Effect of water stress on monoterpene emissions from young potted holm oak (Quercus ilex L.) trees

We investigated the effects of a short period of water stress on monoterpene emissions from Quercus ilex, a common oak species of the Mediterranean vegetation and a strong emitter of monoterpenes. The experiment was carried out on two young saplings with a branch enclosure system under semi-controlled conditions. Under unstressed conditions, small qualitative (cis--ocimene, trans--ocimene, -caryophyllene and 1,8-cineol) and large quantitative (as much as 40% for the main compounds emitted) differences were observed between the two apparently similar trees. Nevertheless these differences did not affect the short-and long-term responses to temperature and water stress. Daily courses of emissions and gas exchanges were similar before and after the stress. During the most severe stress, emissions were reduced by a factor of two orders of magnitude and the log-linear relationship between emissions and temperature no longer existed. Photosynthesis and transpiration rates decreased as soon as the soil started to dry, whereas monoterpene emissions slightly increased for few days and then dropped when the daily CO₂ balance approached zero. We concluded that under water stress monoterpene emissions were highly limited by monoterpene synthesis resulting from a lack of carbon substrate and/or ATP. After rewatering, both emissions and gas exchanges recovered immediately, but to a level lower than the pre-stress level. These results have many implications for monoterpene emission modelling in the Mediterranean area, since the dry period generally extends from May to August. If our results are confirmed by field experiments, water stress could lead to a large overestimation of the emissions under summer conditions, when the algorithms based on light and temperature would give high emission rates.     

Light and temperature dependence of the emission of cyclic and acyclic monoterpenes from holm oak (Quercus ilex L.) leaves

In a laboratory study, we investigated the monoterpene emissions from Quercus ilex, an evergreen sclerophyllous Mediterranean oak species whose emissions are light dependent. We examined the light and temperature responses of individual monoterpenes emitted from leaves under various conditions, the effect of heat stress on emissions, and the emission-onset during leaf development. Emission rate increased 10-fold during leaf growth, with slight changes in the composition. At 30 °C and saturating light, the monoterpene emission rate from mature leaves averaged 4·1 nmol m^–2 s^–1, of which α-pinene, sabinene and β-pinene accounted for 85%. The light dependence of emission was similar for all monoterpenes: it resembled the light saturation curve of CO₂ assimilation, although monoterpene emission continued in the dark. Temperature dependence differed among emitted compounds: most of them exhibited an exponential increase up to 35 °C, a maximum at 42 °C, and a slight decline at higher temperatures. However, the two acyclic isomers cis-β-ocimene and trans-β-ocimene were hardly detected below 35 °C, but their emission rates increased above this temperature as the emission rates of other compounds fell, so that total emission of monoterpenes exponentially increased from 5 to 45 °C. The ratio between ocimene isomers and other compounds increased with both absolute temperature and time of heat exposure. The light dependence of emission was insensitive to the temperature at which it was measured, and vice versa the temperature dependence was insensitive to the light regime. The results demonstrated that none of the models currently applied to simulate isoprene or monoterpene emissions correctly predicts the short-term effects of light and temperature on Q. ilex emissions. The percentage of fixed carbon lost immediately as monoterpenes ranged between 0·1 and 6·0% depending on temperature, but rose up to 20% when leaves were continuously exposed to temperatures between 40 and 45 °C.     

Influence of different post-planting treatments on the development in Holm oak afforestation

This work examines the influence of different post-planting treatments in the development of afforested Holm oak, Quercus ilex L. subsp. ballota (Desf.) Samp., in an abandoned agricultural field in SE Spain. A total of nine post-planting treatments were carried out: tilling spring and autumn in different years, spreading of organic and inorganic mulch, irrigation in different periods. Soil preparation prior to planting consisted of digging of a hole with a backhoe excavator of 80 HP and homogenized the profile characteristics. Survival, nutrient concentrations in both leaves and soil, and foliage surface have been studied. Dry-season irrigation every 2 weeks provided the best survival rate (98.3%), while mulching with solid urban wastes caused the worst results (6.6%). The supply of forest debris increased the N, K, and Mn concentrations at all depths with respect to the other treatments, and the solid urban waste had the same effect in N, P, K, Na, Cu, Fe, and Zn. All the foliar nutrients decreased in 2005 with respect to the 2004 sampling, except in N, Cu, and Zn. The use of foliar nutrient concentrations, together with the increase in leaf surface is proposed as a non-destructive index of development and of forestation stands in the first stages of growth. The application of forest debris, and particularly the placement of boulders in the beds of individual trees proved most effective in sapling development.     

The holm oak leaf proteome: analytical and biological variability in the protein expression level assessed by 2-DE and protein identification tandem mass spectrometry de novo sequencing and sequence similarity searching

As a first approach in establishing the holm oak leaf proteome, we have optimised a protocol for this plant and tissue which includes the following steps: trichloroacetic acid-acetone extraction, two-dimensional gel electrophoresis (2-DE) on pH 5 to 8 linear gradient immobilised pH gradient strips as the first dimension, and sodium dodecyl sulfate-polyacrylamide gel electrophoresis on 13% polyacrylamide gels as the second one. Proteins were detected by Coomassie staining. Gel images were recorded and digitalized, and the protein spots quantified by using a linear regression equation of protein quantity on spot volume obtained against standard proteins. Analytical variance was calculated for one-hundred protein spots from three replicate 2-DE gels of the same protein extract. Biological variance was determined for the same protein spots from independent tissue extracts corresponding to leaves from different trees, or the same tree at different orientations or sampling times during a day. Values of 26% for the analytical variance and 58.6% for the biological variance among independent trees were obtained. These values provide a quantified and statistical basis for the evaluation of protein expression changes in comparative proteomic investigations with this species. A representative set of the major proteins, covering the isoelectric point range of 5 to 8 and the relative molecular mass(r) range of 14 to 78 kDa, were subjected to liquid chromatography-tandem mass spectrometry analysis. Due to the absence of Quercus DNA or protein sequence databases, a method based on the procedure reported by Liska and Shevchenko including de novo sequencing and BLAST similarity searching against other plant species databases was used for protein identification. Out of 43 analysed spots, 35 were positively identified. The identified proteins mainly corresponded to enzymes involved in photosynthesis and energetic metabolism, with a significant number corresponding to RubisCO.     

The role of climate change in the widespread mortality of holm oak in open woodlands of Southwestern Spain

Forest decline and increasing tree mortality are of global concern and the identification of the causes is necessary to develop preventive measures. Global warming is an emerging factor responsible for the increasing tree mortality in drought-prone ecosystems. In the southwestern Iberian Peninsula, Mediterranean holm oak open woodlands currently undergo large-scale population-level tree die-off. In this region, temperature and aridity have increased during recent decades, but the possible role of climate change in the current oak mortality has not been investigated.To assess the role of climate change in oak die-off in managed open woodlands in southwestern Spain, we analyzed climate change-related signals in century-long tree ring chronologies of dead holm oaks. We examined the high/low-frequency variability in growth and the relationship between growth and climate.Similar to other Mediterranean forests, growth was favored by precipitation from autumn of the year prior to ring formation to spring of the year of ring formation, whereas high temperatures during spring limited growth. Since the 1970s, the intensity of the high-frequency response to water availability increased simultaneously with temperature and aridity. The growth trends matched those of climatic changes. Growth suppressions occurred during droughts in the 1970s, 1980s and 1990s. Widespread stand-level, age-independent mortality occurred since 2005 and affected trees that cannot be considered old for the species standards.The close relationship between growth and climate indicate that climate change strongly controlled the growth patterns. This suggests that harsher climatic conditions, especially increased aridity, affected the tree performance and could have played a significant role in the mortality process. Climate change may have exacerbated or predisposed trees to the impact of other factors (e.g. intense management and pathogens). These observations could suggest a similar future increase in oak mortality which may occur in more northern oak open woodlands if aridity further increases.     

Testing the effect of restoration‐focused silviculture on oak regeneration and groundlayer plant communities in urban–exurban oak woodlands

Throughout their global range, oak‐dominated ecosystems have undergone state changes in stand structure and composition. Land managers face an especially acute challenge in restoring oak ecosystems and promoting oak regeneration in urban–exurban areas, where high‐intensity silvicultural treatments are often not feasible. To investigate low‐intensity management alternatives which could be widely applied in urban–exurban forests, a large‐scale adaptive management experiment was implemented in Lake County, IL, in 2012. Five canopy manipulation treatments of varying intensity, timing, and spatial aggregation were replicated across three study areas and oak seedlings were under‐planted into treatment units following management. Responses of understory light environment, shrub and groundlayer plant communities, and survival and growth of underplanted oak seedlings were evaluated. Understory light availability, canopy openness, total groundlayer plant cover, and groundlayer species diversity all differed among treatments. However, although understory light availability was significantly increased by canopy manipulation, groundlayer communities and oak seedling survival and growth did not differ among treatments. High overall seedling survival rates suggest current conditions are amenable to oak regeneration, but long‐term monitoring will be needed to assess the potential for seedlings to transition to the sapling and canopy layers. Early results demonstrate that canopy‐focused silvicultural treatments can affect the understory light environment and, to some degree, groundlayer plant communities. However, underplanting of oak seedlings paired with subcanopy thinning may be sufficient to restore an oak seedling layer, and (when necessary or preferred) canopy manipulation could potentially be deferred until later in the restoration timeline to promote oak recruitment.     

Comparison of the hydrological characteristics of three small experimental holm oak forested catchments in NE Spain in relation to larger areas

Precipitation and streamflow have been measured in three small (0.04–0.52 km²) experimental catchments covered by dense holm oak (Quercus ilex L.) forests. Two of them are in the Prades mountains and one in the Montseny mountains (NE Spain). Here we test the hydrological representativeness of these catchments against the streamflow record at two nearby larger (34–60 km²) catchments, one from each massif. Comparisons of (i) annual streamflow, (ii) seasonal distribution of streamflow, and (iii) flow duration curves were made. At Prades, for the period of common record, mean annual precipitation was about 580 mm, and mean annual streamflow 44–81 mm at the two experimental catchments and 102 mm at the larger one. Most streamflow occurred during winter and spring in the three catchments. At Montseny, rainfall was higher, and mean annual streamflow was 495 mm in the experimental catchment, and 760 mm in the larger catchment, though these data were obtained in different periods in each catchment. Streamflow was roughly equal in autumn, winter and spring. At both sites flow duration curves were fairly similar in the small experimental catchments and the larger catchments. The higher streamflow at Montseny is reflected in its flow duration curves being well above those at Prades. The experimental catchments at Prades are thus fairly representative of the larger nearby catchment for the investigated hydrological characteristics. At Montseny, hydrological differences between the experimental catchment and the larger catchment are probably due to the higher mean altitude of the latter and to the non-overlapping periods of their streamflow records.     

Variation in the genetic structure and reproductive biology of holm oak populations

Five polymorphic enzymatic loci were studied in 30 populations of Quercus ilex distributed from Crete to North Africa. A high within-population genetic diversity was found for each locus studied. Most of the alleles were common to all the populations, indicating that the holm oak corresponds to a single genetic entity. Observation of breeding system characteristics showed that the holm oak is monoecious, wind-pollinated, and likely to possess genetic autoincompatibility. Furthermore, studies of flowering phenology in a single population showed that the period of flowering varied notably among trees. An average of 29% of the trees showed inter-annual variation in flowering time. The male, female or vegetative investment also varied widely among trees from one year to another. This results in a diversification of pollen source received by a given tree each year, and also from one year to another.     

Evaluating the benefits of CAP reforms: Can afforestations restore bird diversity in Mediterranean Spain?

Current Common Agricultural Policy (CAP) subsidies in the Mediterranean region tend to prioritize afforestation on former arable land with oaks rather than pines because pine plantations would maintain lower biological diversities than native forests. Nevertheless, no thorough evaluations of the conservation values of pine plantations as compared to oak remnants have been carried out to date. We analyze the diversity and conservation value of bird assemblages breeding in 200 remnants of Holm oak Quercus ilex woodlands and 82 mature (>50-year-old) pine plantations in central Spain, a Mediterranean region mostly devoted to arable farming. Species–area relationships were compared between forest types. The conservation value of bird assemblages was assessed using the “Species of European Conservation Concern” (SPEC) classification of Burfield and van Bommel [(2004). Birds in Europe: Populations estimates, trends and conservation status. Cambridge: BirdLife International]. Overall numbers of bird species maintained by oak and pine archipelagoes were rather similar, but species–area relationships differed between forest types. Intercepts were higher in oak fragments, whereas slopes were steeper in pine plantations. Small oak fragments held more species (mainly Mediterranean Sylvia warblers) than plantations, whereas large plantations held more species than large oak remnants. Differences in species–area relationships seemed to be due to differences in vegetation structure, especially understorey shrub cover and tree height and cover. We recorded nine SPECs, all exclusive (6) or near-exclusive (3) to oak woodlands, although such woodlands do not appear to be critical for their conservation. Hence, we conclude that pine afforestations have played a role for maintaining and restoring forest bird communities in the farming landscapes of central Spain. Promoting large and shrubby plantations would enhance their conservation value for breeding birds, together with promoting growth, regeneration and expansion of Holm oak remnants by means of set-aside measures previous or alternative to oak reafforestation. The increasing importance of non-commercial as compared to commercial values of Mediterranean forests would justify subsidizing the proposed policy.     

Red oak seedlings as indicators of deer browse pressure: Gauging the outcome of different white‐tailed deer management approaches

After decades of high deer populations, North American forests have lost much of their previous biodiversity. Any landscape‐level recovery requires substantial reductions in deer herds, but modern societies and wildlife management agencies appear unable to devise appropriate solutions to this chronic ecological and human health crisis. We evaluated the effectiveness of fertility control and hunting in reducing deer impacts at Cornell University. We estimated spring deer populations and planted Quercus rubra seedlings to assess deer browse pressure, rodent attack, and other factors compromising seedling performance. Oak seedlings protected in cages grew well, but deer annually browsed ≥60% of unprotected seedlings. Despite female sterilization rates of >90%, the deer population remained stable. Neither sterilization nor recreational hunting reduced deer browse rates and neither appears able to achieve reductions in deer populations or their impacts. We eliminated deer sterilization and recreational hunting in a core management area in favor of allowing volunteer archers to shoot deer over bait, including at night. This resulted in a substantial reduction in the deer population and a linear decline in browse rates as a function of spring deer abundance. Public trust stewardship of North American landscapes will require a fundamental overhaul in deer management to provide for a brighter future, and oak seedlings may be a promising metric to assess success. These changes will require intense public debate and may require new approaches such as regulated commercial hunting, natural dispersal, or intentional release of important deer predators (e.g., wolves and mountain lions). Such drastic changes in deer management will be highly controversial, and at present, likely difficult to implement in North America. However, the future of our forest ecosystems and their associated biodiversity will depend on evidence to guide change in landscape management and stewardship.     

Growth and yield of young Quercus ilex coppice stands in the Tafferte forest (Morocco)

The coppice method is the silvicultural system commonly applied to Quercus ilex (holm oak) forests for the production of firewood and charcoal. Although the holm oak is the species that covers the largest area in Morocco (1400 000 hectares), little is known about its growth and yield. The objective of this study is to quantify the stand growth and yield of young coppice holm oak stands in the Tafferete forest in the eastern part of the Middle Atlas (Morocco).Equations relating dominant height to age, and basal area, volume, and number of trees to dominant height were developed and used to derive young holm oak coppice yield estimates.The growth and yield information developed in this study may have a greater utility in defining cultural measures that should be applied in the early stage of holm-oak coppice stands.     

Biomass,production and nutrient distribution of a natural oak forest in central Korea

Biomass, production, and nutrient distribution of a pure Quercus variabilis Bl. stand (stand 1) and two mixed Q. variabilis–Q. mongolica Fisch. stands (stand 2 and 3) were investigated in central Korea. Stand 1 naturally occurred on a site with a southern aspect while stand 2 and stand 3 occurred on sites with a northern aspect. Total (overstory+understory vegetation) biomass (tha^-1) and annual production (tha^–1year^–1) were 137.8 and 11.1 for stand 1, 216.2 and 16.6 for stand 2, and 253.3 and 19.7 for stand 3. Nutrient contents (kgha^–1) in the vegetation were distributed as follows: K, 478–860; N, 471–839; Ca, 428–791; Mg, 72–125; Na, 77–141; and P, 37–71, and were greatest in stand 3 followed by stand 2, and stand 1. Stand density influenced the differences in biomass, annual production and nutrient contents in the vegetation. Forest floor dry mass and N content (kgha^–1) were 13400 and 169 for stand 1, 10400 and 133 for stand 2, and 11200 and 127 for stand 3. Total amounts of N, P and Na in the ecosystem were greatest in the upper 40cm of mineral soil followed by the vegetation and forest floor. However, the vegetation contained a greater amount of K than the mineral soil. It appeared that microenvironments, such as, aspect influenced the distribution of natural oak species within a relatively small area and resulted in differences in biomass, production and nutrient distribution among the stands.     

The role of reed management and habitat quality on brood parasitism and chick survival of the brood parasitic Common Cuckoo

Despite efforts on ecosystem restoration and management, biodiversity loss remains one of the major environmental concerns of our time. Beyond the focus on threatened species, animals that indicate regional biodiversity hotspots and population trends, such as brood parasites, should also be targeted by conservation actions. We studied how reed habitat quality and management influence brood parasitism rate and offspring survival in Common Cuckoos Cuculus canorus parasitizing nests of Great Reed Warblers Acrocephalus arundinaceus in six reed habitats in an intensive agricultural landscape. Data collected from 45 sites over 13 years showed that the brood parasitism rate was highest on large canals and was positively influenced by the availability of potential perches (Cuckoo vantage points) and the height where host nests were built. Cuckoo chick survival decreased with water depth and was not affected by other factors. Our results suggest that the habitat-dependent detectability of host nests was central in brood parasitism rate and that water level was central in Cuckoo chick survival. Our study shows that a maintenance of intermediate water levels is the most optimal for maintaining Cuckoo populations in intensive agricultural landscapes. Because brood parasites are excellent bioindicators as their presence predicts regional hotspots of taxonomic and functional diversity as well as population trends in bird communities, knowledge on their habitat requirements is relevant in management targeting diverse bird communities.     

Acorn preference under field and laboratory conditions by two flightless Iberian dung beetle species (Thorectes baraudi and Jekelius nitidus): implications for recruitment and management of oak forests in central Spain

1. Recently, a mutualistic relationship has been described between some dung beetles (Thorectes lusitanicus and Mycotrupes lethroides) and oak species (Quercus suber, Q. canariensis, and Q. rubra), which could be crucial for ensuring seedling recruitment and sustaining the equilibrium of oak populations. For T. lusitanicus, a diet based on acorns during the reproductive period improved resistance to low‐temperature conditions and improved ovarian development. 2. In this paper, we conducted field and laboratory experiments to investigate the interaction between two potential acorn‐eating beetles, Thorectes baraudi and Jekelius nitidus, with Quercus suber. We determined the feeding preferences of both beetle species and estimated the rates of acorn manipulation by beetles according to habitat structure and several characteristics of the acorn, such as seed size and acorn infestation by weevils. 3. Results demonstrated the positive interaction between the dung beetle Thorectes baraudi and Quercus trees. Thorectes baraudi was clearly more attracted to volatiles of acorns than to dung. Jekelius nitidus, on the contrary, was either not or anecdotally attracted to acorns. On the contrary, in the case of Jekelius nitidus, the acorn attraction could be considered anecdotal or even accidental. Our field results demonstrated the acorn burying behaviour of T. baraudi in the oak forests of the Cabañeros National Park (Spain), suggesting a potential role of this beetle species as an active secondary acorn disperser. 4. This unexpected behaviour could be particularly important in Mediterranean oak forests and savannahs, where most Quercus species are strongly recruitment limited because of serious overgrazing problems.     

The effect of low temperatures on the photosynthetic apparatus of Quercus ilex subsp. ballota at its lower and upper altitudinal limits in the Iberian peninsula and during a single freezing-thawing cycle

We investigated the response of the photosynthetic apparatus during an episode of extreme low winter temperature in Quercus ilex subsp. ballota (Desf.) Samp., a typical Mediterranean evergreen species in the Iberian peninsula. Both plants in a woodland located at high altitude (1,177 m. a.s.l.) and potted plants obtained from acorns of the same populations grown at low altitude (225 m. a.s.l.) were analyzed. Net CO₂ assimilation rate was negative and there was a marked decrease in photosystem II (PSII) efficiency during winter in leaves of the woodland population (high altitude individuals). These processes were accompanied by increases in non-photochemical quenching (NPQ) and in the de-epoxidated carotenoids within the xanthophyll cycle, mechanisms aimed to dissipate excess energy. In addition, these de-epoxidated carotenoids were largely preserved during the night. There was no chlorophyll bleaching during the winter, which suggests that leaves were not experiencing photoinhibitory damage. In fact, the net photosynthetic rate and the PSII efficiency recovered in spring. These changes were not observed, or were much more reduced, in individuals located at lower altitude after a few frosts. When the response to rapid temperature changes (from 20°C to –5°C and from –5°C to 20°C) was studied, it was found that the maximum potential PSII efficiency was fairly stable, ranging from 0.70 to 0.75. The rest of the photosynthetic parameters measured, actual and intrinsic PSII efficiency, photochemical and NPQ, responded immediately to the changes in temperature and, also, the recovery after cold events was practically immediate.     

农田非作物生境调控与害虫综合治理

就害虫综合治理的研究而言,农田生态系统可以区分出作物生境和邻近作物的非作物生境。昆虫从作物生境迁移到非作物生境,与作物生境缺乏食物或受人类栽培活动干扰而引起的自然迁移、转换寄主和寻求庇护场所有关。许多研究表明,与特定作物田块相联系的植被类型和结构可影响害虫及其天敌迁居的种类、数量和时间。因此,我们可以通过改变大田周围非作物生境的植被组成及特征来调控农业生态系统中害虫与天敌的相互关系,提高天敌对害虫的控制效能。由于景观的空间格局对节肢动物的生物学特性有直接或间接的影响,所以,以景观为单元要比以同类作物的田块为单元更加适于害虫综合治理的研究和实施。在现代农业景观区域内重新引入和相嵌一些非栽培植物或廊道,可为众多有益节肢动物的繁殖、取食和避害提供多种类型的非作物栖境。