Genetic variation of pedunculate oak (Quercus robur L.) in the urban woodlands of Helsinki

We present a genetic profile of pedunculate oak (Quercus robur L.) in three woodlands of the Helsinki region. We genotyped all mature trees and samples of seedlings to evaluate patterns of genetic variation and gene flow, and evaluate the long-term stability and natural regeneration of these stands. Although five microsatellite loci showed no significant differences in heterozygosity between generations in most locations, significant (p?per se does not interfere with gene flow over the distances represented here. Based on our findings, a minimal-maintenance program reliant upon natural regeneration of oak in the public parks of Helsinki seems plausible.     

Crookedness of pedunculate oak (Quercus robur L.) growing under a canopy of Scots pine (Pinus sylvestris L.)

Natural regeneration of pedunculate oak growing under a canopy of Scots may be used for conversion into a mixed or predominantly broadleaved forest. From an economic perspective it is desirable that the oaks have straight stems. In this paper we present a case study for analysing oak stem curvature and its causes. The study was conducted in a mature pine stand with understorey oaks of age ca. 60 years in Poland. The oaks were classified into two main groups as either straight or crooked. The following variables were measured or assessed for the oaks: (1) stem curvature (crookedness), total tree height, diameter at breast height (DBH) and some tree crown characteristics, (2) the growth of DBH and height in each year of the life of each tree and (3) the competitive pressure of surrounding trees. Durable stem curvatures were formed at the culmination of the height increment. The first 20 years of the life of the oak were crucial in terms of stem quality. The annual height increment was larger in the years when the curvature was formed than in adjacent years. The factors that caused crooked stems were the same for straight and crooked oaks, but the magnitude of the response was different. When planning the conversion of Scots pine into oak, full overhead light should be provided as early as possible, but not later than 20 years since the establishment of the regeneration.     

Effects of pruning and stand density on the productionof new epicormic shoots in young stands of pedunculate oak (Quercus robur L.)

Context

Avoidance or control of epicormic shoots is among the major silvicultural challenges for the production of high-quality oak timber. In northern Europe, contemporary oak silviculture aims to produce valuable timber on a relatively short rotation, applying early, heavy thinning combined with artificial pruning.

Aims

The aim of this study was to analyse the effects of pruning and stand density on the production of new epicormic shoots on young trees of pedunculate oak (Quercus robur L.).

Methods

The study was based on two field experiments in even-aged stands of pedunculate oak subjected to different thinning practices and early selection of potential future crop trees. From ages 13 to 15 years, stem density was reduced to 300 trees ha^?1, 1,000 ha^?1 or stands remained unthinned. Pruning was conducted on selected trees at ages 22–24 years. At that age, the stem density in unthinned control plots ranged from 2,500 to 3,100 ha^?1. All treatments were replicated twice within each experiment.

Results

Pruning led to an overall increase in the total production of new epicormic shoots. More epicormic shoots were produced in the lower part of the stem (0–3 m in height) than in the upper part (3–6 m). The number of new epicormic shoots increased with increasing stand density.

Conclusion

Early, heavy thinning combined with high pruning at regular intervals may help shorten the rotation length for pedunculate oak without further reduction in wood quality than that which is caused by wider annual growth rings.     

Changes in water status and proline and abscisic acid concentrations in developing somatic embryos of pedunculate oak (Quercus robur) during maturation and germination

Somatic embryos of oak (Quercus robur L.) were matured on P24 media differing in gel strength (0.8, 0.9 and 1.0% (w/v) agar). Viscosity and osmotic potential (Psipi,medium) of the media were determined. Developing cotyledonary embryos were analyzed at maturity Stages I-III for water content, osmotic potential (Psipi,embryo) and concentrations of abscisic acid (ABA) and proline. Proliferation of embryogenic tissue, germination rates and the number of embryos formed were also determined in order to relate embryo quality to physiological parameters. Viscosity increased with agar concentration, a phenomenon apparently related to water availability. Many Stage III embryos with high germination potentials were obtained on P24 medium containing 1.0% agar. Embryo water content decreased progressively from 94 to 80% during embryo maturation. Stage I and II embryos that matured on media containing 0.8 or 0.9% agar had similar values of Psipi,embryo, whereas Psipi,embryo of Stage III embryos that matured on medium containing 1.0% agar was significantly lower, although Psipi,medium was unaffected by gel strength. Stage III embryos showed a nearly 16-fold increase in proline concentration and a 50% decrease in ABA concentration compared with Stage I embryos. We conclude that tissue water status and a complex relationship between ABA and proline concentrations, modulated by medium gel strength, are important factors in the maturation process and the quality of oak somatic embryos.     

Reduced photosynthesis in old oak (Quercus robur): the impact of crown and hydraulic architecture

We tested the hypothesis that changes in crown architecture of old pedunculate oak trees (Quercus robur L. ssp. robur Kl. et Kr. et Rol.) reduce leaf specific hydraulic conductance of shoots, thereby limiting stomatal conductance and assimilation of affected shoots. At the end of summer 1999, hydraulic conductance and leaf specific hydraulic conductance, measured with a high-pressure flow meter in 0.5- to 1.5-m long shoots, were 27 and 39% lower, respectively, in shoots of low vigor compared with vigorously growing shoots in a 165-year- old stand in southeastern Germany. Two types of bottlenecks to water transport can be identified in shoots of old oak trees, namely nodes and abscission zones. The reduction in hydraulic conductance was especially severe in shoots with diameters of less than 2 mm. Maximum stomatal conductance and maximum net assimilation rate increased significantly with hydraulic conductance and leaf specific hydraulic conductance. Our data support the hypothesis that changes in shoot and consequently crown architecture observed in aging trees can limit photosynthesis by reducing shoot hydraulic conductance. Thus, in addition to increasing pathway length and lower conductivity of xylem in old trees, structural changes in shoot and crown architecture need to be considered when analyzing water relations and photosynthesis in mature and declining trees.     

Intra- and interspecific diversity in the response to waterlogging of two co-occurring white oak species (Quercus robur and Q. petraea)

Quercus robur L. and Quercus petraea (Matt.) Liebl. are sympatric oak species with different ecological requirements. Quercus robur is more tolerant to waterlogging than Q. petraea. This ecological divergence may play a role in the maintenance of the two species despite the absence of an insurmountable reproductive barrier between them. We predicted that the genetic architecture of traits related to waterlogging tolerance differs between the species. To gain insight into this architecture in the absence of genetic markers for waterlogging tolerance, we compared populations of seedlings of each species for diversity in the expression of quantitative phenotypic traits induced by severe hypoxia. To determine the capacity for hypertrophied lenticel formation, we applied gas-impermeable mastic to stems of seedlings. Two months after application, the mastic treatment had induced the formation of 3 (+/- 2) cm(-2) hypertrophied lenticels in the absence of root hypoxia. Leaf epinasty during root hypoxia was an early predictor of seedling mortality. Four weeks of waterlogging resulted in greater epinasty in Q. petraea than in Q. robur, but fewer hypertrophied lenticels (16 +/- 6 versus 21 +/- 9 cm(-2)) and adventitious roots (2.7 +/- 4.7 versus 5.2 +/- 5.9). Differences between species in these traits were associated with differences in the frequencies of extreme phenotypes rather than with a generally higher tolerance to waterlogging in Q. robur seedlings.     

Genetic control of wood colour, density and heartwood ellagitannin concentration in European oak (Quercus petraea and Q. robur)

Pressler cores of young clones of Quercus petraea and Q. roburwere analysed from two sites in Germany. Variation of wood colour(defined by the CIELAB system), density and heartwood ellagitanninswas examined between clones, species and sites. Similar studieswere made of cores from another trial of 20 half-sib familiesof parent trees deriving from five German forests. The resultsfrom both the clonal and progeny trials indicate that heartwoodellagitannin content and wood density are under strong geneticcontrol, while sapwood and heartwood colour are less so. A largeproportion of between-clone variation in wood density and ellagitanninlevels occurred between the two species, while these propertiesvaried little among ramets of the same clone grown on two contrastingsites, despite a difference in growth rates. A large proportionof the total variation among progeny was attributed to forestorigins and could not be attributed clearly to either geneticor environmental causes because of limitations of the samplingdesign     

Visitor preferences of thinning practice in young even-aged stands of pedunculate oak (Quercus robur L.): comparing the opinion of forestry professionals in six European countries

This study compared visitor preferences of forestry professionals across six European countries (Sweden, Denmark, Great Britain, Austria, Romania and Portugal) using a questionnaire survey. The 598 interviewees were asked to rank photographs depicting recently thinned experimental plots in a 13-year old stand of pedunculate oak (Quercus robur L.) according to the criterion: “Which forest environment do you prefer as a visitor?” The plots represented five different residual stem densities: 7000 (no thinning, very high stem density), 5300 (heavy thinning, high stem density), 1000 (very heavy thinning, medium stem density), 300 (extremely heavy thinning, low stem density/open stand) and 100 (solitary trees, very low stem density/very open stand) stems ha^?1. The results indicated geographical variation in the preferences for different thinning practices in young stands of oak. Portuguese, Austrian and Romanian respondents generally favoured thinned, but dense stands, whereas Danish and British respondents preferred very heavily thinned stands. Swedish respondents preferred open stands resulting from extremely heavy thinning. Photographs taken along rows were favoured to photographs across rows, indicating a preference for scenes offering perspective and accessibility. The results indicate a variation of visitor preferences among forestry professionals for different silvicultural regimes. We interpret this in the context of national traditions and forestry paradigms that influence the shaping of preferences.     

Low temperature during winter elicits differential responses among populations of the Mediterranean evergreen cork oak (Quercus suber)

Populations of cork oak (Quercus suber L.) were assessed for seasonal and inter-population variability in, and temperature responses of, the ratio between light-induced variable and maximum fluorescence of chlorophyll, Fv/Fm, considered a surrogate for the maximum photochemical efficiency of photosystem II (PSII). Seedlings from 10 populations throughout the distribution range of Q. suber in the Mediterranean basin were grown in a common garden in central Spain. The Fv/Fm ratio of dark-adapted leaves was measured at dawn every month for 2 years. Air temperature was recorded at a nearby climatic station. During the summer, when maximum air temperatures reached 40 degrees C, there were no significant differences in Fv/Fm among populations, but significant differences were seen during the winter. In colder months, Fv/Fm ranged in all populations between 0.5-0.6 and 0.2-0.3 in 2001 and 2002, respectively. The variance explained by the population effect was greatest during winter months, especially in 2002, reaching a peak value of 10% when minimum air temperature was below -10 degrees C. Populations originating from warmer sites showed the largest decline in Fv/Fm between the end of 2001 and the beginning of 2002. Thus, a negative linear relationship was established between mean annual temperature at the population source and population mean Fv/Fm recorded in the coldest month in 2002 and normalized by the Fv/Fm spring measurement.     

Genetic diversity and differentiation of fall webworm (Hyphantria cunea Drury) populations

The molecular variation and genetic relationships among five populations of the fall webworm (Hyphantria cunea Drury) in China were assessed using microsatellite technology. Ten microsatellite primers, producing polymorphic bands, were used across 300 samples. The percentage of polymorphic loci (PPB) was 98.36%; the percentage of polymorphic loci in five populations ranged from high to low in the following order: Cangzhou population, Yantai population, Qinhuangdao population, Dandong population, and Shijiaz...     

Genetic diversity and differentiation of fall webworm (Hyphantria cunea Drury) populations

The molecular variation and genetic relationships among five populations of the fall webworm (Hyphantria cunea Drury) in China were assessed using microsatellite technology. Ten microsatellite primers, producing polymorphic bands, were used across 300 samples. The percentage of polymorphic loci (PPB) was 98.36%; the percentage of polymorphic loci in five populations ranged from high to low in the following order: Cangzhou population, Yantai population, Qinhuangdao population, Dandong population, and Shijiazhuang population. The results showed that 34.38% of the total genetic variation of the fall webworm (GST) occurs among populations, while most variation (65.62%) exists within populations. Nei’s genetic distances ranged from 0.1386 to 0.3224. Using the unweighted pair group method with arithmetic mean (UPGMA), Nei’s genetic distances were analyzed by a clustering technique and the dendrogram shows that population differentiation is closely related to the time and geographic origin of the invasion. The major factors impacting genetic diversity of fall webworm populations are longitude, the plain area ratio, annual precipitation, latitude and time of invasion. The formation of genetic structure is correlated with characteristics of the life history and invasion ecology of the species.     

Interactive effects of elevated CO2 concentration and nitrogen supply on partitioning of newly fixed 13C and 15N between shoot and roots of pedunculate oak seedlings (Quercus robur)

Pedunculate oak (Quercus robur L.) seedlings were grown for 3 or 4 months (second- and third-flush stages) in greenhouses at two atmospheric CO2 concentrations ([CO2]) (350 or 700 micromol mol(-1)) and two nitrogen fertilization regimes (6.1 or 0.61 mmol N l(-1) nutrient solution). Combined effects of [CO2] and nitrogen fertilization on partitioning of newly acquired carbon (C) and nitrogen (N) were assessed by dual 13C and 15N short-term labeling of seedlings at the second- or third-flush stage of development. In the low-N treatment, root growth, but not shoot growth, was stimulated by elevated [CO2], with the result that shoot/root biomass ratio declined. At the second-flush stage, overall seedling biomass growth was increased (13%) by elevated [CO2] regardless of N fertilization. At the third-flush stage, elevated [CO2] increased growth sharply (139%) in the high-N but not the low-N treatment. Root/shoot biomass ratios were threefold higher in the low-N treatment relative to the high-N treatment. At the second-flush stage, leaf area was 45-51% greater in the high-N treatment than in the low-N treatment. At the-third flush stage, there was a positive interaction between the effects of N fertilization and [CO2] on leaf area, which was 93% greater in the high-N/elevated [CO2] treatment than in the low-N/ambient [CO2] treatment. Specific leaf area was reduced (17-25%) by elevated [CO2], whereas C and N concentrations of seedlings increased significantly in response to either elevated [CO2] or high-N fertilization. At the third-flush stage, acquisition of C and N per unit dry mass of leaf and fine root was 51 and 77% greater, respectively, in the elevated [CO2]/high-N fertilization treatment than in the ambient [CO2]/low-N fertilization treatment. However, there was dilution of leaf N in response to elevated [CO2]. Partitioning of newly acquired C and N between shoot and roots was altered by N fertilization but not [CO2]. More newly acquired C and N were partitioned to roots in the low-N treatment than in the high-N treatment.     

Patterns of stomatal conductance among blue oak (Quercus douglasii) size classes and populations: implications for seedling establishment

Quercus douglasii Hook. & Arn. (blue oak) is a deciduous white oak that is currently failing to regenerate throughout much of its range in California, USA. Patterns of water use were observed in adult trees, saplings and seedlings to determine if ontogenetic changes in water use occur, which might be important in the establishment of this long-lived perennial species in a Mediterranean-type system. Seasonal and diurnal stomatal conductance (g(s)), late-season predawn xylem water potentials (Psi(pre)), carbon isotopic ratio (delta(13)C) and soil water status were compared among the three size classes at three sites differing in mean precipitation and soil water characteristics. Comparisons were also made between microsites with and without regeneration (defined by the presence or absence of saplings). Overall patterns of water use were consistent among the three sites, except that, at the site with the highest rainfall, Q. douglasii plants had higher g(s) and more positive Psi(pre) values. Although no differences in water use patterns were found between regeneration and non-regeneration microsites, the observed ontogenetic differences in water use may have important implications for Q. douglasii establishment. Compared with adult trees and saplings, seedlings had higher gas exchange rates during periods of high soil water content (early in the season and in the morning). Seedling g(s) was correlated with percent extractable soil water (ESW) throughout the season; adult tree and sapling g(s) was correlated with ESW between June and September. Despite experiencing greater water stress (indicated by more negative Psi(pre) values) than older trees, seedlings had more negative delta(13)C values, implying lower water-use efficiencies.     

Drought and the diurnal patterns of stem CO2 efflux and xylem CO2 concentration in young oak (Quercus robur)

A young potted oak (Quercus robur L.) tree was subjected to drought by interrupting the water supply for 9 days. The tree was placed in a growth chamber in which daily patterns of temperature and radiation were constant. The effects of drought on the water and carbon status of the stem were examined by measuring stem sap flow rate, stem water potential, stem diameter variations, stem CO(2) efflux rate (F(CO2)) and xylem CO(2) concentration ([CO(2)*]). Before and after the drought treatment, diurnal fluctuations in F(CO2) and [CO(2)*] corresponded well with variations in stem temperature (T(st)). Daytime depressions in F(CO2) did not occur. During the drought treatment, F(CO2) still responded to stepwise changes in temperature, but diurnal fluctuations in F(CO2) were no longer correlated with diurnal fluctuations in T(st). From the moment daily growth rate of the stem became zero, diurnal fluctuations in F(CO2) became closely correlated with diameter variations, exhibiting clear daytime depressions. The depressions in F(CO2) were likely the result of a reduction in metabolic activity caused by the lowered daytime stem water status. Xylem [CO(2)*] showed clear daytime depressions in response to drought. When the tree was re-watered, F(CO2) and [CO(2)*] exhibited sharp increases, coinciding with an increase in stem diameter. After resumption of the water supply, daytime depressions in F(CO2) and [CO(2)*] disappeared and diurnal fluctuations in F(CO2) and [CO(2)*] corresponded again with variations in T(st).     

Recreational preferences depending on thinning practice in young even-aged stands of pedunculate oak (Quercus robur L.): comparing the opinions of forest and landscape experts and the general population of Denmark

The aim of this study was to investigate to which extent forestry professionals, other “green” professionals (mainly landscape architects) and natural resource students in Denmark agree with the general population regarding their recreational preferences for contrasting stem densities in young, even-aged stands of pedunculate oak (Quercus robur L.). The study was based on five pairs of colour photographs from each of the five recently thinned experimental plots in a 13-year-old stand. The plots represented five different residual stem densities: 7000 (unthinned), 5300 (traditional thinning), 1000, 300 and 100 stems ha^?1. All cut trees were left on the ground. The analysis was carried out as a survey of forest and landscape experts (n=158) and the general public (n=243) based on questionnaires. Interviewees ranked the photographs according to the criterion: “Which forest environment do you prefer as a visitor?” Principal component factor analysis showed that the general public tended to perceptually group photos according to similar overall pattern of openness, presence of row structure and stand accessibility (including presence/absence of slash). As long as a stand appeared accessible, stand density had little influence on their preferences for a wide range of stand densities (5300?300 stems ha^?1). In contrast, foresters, other “green” professionals and students seemed to perceptually group photos according to treatment type, and the presence of slash had little influence on their preferences. This suggests an influence of professional background, making experts visually analyse and evaluate stands according to contemporary management standards and paradigms within their own profession.     

Decline of genetic diversity of sessile oak at the retracting (xeric) limits

Context

Genetic diversity of sessile oak (Quercus petraea) populations in Hungary was assessed close to the retracting, low-elevation, low-latitude (xeric) distribution limits.

Objective

We aimed at tracing an assumed effect of climatic factors on genetic diversity, particularly at the southern, low elevation limits of distribution.

Methods

Genetic diversity at isozyme-coding loci was analysed in populations, and related to the climate of the sites where the populations grow. A locus-wise analysis proved to be essential to follow responses.

Results

A climate-related cline was found at seven isoenzyme-coding gene loci. Declining allelic numbers and heterozygosity indicated lower diversity at warmer and drier sites. The majority of loci were responsive to precipitation factors, others to temperature. Genetic clustering was neither related to geographic distance nor to random or historic effects.

Conclusions

The results suggest that climatic stress may elicit a genetic diversity loss in populations, which may reduce their plasticity and adaptive potential. The selective pressure may override historic effects and gene flow. With respect to expected climate change, the correlation of diversity with some climatic factors gains specific importance. If supported by further investigations, the results might be utilised for reconsidering conservation strategies and rules for use of forest reproductive material.     

High-pruning of European beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.): work efficiency for target pruning as a function of tree species,pruning height,branch characteristics,pole saw type and operator

The objective of this study was to establish an operational model of productive work time per tree (work efficiency) for high-pruning of young European beech and pedunculate oak depending on tree species, pruning height, branch characteristics, pole saw type and operator. The final model included all of these independent variables with branch characteristics specified in terms of number of live branches and cross-sectional area of the thickest branch at the cut. Work time increased with increasing values of each of the three numeric variables. For a given pruning height the size of the largest branch was for all practical purposes more influential than the number of live branches. Beech took 28% longer to prune than oak. The German Ergo-Schnitt saw was 21% slower than the Japanese Silky Hayauchi saw. The variation in worker performance within our study was larger than that attributed to tree species and pruning equipment.     

Low genetic diversity and high genetic differentiation among severely fragmented populations of the critically endangered tree Talbotiella gentii (Fabaceae)

The genetic diversity among 17 populations (including 10 newly discovered) of Talbotiella gentii (Fabaceae, Caesalpinioideae), a rare tree species endemic to Ghana, was assessed using random amplified polymorphic DNA (RAPD). A total of 80 polymorphic bands were generated from eight primers and analysed by cluster and correspondence analysis. This showed a clear distinction between populations, with two major groups associated with geographical origin. AMOVA analysis showed that 94.05% of the genetic variation was partitioned among populations, while only 5.95% was found within populations suggesting little gene flow between populations. Possible reasons for the high population genetic differentiation and the low levels of genetic variation within populations are inbreeding and genetic drift. Of a total of 26 known populations, 14 are now extinct, five during the course of this study. Action to prevent complete extinction of the species is therefore urgent.     

Variation in the radial patterns of sap flux density in pubescent oak (Quercus pubescens) and its implications for tree and stand transpiration measurements

Radial variation in sap flux density across the sapwood was assessed by the heat field deformation method in several trees of Quercus pubescens Wild., a ring-porous species. Sapwood depths were delimited by identifying the point of zero flow in radial patterns of sap flow, yielding tree sapwood areas that were 1.5-2 times larger than assumed based on visual examinations of wood cores. The patterns of sap flow varied both among trees and diurnally. Rates of sap flow were higher close to the cambium, although there was a significant contribution from the inner sapwood, which was greater (up to 60% of total flow) during the early morning and late in the day. Accordingly, the normalized difference between outer and inner sapwood flow was stable during the middle of the day, but showed a general decline in the afternoon. The distribution of sap flux density across the sapwood allowed us to derive correction coefficients for single-point heat dissipation sap flow measurements. We used daytime-averaged coefficients that depended on the particular shape of the radial profile and ranged between 0.45 and 1.28. Stand transpiration calculated using the new method of estimating sapwood areas and the radial correction coefficients was similar to (Year 2003), or about 25% higher than (Year 2004), previous uncorrected values, and was 20-30% of reference evapotranspiration. We demonstrated how inaccuracies in determining sapwood depths and mean sap flux density across the sapwood of ring-porous species could affect tree and stand transpiration estimates.