Factors affecting post-fire crown regeneration in cork oak (Quercus suber L.) trees

Cork oak (Quercus suber) forests are acknowledged for their biodiversity and economic (mainly cork production) values. Wildfires are one of the main threats contributing to cork oak decline in the Mediterranean Basin, and one major question that managers face after fire in cork oak stands is whether the burned trees should be coppiced or not. This decision can be based on the degree of expected crown regeneration assessed immediately after fire. In this study we carried out a post-fire assessment of the degree of crown recovery in 858 trees being exploited for cork production in southern Portugal, 1.5 years after a wildfire. Using logistic regression, we modelled good or poor crown recovery probability as a function of tree and stand variables. The main variables influencing the likelihood of good or poor crown regeneration were bark thickness, charring height, aspect and tree diameter. We also developed management models, including simpler but easier to measure variables, which had a lower predictive power but can be used to help managers to identify, immediately after fire, trees that will likely show good crown regeneration, and trees that will likely die or show poor regeneration (and thus, potential candidates for trunk coppicing).     

Fingerprinting and genetic variability in cork oak (Quercus suber L.) elite trees using ISSR and SSR markers

Quercus suber L., is a socially, economically and ecologically important forest species in rural areas of the Mediterranean basin. Fifty three elite-trees from nine stands of four provenance regions in the Community of Extremadura (Spain) were analysed with the aim to establish their DNA-fingerprinting and the genetic relationships among them. Two types of molecular markers, microsatellites and intermicrosatellites, were used for tree genotyping. Microsatellite markers could distinguish 94% of the trees. Observed and expected heterozygosities, and effective number of alleles were correlated with the polymorphism information content (PIC). Intermicrosatellite patterns were used to construct a dendrogram. They showed high levels of genetic diversity in these elite trees, without any clear relationship with provenance regions. Both types of markers detected a high level of intra-stand variability, which is common in Quercus species, probably due to its mating system, the low distance among stands and the small size of stands. Others factors that could affect this result, such as introgression between Q. suber L. and Q. ilex L., are also commented. The high level of genetic variability detected in these elite trees can be useful for improvement programs. Also the usefulness of SSR and ISSR markers to establish the DNA-fingerprinting of each tree could be focused to select clonal lines of commercial interest.     

The diversification of terpene emissions in Mediterranean oaks: lessons from a study of Quercus suber, Quercus canariensis and its hybrid Quercus afares

Interspecific gene flow is common in oaks. In the Mediterranean, this process produced geographical differentiations and new species, which may have contributed to the diversification of the production of volatile terpenes in the oak species of this region. The endemic North African deciduous oak Quercus afares (Pomel) is considered to be a stabilized hybrid between the evergreen Quercus suber (L.) and the deciduous Quercus canariensis (Willd.), presumably being monoterpene and isoprene emitters, respectively. In a common garden experiment, we examined the terpene emission capacities, terpene synthase (TPS) activities and nuclear genetic markers in 52 trees of these three oak species. All but one of the Q. suber and Q. canariensis trees were found to be genetically pure, whereas most Q. afares trees possessed a mixed genotype with a predominance of Q. suber alleles. Analysis of the foliar terpene emissions and TPS activities revealed that all the Q. canariensis trees strongly produced isoprene while all the Q. suber trees were strong monoterpene producers. Quercus afares trees produced monoterpenes as well but at more variable and significantly lower rates, and with a monoterpene pattern different than that observed in Q. suber. Among 17 individuals tested, one Q. afares tree emitted only an insignificant amount of terpenes. No mixed isoprene/monoterpene emitter was detected. Our results suggest that the capacity and pattern of volatile terpene production in Algerian Q. afares populations have strongly diverged from those of its parental species and became quantitatively and qualitatively reduced, including the complete suppression of isoprene production.     

Water-use efficiency in cork oak (Quercus suber) is modified by the interaction of water and light availabilities

We studied the interaction of light and water on water-use efficiency in cork oak (Quercus suber L.) seedlings. One-year-old cork oak seedlings were grown in pots in a factorial experiment with four light treatments (68, 50, 15 and 5% of full sunlight) and two irrigation regimes: well watered (WW) and moderate drought stress (WS). Leaf predawn water potential, which was measured at the end of each of two cycles, did not differ among the light treatments. Water-use efficiency, assessed by carbon isotope composition (delta(13)C), tended to increase with increasing irradiance. The trend was similar in the WW and WS treatments, though with lower delta(13)C in all light treatments in the WW irrigation regime. Specific leaf area increased with decreasing irradiance, and was inversely correlated with delta(13)C. Thus, changes in delta(13)C could be explained in part by light-induced modifications in leaf morphology. The relationship between stomatal conductance to water vapor and net photosynthesis on a leaf area basis confirmed that seedlings in higher irradiances maintained a higher rate of carbon uptake at a particular stomatal conductance, implying that shaded seedlings have a lower water-use efficiency that is unrelated to water availability.     

Chemical composition and variability of cork from Quercus suber L.

Summary The chemical composition of cork from Quercus suber L. was determined for virgin cork and for reproduction cork. Average chemical composition of virgin cork is: ash 0.7%, total extractives 15.3%, suberin 38.6%, lignin 21.7% and polysaccharides 18.2%. The carbohydrate composition shows that glucose represents 50.6% of all monosaccharides, xylose 35.0%, arabinose 7.0% and galactose and mannose, respectively, 3.6% and 3.4%. Reproduction cork has a composition similar to virgin cork. For the cases studied, reproduction cork had however less extractives and a higher amount of suberin. Significant differences were found for the chemical composition of virgin cork in relation to location of extractives and polysaccharide content. The between-tree variability was large and a variation within the tree was also recorded.The author is grateful for the help of collaborators throughout the chemical analysis of the many samples: Joaquina Ferreira, A. Velez Marques, M. Lurdes Ferreira, Rosa Amorim and Célia Constâncio.The work was financially supported by the Instituto de Ciência e Tecnologia dos Materiais (ICTM), Lisbon, Portugal     

Waxes composition of Quercus suber reproduction cork from different Spanish provenances

Summary The chemical composition of waxes was studied in Quercus suber cork planks collected in seven localities of the three main production areas of Spain. Waxes were extracted with chloroform (CHCl₃) and the extract was submitted to saponification in order to obtain the neutral and acid fractions. The trimethylsilyl derivatives of both fractions were analysed by GC-MS. The seven populations cannot be distinguished by their total contents of waxes (Chloroform extract) and of the neutral and acid fraction. This lack of differences could be due to the important variability among samples of each population. The neutral fraction was mainly composed of fatty alcohols (all the even members from C₁₈ to C₂₆, with traces of the intermediate odd members and some unsaturated groups) and triterpenes (11 components, among them friedelin, betulin and cerin were identified) and a very small amount of monocarboxylic fatty acids (C₁₆ and C₂₄ members) was also present. The main group of components of the acid fraction was that of fatty acids (saturated even C₁₄–C₂₄ and odd C₁₅, C₁₇, C₂₁ members, accompanied by a great amount of unsaturated terms and some ω-hydroxyacids, 18-hydroxy-9,12-octadecadienoic and 18-hydroxy-9-octadecenoic acids). Four triterpenes and the C₂₀ and C₂₄ alcohols were also detected in the acid fraction. Significant differences were found among the corks of the various provenances in the global contents of alcohols and triterpenes and in these individual components contents of the neutral fraction. Concerning the differences among the various provenances in the contents of the groups of fatty acids, alcohols, and triterpenes in the acid fraction, a higher number of differences were obtained in the comparisons of the contents of acids and triterpenes, whereas the differences were practically null in the case of the alcohols. When the differences were studied considering the individual compounds, the relation with the behaviour of the contents of the groups of components was not as clear as it happens in the neutral fraction. In this case, the differences were scarcer and they did not show a unique pattern, neither in the components, nor the populations. There is no correspondence in the differences among the provenances regarding the components of the neutral fraction or those of the acid fraction. No clear relationship was found between the geographical proximity of the provenances and chemical similarity of the waxes composition. Received 12 December 1997     

Pathogenicity of fungi isolated from Quercus suber in Catalonia (NE Spain)

Summary Thirty‐four fungal species isolated from cork oak (Quercus suber) in Catalonia (NE Spain) during 1992–95 were tested for pathogenicity either in stem, leaf or root inoculations. Eleven species were found to be pathogenic on stem: Biscogniauxia mediterranea, Botryosphaeria stevensii, Diatrype cf. stigma, Endothia gyrosa, Fusarium solani, Graphium sp., Ophiostoma quercus, Phomopsis sp., Phytophthora cinnamomi, Sporendocladia bactrospora and an unidentified Coelomycete. Three fungi showed pathogenic effects on leaves: Dendrophoma myriadea, Lembosia quercina and Phomopsis quercella. No clear pathogenic effects were detected in the root inoculation experiment. Trunk pathogens were differentiated into two groups according to the effects induced in the inoculated plants; B. stevensii, Phomopsis sp. and P. cinnamomi caused the death of the inoculated plants and induced the formation of large cankers and vascular necroses. The other pathogenic species also produced severe cankers and vascular lesions, but no significant mortality was detected. Water stress increased the lesions caused by B. mediterranea and Phomopsis sp., but limited those of P. cinnamomi and the rest of the inoculated fungi. However, water stress did not significantly affect the damage caused by B. stevensii, which was the most virulent of the species tested. Leaf pathogens only showed their effects if the leaf cuticle was previously damaged. Lembosia quercina caused small dark lesions whereas D. myriadea and P. quercella produced large necrotic areas in well‐watered plants. The lesions caused by the last two fungi were reduced by water stress.     

Carbon sequestration of modern Quercus suber L. silvoarable agroforestry systems in Portugal: a YieldSAFE-based estimation

Modern alley cropping designs, with trees aligned in rows and adapted to operating farming machinery, have been suggested for Europe. This paper explores the potential for adoption of cork oak (Quercus suber L.) agroforestry in Portugal and estimates the potential carbon sequestration. Spatial modeling and Portuguese datasets were used to estimate target areas where cork oak could grow on farmland. Different implementation scenarios were then modeled for this area assuming a modern silvoarable agroforestry system (113 trees ha^?1 thinned at year 20 for establishing 50 trees ha^?1). The YieldSAFE process-based model was used to predict the biomass and carbon yield of cork oak under low and high soil water holding capacity levels. Approximately 353,000 ha are available in Portugal for new cork oak alley cropping. Assuming implementation rates between 10 % of the area with low soil water capacity (60 mm: 15 cm depth, coarse texture) and 70 % of the area with high soil water holding capacity (1,228 mm: 200 cm depth, very fine texture), then carbon sequestration could be 5 × 10⁶ and 123 × 10⁶ Mg CO₂ respectively. Due to higher yields on more productive land, scenarios of limited implementation in high productivity locations can sequester similar amounts of carbon as wide implementation on low productivity land, suggesting that a priori land classification assessments can improve the targeting of land and financial incentives for carbon sequestration.     

Photosynthetic and transpirational responses of red spruce understory trees to light and temperature

Understory red spruce (Picea rubens Sarg.) trees, between 20 and 50 cm in height and 12 years or more in age, were collected from mid- and high-elevation stands in north-central Vermont and placed in a closed-cuvette system to measure photosynthetic and transpirational responses to photosynthetic photon flux density (PPFD) and temperature. Photosynthesis, dark respiration, transpiration and water-use efficiency of trees from both stands responded to changes in PPFD and temperature in similar ways. Trees from both stands exhibited maximum rates of net photosynthesis at temperatures between 15 and 20 degrees C, and exposure to higher temperatures resulted in reduced rates of photosynthesis and increased rates of respiration. Net photosynthetic rates generally increased with increasing light intensity but began to level off at 250 micro mol m(-2) s(-1). Water-use efficiency was maximal when temperature and PPFD were at 15 degrees C and above 400 micro mol m(-2) s(-1), respectively.     

Morphological evaluation of cork oak (Quercus suber): Mediterranean provenance variability in Tunisia

Twenty-six provenances (2 340 plants) of cork oak (Quercus suber spp.) originating from Portugal, Spain, Italy, Morocco, Algeria, and Tunisia were tested for genetic variation among and within provenances by growth traits. Seven morphometrical characters were measured in 90 plants from each provenance. Analysis of variance showed highly significant differences for all characters. The phenotypic coefficient of differentiation reached 0.24 for the form and 0.22 for height, thus revealing a strong structuring between the provenances. Comparative study of growth among the provenances revealed more vigorous growth and better survival rate for those from Morocco, Spain, and Portugal, which may constitute better materials for afforestation. Furthermore, this variability appeared to be geographically structured and would be mainly genetically controlled, as cork oak provenances were cultivated under the same environmental conditions. Our results should be helpful for guide forest managers in afforestation.     

Effects of cadmium on cork oak (Quercus suber L.) plants grown in hydroponics

Cork oak (Quercus suber L.) is an autochthonous tree species that is being used for reforestation in heavy-metal-contaminated areas in Spain. A hydroponics experiment was carried out to characterize the effects of Cd on several morphological and physiological parameters in this species, including shoot length, nutrient concentrations and allocation in different organs, leaf pigment concentrations, photosynthetic efficiency, root ferric chelate reductase (FCR) activity and organic acid concentrations in xylem sap. Four different Cd treatments were applied, adding Cd chelated with EDTA or as chloride salt at two different concentrations (10 and 50 μM Cd). After 1 month of Cd treatment, plant growth was significantly inhibited in all treatments. Results indicate that Cd accumulates in all organs 7- to 500-fold when compared with control plants. The highest Cd concentration was found in the 50 μM CdCl(2) treatment, which led to concentrations of ~30, 123 and 1153 μg Cd g(-1) dry weight in leaves, stems and roots, respectively. In the strongest Cd treatments the concentrations of P and Ca decreased in some plant parts, whereas the Mn leaf concentrations decreased with three of the four Cd treatments applied. The concentrations of chlorophyll and carotenoids on an area basis decreased, whereas the (zeaxanthin plus antheraxanthin)/(total violaxanthin cycle carotenoids) ratio and the non-photochemical quenching increased significantly in all Cd treatments. Cadmium treatments caused significant increases in the activity of the enzyme FCR in roots and in the concentrations of organic acids in xylem sap. Some of the physiological changes found support the fact that Cd induces a deficiency of Fe in cork oak, although the plant Fe concentrations were not reduced significantly. At higher concentrations the effects of Cd were more pronounced, and were more marked when Cd was in the free ion form than when present in the form of Cd-EDTA.     

Interactive responses of Quercus suber L. seedlings to light and mild water stress: effects on morphology and gas exchange traits

? The combined effect of water stress and light on seedlings of forest species is a key factor to determine the best silvicultural and afforestation practices in the Mediterranean area.

? The aims of this work was (1) to determine the optimal light level for the early development of cork oak seedlings under mild water stress and (2) to test if the combined effect of water stress and light followed the trade-off, the facilitation or the orthogonal hypothesis.

? Shade reduced instantaneous photosynthetic rates and water use efficiency in cork oak. However, seedlings grown under moderate shade (15% of full sunlight) were capable to accumulate similar amount of biomass than those grown under more illuminated environments by increasing their specific leaf area. Absolute differences in net photosynthesis between light treatments were higher in well watered than in water stressed seedlings. However, the impact of both factors on overall growth was orthogonal.

? We concluded that cork oak development is impaired under deep shade (5% of full sunlight) but it can be optimal under moderate shade (15% of full sunlight) even under moderate water stress. Implications of these patterns on regeneration, cultivation and afforestation of cork oak are discussed.

     

Waxes composition of reproduction cork from Quercus suber and its variability throughout the industrial processing

Summary The chemical composition of waxes was studied in cork planks from three different trees of Spanish Quercus suber at four different stages of the in‐dustrial processing of the first transformation: stripping, first rest, boiling fol‐lowed by open air rest and boiling followed by store-room rest. Waxes were extracted with chloroform (CHCl₃) and the extract was submitted to saponification in order to obtain the neutral and acid fractions. The trimethylsilyl derivatives of both fractions were analysed by GC-MS. An important decrease of the total content of waxes -CHCl₃ extract-(average values: 10.13 to 4.91%) and of the acid fraction (average values: 2.51 to 0.96%) was observed throughout the industrial processing. Some differences were also found among trees, two of them being richer in total waxes (average, 8.14 and 7.71%) than the other one (average, 4.67%). Concerning the three groups of identified components (fatty acids, fatty alcohols and triterpenes), the contents of the fatty alcohols and acids suffer an important reduction during the first rest (average values: 30.39 to 17.53% and 58.73 to 31.48%, respectively), while after boiling, there is a decrease of the triterpenes concentrations (79.16 to 58.00% -neutral fraction- and 40.23 to 23.20% -acid fraction-), being greater when the second rest is carried out in a store room. Significant differences among trees were only found in the fatty alcohols contents. The neutral fraction was mainly composed by fatty alcohols (all the even members from C₁₈ to C₂₆, with traces of intermediate odd members and some unsaturated groups, C₂₀ and C₂₁) and triterpenes (14 components, among them friedelin, betulin and cerin were identified), and a very small amount of monocarboxylic fatty acids (C₁₆, C₁₈, C₂₂ and C₂₄ members) was also present. The main group of components of the acid fraction was that of fatty acids (saturated even C₁₂–C₂₄ and odd C₁₅, C₁₇, C₂₁ members, accompanied by a great amount of unsaturated terms and some ω-hydroxyacids, 18-hydroxy-9,12-octadecadienoic and 18-hydroxy-9-octadecenoic acids). Four triterpenes, the C₂₀, C₂₄ and C₂₈ alcohols and ferulic acid were also detected in the acid fraction. Some of these individual components of each fraction were also affected by the industrial processing and presented significant differences among the studied trees. Received 12 December 1997     

东喜马拉雅地区高山栎类 (Quercus Sec.suber)种群变异和环境异质性研究(英文)

该文研究分布在我国东喜马拉雅地区 (云南和西藏 )及四川的高山栎类种群间的形态变异和遗传距离 .在自然条件下 ,这些栎类的生长形态随纬度的增加和降水的减少从高大的乔木变成矮小的灌木 .该地区多样的气候和复杂的地形条件为高山栎类提供了各种形态、生长和遗传变异的可能性 .种群内和种群间的遗传距离随生境条件、山地地形和主风方向的变化明显 ,但与水平距离关系不大 .西藏地区以地势险峻和气候多变为特色 ,云南地区以高山峡谷为特色 ,二地区高山栎的种群遗传距离分别为 0 396和 0 36 5,前者显著高于后者 (P <5% ) ,但却都高于四川地区的高山栎种群遗传距离 ,并达到极显著水平 (P <1% ) .环境的异质性越高 ,高山栎类的变异越大 ,种群变异和物种进化与气候、生境、地形地貌和土壤等环境的异质性密切相关 .尽管东喜马拉雅地区是以大陆性季风为主气候特征 ,但高山栎类的叶片、直径和根系对环境异质性的适应变异与分布在地中海地区的、冬雨夏旱气候条件下的冬青栎类 (Q .ilex)十分相似 .不同地区的环境异质性必然导致物种进化和种群变异的差异 ,然而 ,高山栎类种群在两个截然不同气候区内的高度相似性恰恰说明了它们可能来自于一个共同的祖先或从同一个地区演化而来 ,所以它们才有相同的适应策略     

Removal of nutrient limitations by long-term fertilization decreases nocturnal water loss in savanna trees

Under certain environmental conditions, nocturnal transpiration can be relatively high in temperate and tropical woody species. We have previously shown that nocturnal sap flow accounts for up to 28% of total daily transpiration in woody species growing in a nutrient-poor Brazilian Cerrado ecosystem. In the present study, we assessed the effect of increased nutrient supply on nocturnal transpiration in three dominant Cerrado tree species to explore the hypothesis that, in nutrient-poor systems, continued transpiration at night may enhance delivery of nutrients to root-absorbing surfaces. We compared nocturnal transpiration of trees growing in unfertilized plots and plots to which nitrogen (N) and phosphorus (P) had been added twice yearly from 1998 to 2005. Three independent indicators of nocturnal transpiration were evaluated: sap flow in terminal branches, stomatal conductance (g(s)), and disequilibrium in water potential between covered and exposed leaves (DeltaPsi(L)). In the unfertilized trees, about 25% of the total daily sap flow occurred at night. Nocturnal sap flow was consistently lower in the N- and P-fertilized trees, significantly so in trees in the N treatment. Similarly, nocturnal g(s) was consistently lower in fertilized trees than in unfertilized trees where it sometimes reached values of 150 mmol m(-2) s(-1) by the end of the dark period. Predawn gs and the percentage of nocturnal sap flow were linearly related. Nocturnal DeltaPsi(L) was significantly greater in the unfertilized trees than in N- and P-fertilized trees. The absolute magnitude of DeltaPsi(L) increased linearly with the percentage of nocturnal sap flow. These results are consistent with the idea that enhancing nutrient uptake by allowing additional transpiration to occur at night when evaporative demand is lower may avoid excessive dehydration associated with increased stomatal opening during the day when evaporative demand is high.     

Dieback of cork oak (Quercus suber) in Catalonia (NE Spain) caused by Botryosphaeria stevensii

Botryosphaeria stevensii Shoemaker (anamorph: Diplodia mutila Fr. apud Mont.) is reported as the cause of canker and dieback of cork oak (Quercus suber L.) in Catalonia (NE Spain). It also causes wilting of trees after cork is removed for industrial purposes. Symptomatology and details of morphology for both anamorph and teleomorph are given.     

Root system of Quercus suber L. seedlings in response to herbaceous competition and different watering and fertilisation regimes

Different management practices are commonly applied to increase pasture yield of Mediterranean open woodlands, but the consequences of increasing competition for soil resources with these practices on tree recruitment are still unknown. In a greenhouse study, belowground competition of Quercus suber L. seedlings growing together with natural (Oak_NP) or improved pasture (Oak_IP) was evaluated, and their root systems compared with those of seedlings growing in bare soil (Oak_BS). Two watering levels and two regimes of P₂O₅ fertilisation were also tested. Because of competition, the Oak_IP seedlings had their fine root mass density, coarse root length, and shoot mass reduced by up to 40, 36, and 39%, respectively, when compared to Oak_NP seedlings. Oak_NP and Oak_BS seedlings showed similar average root density parameters and shoot mass values, indicating that Q. suber seedlings growing with natural pasture is a low competitive system. High availability of water and phosphorus did not mitigate the strength of competition between herbaceous plants and oak seedlings, and favoured the pasture to the detriment of the trees. Our findings suggest that P₂O₅ fertilisation and irrigation practices performed to improve herbaceous productivity will negatively influence recruitment of Q. suber seedlings.