated the pathogenicity of the isolates on holm and cork oak seedlings. None of the other biotic factors of Mediterranean oak decline that have been previously described were found in the present study and so, in this case, the forest decline model does not seem to be necessary in order to explain the disease observed. The defoliation and mortality of the oaks was primarily caused by P. cinnamomi, although some abiotic factors such as alternating periods of drought and wet weather in the region may play an important role.     

Influence of pruning and the climatic conditions on acorn production in holm oak (Quercus ilex L.) dehesas in SW Spain

Acorn production by Quercus ilex L. ssp. ballota (Desf.) Samp. in SW Spain was assessed, and variations between years and the influence of pruning on it were examined. To this end, an experimental study was conducted at two different sites (Calañas and San Bartolomé, in the province of Huelva) where trees were subjected to traditional (light, moderate or heavy) pruning and also to a new (crown-regeneration) pruning method. Acorn yield was quantified over a period of 5 years in the Calañas plot and 4 in the San Bartolomé plot, and found to average at 95.61 ± 0.76 g DM/m², which is equivalent to 6.5 ± 0.05 kg DM/tree; however, yield figures varied markedly between years depending on the particular climatic conditions. The average acorn production was correlated with the water potential in mid summer (end of July); the annual, spring and autumn rainfall; and the actual evapotranspiration for the period from September (previous year) to August. No significant differences in acorn production between traditional pruning intensities were detected; in fact, there were only hints that heavy pruning might result in decreased acorn yields. The new pruning method used, crown-regeneration, seems promising with a view to increasing acorn yield; however, it should be tested on larger sample sizes before any final conclusions can be drawn in this respect. Based on the results, the present health status of holm oaks in southwestern Spain (a result of sustained decline) and the low value of firewood — which used to be a very important source of income from pruning a few decades ago -, the authors recommend reducing the frequency and intensity of pruning in the dehesas of the study area.     

Bole water content shows little seasonal variation in century-old Douglas-fir trees

Purportedly, large Douglas-fir trees in the American Pacific Northwest use water stored in bole tissues to ameliorate the effects of seasonal summer drought, the water content of bole tissues being drawn down over the summer months and replenished during the winter. Continuous monitoring of bole relative water content (RWC) in two 110-120-year-old Douglas-fir trees with ThetaProbe impedance devices provided an integrated measure of phloem-sapwood water content over 4 years. Seasonal changes in RWC closely tracked cambial activity and wood formation, but lagged changes in soil water content by 2-3 months. The RWC in the combined phloem and sapwood markedly increased during earlywood production in the late spring and early summer to maximum values of 64-77% as plant available soil water (ASW) decreased by approximately 60%. With transition and latewood formation, RWC decreased to minimum values of 59-72%, even as ASW increased in the fall. The difference between minimum RWC in the fall and maximum RWC in midsummer was only approximately 5%. Seasonal changes in bole RWC corresponded to cambial phenology, although decreasing AWS appeared to trigger the shift from earlywood to latewood formation.     

Site-specific water relations and stomatal response of Quercus ilex in a Mediterranean watershed

Intraspecific variations in the water relations and stomatal response of Quercus ilex L. were analyzed under field conditions by comparing trees at two locations within a Mediterranean watershed (l'Avic, Catalonia, NE Spain). Distinct environmental gradients exist between the two sites (referred to as ridge top at 975 m and valley bottom at 700 m) with greater soil depth for water storage, reduced radiation, reduced wind and higher water vapor pressure deficits at the valley bottom than at the ridge top. Osmotic adjustment and changes in tissue elasticiity did not significantly increase drought resistance in the trees studied. The leaf-to-air vapor pressure difference (Deltaw) threshold for inducing stomatal closure was higher at the ridge top (15.6 kPa MPa(-1) +/- 0.5 SE) than at the valley bottom (9.8 kPa MPa(-1) +/- 1.0 SE). However, increases in Deltaw beyond the threshold were followed by greater reductions in leaf conductance of trees at the ridge top than at the valley bottom. At both sites, maximum leaf conductance was related to predawn shoot water potential which, in turn, was related to watershed stream flow. The effects of water deficits during the dry summer of 1989 were more severe in trees at the valley bottom than at the ridge top. During periods of high evaporative demand, site-specific differences in the control of water loss led to more conservative water use by trees at the ridge top and, thus, to even greater drought avoidance (higher predawn water potentials) in late summer.     

Effect of elevated CO2 on monoterpene emission of young Quercus ilex trees and its relation to structural and ecophysiological parameters

We investigated growth, leaf monoterpene emission, gas exchange, leaf structure and leaf chemical composition of 1-year-old Quercus ilex L. seedlings grown in ambient (350 microl l(-1)) and elevated (700 microl l(-1)) CO2 concentrations ([CO2]). Monoterpene emission and gas exchange were determined at constant temperature and irradiance (25 degrees C and 1000 micromol m(-2) s(-1) of photosynthetically active radiation) at an assay [CO2] of 350 or 700 microl l(-1). Measurements were made on intact shoots after the end of the growing season between mid-October and mid-February. On average, plants grown in elevated [CO2] had significantly increased foliage biomass (about 50%). Leaves in the elevated [CO2] treatment were significantly thicker and had significantly higher concentrations of cellulose and lignin and significantly lower concentrations of nitrogen and minerals than leaves in the ambient [CO2] treatment. Leaf dry matter density and leaf concentrations of starch, soluble sugars, lipids and hemi-cellulose were not significantly affected by growth in elevated [CO2]. Monoterpene emissions of seedlings were significantly increased by elevated [CO2] but were insensitive to short-term changes in assay [CO2]. On average, plants grown in elevated [CO2] had 1.8-fold higher monoterpene emissions irrespective of the assay [CO2]. Conversely, assay [CO2] rapidly affected photosynthetic rate, but there was no apparent long-term acclimation of photosynthesis to growth in elevated [CO2]. Regardless of growth [CO2], photosynthetic rates of all plants almost doubled when the assay [CO2] was switched from 350 to 700 microl l(-1). At the same assay [CO2], mean photosynthetic rates of seedlings in the two growth CO2 treatments were similar. The percentage of assimilated carbon lost as monoterpenes was not significantly altered by CO2 enrichment. Leaf emission rates were correlated with leaf thickness, leaf concentrations of cellulose, lignin and nitrogen, and total plant leaf area. In all plants, monoterpene emissions strongly declined during the winter independently of CO2 treatment. The results are discussed in the context of the acquisition and allocation of resources by Q. ilex seedlings and evaluated in terms of emission predictions.     

Diversification of Pinus halepensis forests by sowing Quercus ilex and Quercus pubescens acorns: testing the effects of different vegetation and soil treatments

The goal of this study was to develop management strategies favouring establishment and survival of holm oak (Quercus ilex L.) and downy oak (Quercus pubescens Willd.)??two species co-occurring in Southern France??in mature Allepo pine forests (Pinus halepensis Mill subsp. halepensis). An experimental design was assessed in a partially cut mature stand of Aleppo pine in which five soil and vegetation treatments??chopping, chopping followed by scarification in one or two directions, prescribed burning, control??and two slash treatments (presence/absence) were applied. A total of 1,600 sowing points, each composed of 3 Q. ilex or Q. pubescens acorns, were installed in the different treatments in November of two consecutive years at 6 and 18 months, after the end of treatments. Survival was monitored 3 and 2 years after sowing, soil surface at the sowing points was characterized at different dates, and predawn leaf water potentials were measured during the dry season. High mortality occurred after the first summer, but survival after 1 year was 2.3?C5.2 higher in Q. ilex than in Q. pubescens, confirming that Q. ilex was better adapted to the drier parts of the Mediterranean area. Survival was significantly influenced by the treatments, but there was a variable response between the two sowing years under most of the treatments. Only intense fire proved the most beneficial treatment for seedling survival in both years. The micro-local soil cover conditions induced by the treatments played a major role in explaining oak survival. In particular, grass cover (mainly Brachypodium retusum) proved to be largely unfavourable to seedling survival and growth, and this detrimental effect was also confirmed by lower predawn leaf water potential values with increasing grass cover. Acorn introduction designed to diversify mature Aleppo pine forest after soil and vegetation treatments therefore has to be considered for treatments that most efficiently impair the pre-existing competing grass cover such as prescribed high-intensity fire treatment.     

Calcium mineral nutrition increases the tolerance of Quercus ilex to Phytophthora root disease affecting oak rangeland ecosystems in Spain

Root rot caused by the soil-borne pathogen Phytophthora cinnamomi is leading to significant oak tree mortality in rangeland ecosystems in south western Spain. Susceptibility to P. cinnamomi infections of Q. ilex seedlings with a standard nutrition, deficient in K⁺, and deficient in Ca²⁺, was tested. Oaks deficient in K⁺ showed high values in Ca²⁺ content and were tolerant to the disease. Nutritional deficiency in Ca²⁺, however, did not lead to a higher K⁺ level in plants and induced poorer root development. In addition, K⁺ plant content does not appear to have any effect on pathogen tolerance. Based on these results, we conclude that satisfactory calcium nutrition may confer Holm oaks with a greater tolerance to root disease caused by P. cinnamomi. For this reason, limestone supplements are recommended as a measure against root rot caused by P. cinnamomi in rangelands in southern Spain, as a good option for control of oak root disease.     

Response of Quercus pyrenaica (melojo oak) to soil water deficit: a case study in Spain

In the present study, carried out from 2004 to 2006, leaf and stem water potential and stem water content were measured in Quercus pyrenaica Willd. individuals in an experimental forested catchment located in Central Western Spain under Mediterranean subhumid conditions. These indicators of tree water status were compared with soil moisture contents measured in the same area from 0 to 1 m depth and from 0 to 2.5 m depth during the last year of the study. The objectives were to clarify the seasonal and year-to-year variations in tree water status, to examine applicability of stem water content as useful water stress indicator and to discuss how deep soil water and root uptake contribute to survival during the long dry summer. Seasonal variations in the tree variables measured revealed a typical pattern, with maximum values at the end of spring followed by a progressive decline during the summer drought in response to the decrease in soil water content (almost exhausted at 0–100 cm depth). The relatively high values and the non-significant variation in predawn leaf water potentials (except for 2005, which was exceptionally dry) indicate that no clear water stress situations occurred. This may be explained in terms of a progressive absorption of water from the deeper layers. The results also suggest that the stem water content is a more sensitive indicator of long-term water limitation than the other variables measured.     

Diurnal and seasonal changes in stem increment and water use by yellow poplar trees in response to environmental stress

To evaluate indicators of whole-tree physiological responses to climate stress, we determined seasonal, daily and diurnal patterns of growth and water use in 10 yellow poplar (Liriodendron tulipifera L.) trees in a stand recently released from competition. Precise measurements of stem increment and sap flow made with automated electronic dendrometers and thermal dissipation probes, respectively, indicated close temporal linkages between water use and patterns of stem shrinkage and swelling during daily cycles of water depletion and recharge of extensible outer-stem tissues. These cycles also determined net daily basal area increment. Multivariate regression models based on a 123-day data series showed that daily diameter increments were related negatively to vapor pressure deficit (VPD), but positively to precipitation and temperature. The same model form with slight changes in coefficients yielded coefficients of determination of about 0.62 (0.57-0.66) across data subsets that included widely variable growth rates and VPDs. Model R2 was improved to 0.75 by using 3-day running mean daily growth data. Rapid recovery of stem diameter growth following short-term, diurnal reductions in VPD indicated that water stored in extensible stem tissues was part of a fast recharge system that limited hydration changes in the cambial zone during periods of water stress. There were substantial differences in the seasonal dynamics of growth among individual trees, and analyses indicated that faster-growing trees were more positively affected by precipitation, solar irradiance and temperature and more negatively affected by high VPD than slower-growing trees. There were no negative effects of ozone on daily growth rates in a year of low ozone concentrations.     

Grazing affects shoot growth,nutrient and water status of Quercus ilex L. in Mediterranean open woodlands

Context

Mediterranean open woodlands (dehesa) have faced a dual process of intensification and abandonment of grazing which has resulted in alteration of the understory vegetation.

Aims

We analysed the effects of land use changes on the physiological status of holm oak in different open woodlands (dehesa) in southern Iberian Peninsula.

Methods

In an area of extensive grazing, we selected six paired plots (one grazed, one abandoned) and grouped them by habitat types according to understory composition (nearly all monospecific Cistus ladanifer L. shrub or mixed shrub). Six plots of moderate and heavy grazing intensity were chosen within a settled area of livestock use. Shoot growth, macronutrient concentrations and water content were assessed in samples of holm oak leaves.

Results

Abandonment of grazing affected some nutrient concentrations and water content of holm oak leaves, but the effects were different according to habitat type. C. ladanifer shrub reduced N and P concentrations and water content while mixed shrub increased P concentration and water content. High grazing intensity improved shoot growth and leaf N and Mg concentrations.

Conclusion

Extensive grazing could be a useful management tool to enhance growth, nutritional and water status of holm oak in a habitat with limited resources such as Mediterranean open woodlands (dehesa).     

The current situation and future perspectives of Quercus ilex and Pinus halepensis afforestation on agricultural land in Spain under climate change scenarios

New Forests - Between 1994 and 2017, 137,455 ha of agricultural land were afforested in Andalusia (Spain), using a great diversity of tree species, under the Common Agricultural Policy...     

Regulation of transpirational water loss in Quercus suber trees in a Mediterranean-type ecosystem

Sap flux density in branches, leaf transpiration, stomatal conductance and leaf water potentials were measured in 16-year-old Quercus suber L. trees growing in a plantation in southern Portugal to understand how evergreen Mediterranean trees regulate water loss during summer drought. Leaf specific hydraulic conductance and leaf gas exchange were monitored during the progressive summer drought to establish how changes along the hydraulic pathway influence shoot responses. As soil water became limiting, leaf water potential, stomatal conductance and leaf transpiration declined significantly. Predawn leaf water potential reflected soil water potential measured at 1-m depth in the rhizospheres of most trees. The lowest predawn leaf water potential recorded during this period was -1.8 MPa. Mean maximum stomatal conductance declined from 300 to 50 mmol m(-2) s(-1), reducing transpiration from 6 to 2 mmol m(-2) s(-1). Changes in leaf gas exchange were attributed to reduced soil water availability, increased resistances along the hydraulic pathway and, hence, reduced leaf water supply. There was a strong coupling between changes in soil water content and stomatal conductance as well as between stomatal conductance and leaf specific hydraulic conductance. Despite significant seasonal differences among trees in predawn leaf water potential, stomatal conductance, leaf transpiration and leaf specific hydraulic conductance, there were no differences in midday leaf water potentials. The strong regulation of changes in leaf water potential in Q. suber both diurnally and seasonally is achieved through stomatal closure, which is sensitive to changes in both liquid and vapor phase conductance. This sensitivity allows for optimization of carbon and water resource use without compromising the root-shoot hydraulic link.     

Impacts of seasonal air and soil temperatures on photosynthesis in Scots pine trees

Seasonal courses of light-saturated rate of net photosynthesis (A360) and stomatal conductance (gs) were examined in detached 1-year-old needles of Scots pine (Pinus sylvestris L.) from early April to mid-November. To evaluate the effects of soil frost and low soil temperatures on gas exchange, the extent and duration of soil frost, as well as the onset of soil warming, were manipulated in the field. During spring, early summer and autumn, the patterns of A360 and gs in needles from the control and warm-soil plots were generally strongly related to daily mean air temperatures and the frequency of severe frost. The warm-soil treatment had little effect on gas exchange, although mean soil temperature in the warm-soil plot was 3.8 degrees C higher than in the control plot during spring and summer, indicating that A360 and gs in needles from control trees were not limited by low soil temperature alone. In contrast, prolonged exposure to soil temperatures slightly above 0 degrees C severely restricted recovery of A360 and especially gs in needles from the cold-soil treatment during spring and early summer; however, full recovery of both A360 and gs occurred in late summer. We conclude that inhibition of A360 by low soil temperatures is related to both stomatal closure and effects on the biochemistry of photosynthesis, the relative importance of which appeared to vary during spring and early summer. During the autumn, soil temperatures as low as 8 degrees C did not affect either A360 or gs.     

不同立地、土壤类型对土壤含水量和苹果新梢生长的影响

为了给苹果园土壤水分管理提供依据,以‘长富2’苹果为试材,选取粘壤土平地、粘壤土梯田、壤土平地、壤土梯田、沙土平地、沙土梯田、上层壤土、下层尾矿砂平地7种不同立地、土壤类型苹果园,研究其对土壤含水量及苹果新梢生长的影响。结果表明:不同时期粘壤土平地的土壤含水量均极显著高于沙土平地、沙土梯田、上层壤土下层尾矿砂平地的土壤含水量,沙土平地、沙土梯田的土壤含水量均极显著低于其他立地、土壤类型。各立地、土壤类型长、中、短新梢均只有一次生长,迅速生长期基本一致,在4月初到5月底;除沙土平地和沙土梯田类型外,其他5种立地、土壤类型徒长新梢均出现了二次生长,第1次迅速生长期与其他新梢一致,第2次生长期在7月至8月。不同立地、土壤类型土壤含水量与苹果各类新梢长度均呈正相关,达到显著相关水平的时期多出现在新梢生长高峰期,其中,4月22日至5月13日,壤土梯田、沙土平地的土壤含水量与长梢长度的相关系数分别为0.968、0.966,5月14日至5月31日,壤土梯田的土壤含水量与徒长新梢长度的相关系数为0.986,壤土平地的土壤含水量与长梢长度的相关系数为0.980。     

Seasonal and spatial variations in leaf nitrogen content and resorption in a Quercus serrata canopy

To elucidate the relationships between spatiotemporal changes in leaf nitrogen (N) content and canopy dynamics, changes in leaf N and distribution in the canopy of a 26-year-old deciduous oak (Quercus serrata Thunb. ex. Murray) stand were monitored throughout the developmental sequence from leaf expansion to senescence, by estimating the leaf mass and N concentrations of all the canopy layers. Seasonal changes were observed in leaf N concentration per unit leaf dry mass (N (m)), which peaked after bud burst, declined for two weeks shortly thereafter, and then remained constant for the rest of the growing season for each canopy layer. Leaf N concentration per unit leaf area (N (a)) was higher in the upper layer than in the lower layer throughout the growing season, and was closely correlated with relative irradiance (RI) in the summer when the air temperature was moderately high. The N concentrations of all leaf layers started to decrease in November, and reached their lowest values in late November, whereas LMA scarcely changed throughout the season. The lowest N concentrations did not differ significantly among the canopy layers. Seasonal changes in the relationship between N (a) and RI were detected, indicating that N (a) is optimized temporally as well as spatially. Nitrogen resorption efficiency was highest in the upper canopy layers where larger amounts of N were invested. Based on the estimates of leaf mass and leaf N concentrations of the canopy layers, total leaf N concentration of the whole canopy was estimated to be 84.1 kg ha(-1) in the summer, and 37.3 kg ha(-1) in late November. Therefore, 46.8 kg ha(-1) of leaf N in the canopy (about 56% of the total N) was resorbed just before leaf abscission.     

Xylem dysfunction in Quercus: vessel sizes, tyloses, cavitation and seasonal changes in embolism

The seasonal progression of xylem dysfunction from tyloses and embolism induced both by cavitation and frost was studied in Quercus rubra L. and Quercus alba L. branches. Vessel lengths and diameters were measured in current-year rings of branches of various ages. Vessels in current-year shoots are about the same size as those in many diffuse porous trees, but vessels in older branches are two to six times larger in diameter and typically more than 10 times longer. Large Quercus vessels were more vulnerable to cavitation than small vessels. The small vessels in current-year shoots were more vulnerable to cavitation than vessels of comparable size in diffuse porous species. Earlywood vessels are completely blocked by tyloses within a year of their formation. Tylose growth starts in winter, but the vessels are not fully blocked until the next summer. Many latewood vessels, by contrast, remain free of complete blockage for several years. In Q. rubra, loss of hydraulic conductivity in current-year shoots due to cavitation reaches 20% by August and > 90% after the first hard frost. Both laboratory and field observations confirm that the role of frost in causing loss of hydraulic conduction by embolism is much more dramatic in Quercus than in conifers and diffuse porous hardwoods.     

Stomatal patchiness in the Mediterranean holm oak (Quercus ilex L.) under water stress in the nursery and in the forest

The evergreen holm oak Quercus ilex L. is the most representative tree in Mediterranean forests. Accurate estimation of the limiting factors of photosynthesis for Q. ilex and the prediction of ecosystem water-use efficiency by mechanistic models can be achieved only by establishing whether this species shows heterogenic stomatal aperture, and, if so, the circumstances in which this occurs. Here, we collected gas-exchange and chlorophyll fluorescence data in Q. ilex leaves from a nursery to measure the effects of stomatal oscillations on PSII quantum yield (Φ(PSII)) under water stress. Stomatal conductance (g(s)) was used as an integrative indicator of the degree of water stress. Images of chlorophyll fluorescence showed heterogeneous Φ(PSII) when g(s) was <50 mmol H(2)O m(-2) s(-1), representative of severe drought and corresponding to a container capacity <45%. Stomatal patchiness was related to a coefficient of variation (CV) of Φ(PSII) values >2.5%. A parallel study in the forest confirmed heterogeneous Φ(PSII) values in leaves in response to declining water availability. Three kinds of Q. ilex individuals were distinguished: those resprouting after a clear-cut (resprouts, R); intact individuals growing in the same clear-cut area as resprouts (controls, C); and intact individuals in a nearby, undisturbed area (forest controls, CF). Patchiness increased in C and CF in response to increasing drought from early May to late July, whereas in R, Φ(PSII) values were maintained as a result of their improved water relations since the pre-existing roots were associated with a smaller aerial biomass. Patchiness was related to a % CV of Φ(PSII) values >4 and associated in the summer with mean g(s) values of 30 mmol H(2)O m(-2) s(-1). Under milder drought in spring, Φ(PSII) patchiness was less strictly related to g(s) variations, pointing to biochemical limitants of photosynthesis. The occurrence of heterogenic photosynthesis caused by patchy stomatal closure in Q. ilex during severe drought should be taken into account in ecosystem modelling in which harsher water stress conditions associated with climate change are predicted.