Combined effects of soil properties and Phytophthora cinnamomi infections on Quercus ilex decline

Aims

The importance of soil properties as determinants of tree vitality and Phytophthora cinnamomi root infections was analysed.

Methods

The study comprised 96 declining stands in western Spain, where declining and non-declining holm oak (Quercus ilex L.) trees were sampled. Soil properties (soil depth, Ah horizon thickness, texture, pH, redox potential, soil bulk density and N-NH₄ ⁺ and N-NO₃ ^? concentrations) and P. cinnamomi infections were assessed.

Results

Tree mortality rates increased with low soil bulk densities, which were also associated with more P. cinnamomi-infected trees. Occurrence of infected trees was higher in fine textured soils and in thick Ah horizons. Fine textured soils favoured trees, but with the presence of P. cinnamomi their health status deteriorated. Soil under declining trees had higher N-NO₃ ^?/N-NH₄ ⁺ ratio values than under non-declining trees. Additional soil properties changes associated to grazing were not related to decline and P. cinnamomi infections.

Conclusions

The implications of P. cinnamomi in holm oak decline and the influence of soil properties as contributors to pathogen activity were demonstrated. Fine soil textures and thick Ah horizons, usually favourable for vigour and vitality of trees growing in the Mediterranean climate, were shown to be disadvantageous soil properties if P. cinnamomi was present. Fine soil textures and thick Ah horizons are frequently related with higher levels of soil moisture, which increase the inoculum of the pathogen and favours root infection. Grazing does not seem to be directly linked to Q. ilex health status or P. cinnamomi root rot.     

The influence of summer and winter stress and water relationships on the distribution of Quercus ilex L.

Several aspects of plant-water relationships and canopy behaviour have been studied over several years in two experimental areas of Quercus ilex L. forests in the northeastern Iberian Peninsula. Water requirements, water status limits necessary for a positive carbon budget and the functional canopy behaviour in the face of abiotic stress were evaluated, in order to determine which factors influence the geographical range of these forests.The results showed that holm oak has a conservative water use, a low cuticular transpiration, a high capacity for osmotic adjustement and xerophytic characteristics in leaf morphology and canopy arrangement. More than 440 mm of annual rainfall are required for these forests to persist. Summer drought and winter cold are thus important abiotic factors limiting the distribution of Quercus ilex. In both cases, drought stress is involved.     

Quercus ilex L.: How season,Plant Organ and Extraction Procedure Can Influence Chemistry and Bioactivities

Quercus species have a plethora of applications, either in wine and wood industries, in human and animal nutrition or in human health. In order to improve the knowledge on this genus, the aim of the present study was to correlate, for the first time, the phenolic composition of different Quercus ilex L. plant tissues (leaves in two maturation stages, acorns, teguments and cotyledons) and different extraction procedures with scavenging and anticholinesterase activities. The hydromethanolic and aqueous extracts obtained showed strong radical scavenging activity against DPPH, superoxide anion radical and nitric oxide radical, leaves exhibiting higher total phenolic content and revealing the best antioxidant properties, followed by tegument and acorns. Concerning the phenolic profile, fifteen compounds were identified and quantified by HPLC‐DAD, ranging from 1568.43 to 45,803.16 mg/kg dried extract. The results indicate that Q. ilex can be a source of strong antioxidant phenolic compounds with possible interest for food and pharmaceutical industries.     

Comparison of temperature effects on soil respiration and bacterial and fungal growth rates

Temperature is an important factor regulating microbial activity and shaping the soil microbial community. Little is known, however, on how temperature affects the most important groups of the soil microorganisms, the bacteria and the fungi, in situ. We have therefore measured the instantaneous total activity (respiration rate), bacterial activity (growth rate as thymidine incorporation rate) and fungal activity (growth rate as acetate-in-ergosterol incorporation rate) in soil at different temperatures (0-45 degrees C). Two soils were compared: one was an agricultural soil low in organic matter and with high pH, and the other was a forest humus soil with high organic matter content and low pH. Fungal and bacterial growth rates had optimum temperatures around 25-30 degrees C, while at higher temperatures lower values were found. This decrease was more drastic for fungi than for bacteria, resulting in an increase in the ratio of bacterial to fungal growth rate at higher temperatures. A tendency towards the opposite effect was observed at low temperatures, indicating that fungi were more adapted to low-temperature conditions than bacteria. The temperature dependence of all three activities was well modelled by the square root (Ratkowsky) model below the optimum temperature for fungal and bacterial growth. The respiration rate increased over almost the whole temperature range, showing the highest value at around 45 degrees C. Thus, at temperatures above 30 degrees C there was an uncoupling between the instantaneous respiration rate and bacterial and fungal activity. At these high temperatures, the respiration rate closely followed the Arrhenius temperature relationship.     

Comparative ecophysiological effects of drought on seedlings of the Mediterranean water-saver Pinus halepensis and water-spenders Quercus coccifera and Quercus ilex

Ecophysiological and structural traits of seedlings of the water-saver Pinus halepensis and the water-spenders Quercus coccifera and Q. ilex were studied in response to water stress under greenhouse conditions. Water deficit reduced stomatal conductance (g _s) and, as a consequence, both net CO₂ assimilation (A) and transpiration rate (E) were also reduced. Water stress also emphasized midday down-regulation of the photochemical efficiency (dynamic photoinhibition) reducing quantum yield of noncyclic electron transport (Φ_PSII), photochemical quenching (qP) and photochemical efficiency of the open reaction centres of PSII () and involved an increase of thermal dissipation of excess energy. However, water stress not only induced dynamic photoinhibition but also brought a reduction in F _v/F _m (chronic photoinhibition). Despite the water-saving strategy of P. halepensis that limited net CO₂ assimilation, this species showed a higher photochemical efficiency and lower photoinhibition than Quercus species. This was not the result of a different photochemical quenching but was linked to a higher value of , indicating a less severe photo-inactivation of PSII. Water stress resulted in a loss of pigment content and in an increase of the carotenoids/chlorophyll ratio, antioxidant capacity and the biomass rate allocated to roots as opposed to that assigned to leaves. P. halepensis showed a lower photoinhibition and antioxidant activity than Quercus species due to its lower pigment content and higher proportion of carotenoids allowing P. halepensis to use, in a more effective way, the lesser excess energy absorbed.     

Different responses to drought and freeze stress of Quercus ilex L. growing along a latitudinal gradient

The vulnerability to drought and freeze stress was measured in young plants of Quercus ilex L. growing in the field in two natural sites within the Italian distribution area of this species, i.e. Sicily (Southern Mediterranean Basin) and Venezia Giulia (Northeastern Italy), respectively. In particular, the resistance strategies adopted by Q. ilex to withstand the two stresses were estimated in terms of seasonal and/or diurnal changes in leaf conductance to water vapour (g_L), water potential (_L) and relative water content (RWC) as well as of xylem embolism in the stem and root hydraulic conductance (K_RL). Sicilian (SI) plants showed to reduce water loss by stomatal closure (g_L decreased) in summer, thus maintaining average RWCs at 88–90%. Moreover, SI plants showed considerable resistance to xylem cavitation in the stem (the loss of hydraulic conductance, PLC, was less than 12% throughout the year) and to maintain the hydraulic conductance of their roots (K_RL), constantly high even in summer. Plants growing in Venezia Giulia (VG plants), on the contrary, underwent leaf dehydration in the winter due to freeze stress so that RWC measured in April was still 78% on a diurnal basis. This was apparently due to consistent xylem embolism in the stem. In fact, PLC was as high as 40% between November and March. Only in the summer was PLC similar to that recorded in SI plants. Moreover, K_RL of VG plants decreased in November from about 1.5 to 0.8×10^–4 kg s^–1 m^–2 MPa^–1, i.e. about 50%, and in February K_RL dropped further to 0.4×10^–4 kg s^–1 m^–2 MPa^–1. On the basis of the above, we conclude that: (a) Q. ilex was more sensitive to freeze than to drought stress so that freeze stress can be considered as a factor limiting the distribution area of this species; (b) drought and freeze stress were faced by Q. ilex adopting two different resistance strategies, i.e. drought avoidance based on water saving in Sicily and freeze tolerance in Venezia Giulia.     

Effect of simazine,lindane and ceresan on soil respiration and nitrification rates

Summary The effect of three pesticidesviz, simazine, lindane and ceresan upto 100-fold of field application was studied on organic matter mineralization as well as on nitrification rates in Delhi alluvial soil. The normal rate of simazine (2 ppm) did not influence CO₂ evolution but higher concentrations suppressed it. Lindane and ceresan inhibited CO₂ production from soil by normal concentrations but the same CO₂ production was enhanced in berseem roots treated soil upto 10 ppm of ceresan.The nitrification of ammonium sulphate in soil due to these pesticides was impaired significantly for varying periods. The inhibitory effect of higher concentrations was marked only upto 3 weeks, subsequently the rate of nitrification was restored. Among the nitrifiers, Nitrobacter were more susceptible as compared to Nitrosomonas.     

Plant diversity and insect herbivores: effects of environmental change in contrasting model systems

There is increasing concern over the potential impact of anthropogenic factors (e.g. increasing nutrient inputs, global climate change) on the rate of loss of diversity in ecosystems. Such losses may affect ecosystem processes. In addition, a change in diversity of one group of organisms may influence the diversity of species of the next trophic level. We examined the extent to which plant species richness influences that of insect herbivores in two systems: a long‐term field experiment on heather moorland and a model community in the Ecotron controlled environment facility. We examined the response of these two plant communities to environmental change, specifically increased levels of nutrients, grazing and atmospheric CO₂. We measured the indirect effects of changes in these factors on insect herbivores, both above‐ and below‐ground. In the moorland system, grazing was the largest influence on plant community structure. The community was dominated by one species, Calluna vulgaris, and loss of cover under heavy grazing allowed competing species to invade. However, grazing regime was not a major influence on the species richness of the insect herbivore community. Site was more important: there were a greater number of Hemiptera species on sites with more mineral soils than on peat sites, possibly because a greater variety of grass and herb species was present on the former sites. In the Ecotron, below‐ground factors were also important drivers of community change: elevated CO₂ increased carbon availability in the soil and there were simultaneous changes in the community composition of soil biota. Above‐ground, some plant species increased in abundance and others decreased, leading to interaction‐specific effects on the insect herbivores. In two very different studies of the effects of environmental change on the interactions between plants and their herbivores, several similar conclusions can be drawn: (1) effects are likely to be site‐ and interaction‐specific; (2) outcomes are likely to be strongly dependent on the initial state and the dominant species of the plant community; and (3) indirect effects, often mediated by below‐ground factors, may have a bigger influence on insect‐plant interactions than more direct effects of above‐ground factors.     

Changes in the protein profile of Quercus ilex leaves in response to drought stress and recovery

To characterize the molecular response of holm oak to drought stress and its capacity to recover 9-month-old Quercus ilex seedlings were subjected to three treatments for a 14-d period: (i) continuous watering to field capacity (control plants, W), (ii) no irrigation (drought treatment, D), and (iii) no irrigation for 7d followed by a watering period of 7d (recovery treatment, R). In drought plants, leaf water potential decreased from -0.72 (day 0) to -0.99MPa (day 7), and -1.50MPa (day 14). Shoot relative water content decreased from 49.3% (day 0) to 47.7% (day 7) and 40.8% (day 14). Photosystem II quantum yield decreased from 0.80 (day 0) to 0.72 (day 7) and 0.73 (day 14). Plants subjected to water withholding for 7d reached, after a 7-d rewatering period, values similar to those of continuously irrigated control plants. Changes in the leaf protein pattern in response to drought and recovery treatments were analyzed by using a proteomic approach. Twenty-three different spots were observed when comparing the two-dimensional electrophoresis profile of control to both drought and recovered plants. From these, 14 proteins were identified from tryptic peptides tandem mass spectra by using the new Paragon algorithm present in the ProteinPilot software. The proteins identified belong to the photosynthesis, carbohydrate and nitrogen metabolism, and stress-related protein functional categories.     

On the monoterpene emission under heat stress and on the increased thermotolerance of leaves of Quercus ilex L. fumigated with selected monoterpenes

Leaves of the monoterpene emitter Quercus ilex were exposed to a temperature ramp with 5 °C steps from 30 to 55 °C while maintained under conditions in which endogenous emission of monoterpenes was allowed or suppressed, or under fumigation with selected exogenous monoterpenes. Fumigation with monoterpenes reduced the decline of photosynthesis, photorespiration and monoterpene emission found in non-fumigated leaves exposed to high temperatures. It also substantially increased respiration when photosynthesis and photorespiration were inhibited by low O₂ and CO₂-free air. These results indicate that, as previously reported for isoprene, monoterpenes may help plants cope with heat stress. Monoterpenes may enhance membrane stability, thus providing a rather non-specific protection of photosynthetic and respiratory processes. Monoterpene emission was maximal at a temperature of 35 °C and was inhibited at higher temperatures. This is likely to be the result of the temperature dependency of the enzymes involved in monoterpene synthesis. In contrast to other monoterpenes, cis- and trans- β -ocimene did not respond to exposure to high temperatures. Cis- β -ocimene also did not respond to low O₂ or to fumigation. These results indicate that cis and trans- β -ocimene may have a different pathway of formation that probably does not involve enzymatic synthesis.     

Geographical and seasonal influences on the distribution of fungal endophytes in Quercus ilex

A systematic survey of the endophytic assemblages of Quercus ilex in central Spain has been performed, with the goal of evaluating the importance of geographical and seasonal factors on these fungal communities. Four sampling sites were selected; one of them was sampled twice, in the spring and the autumn. The collected plant material consisted of bark, twigs and leaves from eight trees per site. Fungal strains were isolated with the use of a surface-sterilization method with sodium hypochlorite. A total of 2921 fungal strains grouped into 149 'species' or morphological types were recovered. The 10 dominant species, with isolation frequencies >1.5%, were Pyrenochaeta sp., Periconiella anamorph of Biscogniauxia mediterranea (De Not.) Kuntze, Pseudonectria sp., Cryptosporiopsis quercina Petrak, Alternaria alternata (Fr:) Keissl., two undetermined coelomycetes, Penicillium funiculosum Thom, Diplodia mutila Fr. apud Mont. and Ascochyta sp. Medians of fungal species per tree were significantly different among the sampled sites. The isolation frequencies of the dominant species, as well as other less frequent species, were significantly dependent on the sampling site. The degree of endophytic infection and the diversity of fungal species were significantly higher in the spring. The frequencies of all dominant species at one of the sites depended significantly on the season, except for C. quercina , Acremonium sclerotigenum (F & V Moreau ex Valenta) Gams. and D. mutila . Cluster analysis of the whole endophytic mycoflora of the sampled trees suggested that the geographical factor affects the endophytic distribution patterns more significantly than the seasonal factor.     

Seasonal emission of monoterpenes by the Mediterranean tree Quercus ilex in field conditions: Relations with photosynthetic rates, temperature and volatility

The relationships of monoterpene emission with temperature, light, photosynthesis and stomatal conductance (g_s) were studied in Quercus ilex L. trees throughout the four annual seasons under field conditions. The highest monoterpene emission was measured in spring and summer (midday average of 11 μg [g DW]^?1 h^?1), whereas the lowest rates were found in autumn and winter (midday averages of 0.51 and 0.23 μg [g DW]^?1 h^?1, respectively). In spring and summer, limonene was the monoterpene emitted at highest rate (midday averages of 5.27–6.69 μg [g DW]^?1 h^?1), whereas α-pinene was emitted the most in autumn and winter (midday averages of 0.31 μg [g DW]^?1 h^?1). The monoterpenes limonene, α-pinene and β-pinene represented about 75–95% of total detected monoterpenes. The total monoterpene emission rates represented about 0.04% of carbon fixed in autumn, 0.17% in winter, 0.84–2.51% in spring and 1.22–5.13% in summer. Significant correlations of total monoterpene emission with temperature were found when considering either summer emission or the emission over the entire year, whereas significant correlations with net photosynthetic rates were only found when considering summer season. Among individual terpenes, the most volatile, α-pinene and β-pinene, were more correlated with temperature than with net photosynthetic rates whereas the less volatile limonene was more correlated with net photosynthetic rate. Thus, under field conditions it seems that dependency of monoterpene emission on photosynthetic rate or temperature is partly related with volatility of the compounds. Influences of seasonality, temperature, photosynthetic rates and volatility should be considered in inventories and models of emission rates in Mediterranean ecosystems.     

Importance of microhabitat and acorn burial on Quercus ilex early recruitment: non-additive effects on multiple demographic processes

This study investigates the effect of microhabitat and seed burial on the main demographic processes operating during the early recruitment of Quercus ilex, such as postdispersal seed predation, seed germination, and seedling emergence, survival and growth. The effect of burial was positive over all the processes analysed in this study, since predation rate was lower (63.6% vs. 88%), whereas germination (53.1% vs. 21.8%) and emergence (32.0% vs. 5.5%) were higher for buried acorns. The quality of some microhabitats remained similar throughout the stages and processes studied. Thus, afforestation provided especially suitable microhabitats for oak establishment, since seed predation was lower, while germination, emergence and seedling survival were higher, than in any other microhabitat. By contrast, the quality of some microhabitats, such as open sites and Holm oaks, differed between recruitment stages. Acorns in open sites escaped predation and germinated easily, but most seedlings died due to summer drought. Similarly, although acorns under Holm oaks can germinate and survive drought, they cannot survive to postdispersal predators. This uncoupling results in a post-dispersal change in the spatial distribution of Q. ilex recruits. Furthermore, there were significant interactions between burial and microhabitat for some demographic processes. The recruitment was in afforestations high irrespective of burial, suggesting that burial is not as beneficial in high-quality habitats as it is in lower-quality ones. An accurate understanding of plant recruitment requires the determination not only of the direct effects of limiting factors but also the potential interactions occurring between them.     

Drought stress does not protect Quercus ilex L. from ozone effects: results from a comparative study of two subspecies differing in ozone sensitivity

Long‐term effects of ozone (O₃) exposure and drought stress were assessed on two subspecies of Quercus ilex: ssp. ilex and ssp. ballota. Two‐year‐old seedlings were continuously exposed for 26 months in open‐top chambers to three O₃ treatments: charcoal filtered air, non‐filtered air and non‐filtered air supplemented with 40 nl·l^?1 O₃. Additionally, two irrigation regimes were adopted: half of the plants were well‐watered and the others received half of the water supplied to control plants. Growth, shoot water potential and gas exchange rates were assessed seasonally, and biomass accumulation was determined at the end of the experiment. Drought stress caused higher reductions of gas exchange, growth and biomass accumulation than O₃ exposure in both subspecies. The combination of O₃ and drought stress caused further decreases of accumulated aboveground biomass but no additive effects were observed on gas exchange rates or root biomass. Thus, drought stress did not protect Q. ilex from O₃ effects on biomass when the response of the whole plant was considered. Q. ilex ssp. ballota was more sensitive to O₃ and ssp. ilex was more affected by drought stress. The different O₃ sensitivity was not only related to pollutant uptake but also to the ability of plants for resource acquisition and allocation. Based on biomass dose–response functions, Q. ilex is more resistant to O₃ than other European evergreen tree species, however, O₃ represents an additional stress factor that might be impairing plant ability to withstand current and future climate change.     

Phosphorus limitation and competitive capacities of Pinus halepensis and Quercus ilex subsp. rotundifolia on different soils

Aleppo pine (Pinus halepensis) and the evergreen holm oak (Quercus ilex) dominate forest areas of the Mediterranean Basin. Both species regenerate abundantly after fires: pine through seedlings and holm oak through resprouts. Cumulative nutrient losses caused by frequent fires may have decreased soil nutrient availability in such areas. To assess the role of nitrogen and phosphorus as limiting factors for growth of these species during post-fire recovery, a field fertilisation and competition experiment was conducted in a 5-year post-fire shrubland on calcareous soil, where naturally-regenerated saplings of Aleppo pine and resprouts of interior holm oak (Quercus ilex subsp. rotundifolia) coexist. Three years after fertilisation, relative basal area increment was 56% greater in pines fertilised with 250 kg P ha^–1 than in non fertilised ones. N fertilisation had small or no effects. Interactions between N and P fertilisation were not observed. Growth of Aleppo pine only increased with P fertilisation when neighbours were removed. Hence, the negative effect of neighbours on growth was greater when P availability was enhanced by fertilisation. In contrast, holm oak was able to grow more (110%) in response to increased P supply even without neighbour removal. A common garden experiment was then conducted with potted seedlings to investigate whether the suggested higher competitive capacity of holm oak for P held under a range of P amendments on different soils and competitive situations. P fertilisation increased seedling biomass yield of both species. When P availability increased, a negative effect of neighbours on growth was observed for holm oak and in 70 a lesser extent for Aleppo pine. In conclusion, in the field, holm oak resprouts showed higher competitive ability for P uptake compared to Aleppo pine saplings, but in potted seedlings in common garden conditions this trend was not observed. Therefore holm oak is not always competitively superior to Aleppo pine for P. Potted seedlings of both species had a notable plasticity in shoot/root biomass allocation, but only holm oak increased its proportional allocation to roots when neighbours were present. P availability can be a key factor in growth and competitive relations of these two species, but effects differ depending on soil type, individual age, regeneration type (i.e., seedling versus resprouts), and competitive situation. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of nitrogen deposition,drought and their interaction,on functional and structural traits of Fraxinus ornus L. and Quercus ilex L.

A controlled experiment was conducted in order to understand how functional and structural traits of species with different leaf habits (Fraxinus ornus and Quercus ilex) shift as a consequence of nitrogen (N) addition (30 kg ha yr^?1) and to explore the effect that N has on the water stress response. The experiment was divided in two stages: stage I, N addition under well-watered condition; stage II, N addition under drought. Functionality of the photosynthetic machinery, growth and biomass partitioning were assessed. The N content at leaf level increases in F. ornus only, which invests resources on photosynthetic machinery, whereas Q. ilex tends to store N in non-photosynthetic biomass, increasing relative growth rate and biomass, resulting in different allometric ratio. This effect may play a role in water stress response. Stomatal conductance of Q. ilex treated with N and subjected to water stress is lower relative to drought treatment. On the contrary, F. ornus takes advantage of N addition that has ameliorative effects on its functionality when drought was imposed. The obtained results, highlighting response mechanisms to multiple stress factors, should help to better understand and assess the performance of forest ecosystems under the foreseen environmental changes.     

环境因子和生物因子对黄河三角洲滨海湿地土壤呼吸的影响

采用Li-8150多通道土壤呼吸自动测量系统对黄河三角洲滨海湿地土壤呼吸进行全年连续测定,同步测量了温度、土壤含水量、地上生物量以及叶面积指数等环境因子和生物因子.结果表明: 土壤呼吸日动态在全年尺度上多呈单峰型,但在受到土壤封冻和地表积水干扰时,土壤呼吸日动态呈多峰型.土壤呼吸具有明显的季节动态特征,总体呈单峰型,年平均土壤呼吸速率为0.85 μmol CO₂·m^-2·s^-1,生长季平均土壤呼吸速率为1.22 μmol CO₂·m^-2·s^-1.在全年尺度上,土壤温度是滨海湿地土壤呼吸的主要控制因子,可解释全年土壤呼吸87.5%的变化.在生长季尺度上,土壤含水量和叶面积指数对土壤呼吸的协同影响达到85%.