Carbon isotope discrimination and wood anatomy variations in mixed stands of Quercus robur and Quercus petraea

The two most common oak species in western Europe, Quercus robur and Quercus petraea, display different ecological behaviours, particularly with respect to their responses to drought. The ecophysiological basis of this niche difference is not understood well. Here we test the hypothesis that these two species present distinct water use efficiencies (WUEs), using the carbon isotope discrimination approach. Leaves and 13 dated ring sequences were sampled in 10 pairs of adult trees growing side by side. Carbon isotope composition was measured on cellulose extracts. In addition, relationships between carbon isotope discrimination and wood anatomy were assessed at the tree level. Quercus robur displayed a 1·0‰ larger isotopic discrimination than Q. petraea, and therefore a lower intrinsic WUE (?13%). This interspecific difference of isotopic discrimination was quite stable with time and independent of tree radial growth and climate fluctuations. A strong positive correlation was observed between average tree values of earlywood vessel surface area and ¹³C isotopic discrimination. This correlation was even higher with ¹³C of the 1976 dry year (r = 0·86). These observations led to the hypothesis that hydraulic properties of xylem could exert a constraint on leaf gas exchange, resulting in a larger WUE for individuals with smaller vessel cross‐section area.     

Three divergent rDNA clusters predate the species divergence in Quercus petraea (Matt.) Liebl. and Quercus robur L

Quercus petraea and Quercus robur are two closely related oak species that frequently hybridize. We sequenced 70 clones containing the 5.8S and ITS2 regions of ribosomal DNA (rDNA) from these two species and did not detect a species-specific difference. Surprisingly, three divergent (up to 12.6%) rDNA families were identified in both species, indicating that they predate the speciation event. Despite a large between-rDNA-families divergence, rDNA sequences were very similar within families, suggesting ongoing concerted evolution. Expression analysis, relative-rate tests, and mutation spectrum analyses indicated that only a single rDNA family is functional. We propose that past hybridization events, combined with nucleolar dominance, were the evolutionary processes underlying the contemporary rDNA variability in Q. petraea and Q. robur.     

Characterisation and natural variation of a dehydrin gene in Quercus petraea (Matt.) Liebl

For the first time in sessile oak [Quercus petraea (Matt.) Liebl.], the isolation and characterisation of a full-length dehydrin gene and its promoter region, as well as its allelic variation in natural populations, is reported. Dehydrins (Dhn) are stress-related genes important for the survival of perennial plants in a seasonal climate. A full-length dehydrin gene (Dhn3) was characterised at the nucleotide level and the protein structure was modelled. Additionally, the allelic variation was analysed in five natural populations of Quercus petraea (Matt.) Liebl. sampled along an altitudinal gradient in the French Pyrenees. The analysed sequences contain typical domains of the K(n) class of dehydrins in the coding region. Also, the 5'untranslated region (promoter) of the gene was amplified, which shows typical motifs essential for drought- and cold-responsive gene expression. Single nucleotide substitutions and indels (insertions/deletions) within the coding region determine large biochemical differences at the protein level. However, only low levels of genetic differentiation between populations from different altitudes were detectable.     

Regeneration Patterns of European Oak Species (Quercus petraea (Matt.) Liebl., Quercus robur L.) in Dependence of Environment and Neighborhood

Quercus robur L. (pedunculate oak) and Quercus petraea (Matt.) Liebl. (sessile oak) are two European oak species of great economic and ecological importance. Even though both oaks have wide ecological amplitudes of suitable growing conditions, forests dominated by oaks often fail to regenerate naturally. The regeneration performance of both oak species is assumed to be subject to a variety of variables that interact with one another in complex ways. The novel approach of this research was to study the effect of many ecological variables on the regeneration performance of both oak species together and identify key variables and interactions for different development stages of the oak regeneration on a large scale in the field. For this purpose, overstory and regeneration inventories were conducted in oak dominated forests throughout southern Germany and paired with data on browsing, soil, and light availability. The study was able to verify the assumption that the occurrence of oak regeneration depends on a set of variables and their interactions. Specifically, combinations of site and stand specific variables such as light availability, soil pH and iron content on the one hand, and basal area and species composition of the overstory on the other hand. Also browsing pressure was related to oak abundance. The results also show that the importance of variables and their combinations differs among the development stages of the regeneration. Light availability becomes more important during later development stages, whereas the number of oaks in the overstory is important during early development stages. We conclude that successful natural oak regeneration is more likely to be achieved on sites with lower fertility and requires constantly controlling overstory density. Initially sufficient mature oaks in the overstory should be ensured. In later stages, overstory density should be reduced continuously to meet the increasing light demand of oak seedlings and saplings.     

Water relations and gas exchange in Fagus sylvatica L. and Quercus petraea (Mattuschka) Liebl. in a mixed stand at their southern limit of distribution in Europe

Water status and gas exchange of beech (Fagus sylvatica L.) and sessile oak [Quercus petraea (Mattuschka) Liebl.] were studied in a mixed stand in the Montejo de la Sierra forest (central Iberian Peninsula), one of the southernmost locations of both species in Europe. Gas exchange and water potential were measured in leaves at different canopy levels over several days in two growing seasons. The daily variation pattern was established with the measurements of three selected dates per year, representative of the soil moisture content situations in early, mid- and late summer. A similar daily time course of leaf water potential was found for the two species. Nevertheless, beech showed a most noticeable decrease of water potential at midmorning and maintained lower leaf water potential than oak in the early afternoon. In 1994 the sessile oak saplings showed higher values of predawn water potential (Ψ_pd) than beech at the end of summer, when soil moisture content was lowest (20 cm depth). Beech showed a significantly lower net assimilation rate (A) than sessile oak for leaves under the same PPFD. Maximum net photosynthesis values (A _max) for beech and sessile oak on sunny leaves were 10.1±0.4 μmol m^–2 s^–1 and 17.8±1.7 μmol m^–2 s^–1 respectively, and those for water vapour stomatal conductance (g _wv) were 265±31 mmol m^–2 s^–1 and 438±74 mmol m^–2 s^–1. Differences in A and g _wv between the two species were maintained throughout the day on all measurement dates. No clear relationship was found between water status of saplings and stomata performance; there was only a negative correlation between Ψ_pd and g _wvmid in beech. Nevertheless, a significant response to the air vapour pressure gradient between leaf and air was translated into stomata closure on an hourly basis, more intensively in beech. Received: 4 March 1999 / Accepted: 21 December 1999     

Species distinction in Irish populations of Quercus petraea and Q. robur: morphological versus molecular analyses

BACKGROUND AND AIMS: Populations of oak (Quercus petraea and Q. robur) were investigated using morphological and molecular (AFLP) analyses to assess species distinction. The study aimed to describe species distinction in Irish oak populations and to situate this in a European context. METHODS: Populations were sampled from across the range of the island of Ireland. Leaf morphological characters were analysed through clustering and ordination methods. Putative neutral molecular markers (AFLPs) were used to analyse the molecular variation. Cluster and ordination analyses were also performed on the AFLP markers in addition to calculations of genetic diversity and F-statisitcs. KEY RESULTS: A notable divergence was uncovered between the morphological and molecular analyses. The morphological analysis clearly differentiated individuals into their respective species, whereas the molecular analysis did not. Twenty species-specific AFLP markers were observed from 123 plants in 24 populations but none of these was species-diagnostic. Principal Coordinate Analysis of the AFLP data revealed a clustering, across the first two axes, of individuals according to population rather than according to species. High F(ST) values calculated from AFLP markers also indicated population differentiation (F(ST) = 0.271). Species differentiation accounted for only 13 % of the variation in diversity compared with population differentiation, which accounted for 27 %. CONCLUSIONS: The results show that neutral molecular variation is partitioned more strongly between populations than between species. Although this could indicate that the populations of Q. petraea and Q. robur studied may not be distinct species at a molecular level, it is proposed that the difficulty in distinguishing the species in Irish oak populations using AFLP markers is due to population differentiation masking species differences. This could result from non-random mating in small, fragmented woodland populations. Hybridization and introgression between the species could also have a significant role.     

Identification of triterpene saponins in Quercus robur L. and Q. petraea Liebl. Heartwood by LC-ESI/MS and NMR

A detailed structural study of the major acidic triterpene saponins from European oak heartwood, Quercus robur L. and Q. petraea Liebl., revealed the presence of 2 alpha,3 beta,19 alpha-trihydroxyolean-12-ene-24,28-dioic acid (1), 2 alpha,3 beta,19 alpha-23-tetrahydroxyolean-12-ene-24,28-dioic acid (2), and their corresponding glycosides, 28-beta-D-glucopyranosyl-2 alpha,3 beta,19 alpha-trihydroxyolean-12-ene-24,28-dioic acid (3), and 28-beta-D-glucopyranosyl-2 alpha,3 beta,19 alpha,23-tetrahydroxyolean-12-ene-24,28-dioic acid (4). Compounds were isolated and purified by successive flash chromatography and semi-preparative HPLC, and structurally determined by NMR and LC-ESI/MS in the negative ion mode. Compounds 2 and 4 have been identified for the first time in Quercus species and are new compounds.     

Effect of indole-3-acetic acid (IAA) and sucrose on vessel size and density in isolated stem segments of oak (Quercus robur)

The effects of several concentrations of indole-3-acetic acid (IAA) and sucrose on xylogenic cambial activity and secondary xylem differentiation were investigated in isolated stem segments of Quercus robur L. supplied with liquid medium in aseptic conditions. After 5 weeks of culture auxin controlled cambial cell division and the number and size of vessel elements even without sugar in the medium. Sucrose modified these IAA effects, although little cambial activity occurred without auxin. The xylem increment correlated with changes of auxin concentration with the optimum at 28.5 μ M IAA. The formation of wide vessels was correlated with the optimal concentration of auxin. The frequency of vessel differentiation increased with auxin concentration. High concentrations of sucrose (0.24 M and 0.96 M ) reduced both the number of vessels and their diameter. The frequency of vessel formation was inhibited more than the vessel size by changes of sugar concentration. The vessels formed under low concentrations of IAA were circular in transverse section. With increase in IAA concentration the shape of the vessel cross-section changed to oval with the largest dimension in the radial direction.     

AFLP markers demonstrate local genetic differentiation between two indigenous oak species [ Quercus robur L. and Quercus petraea (Matt.) Liebl.] in Flemish populations

The nuclear genetic variation within and between four sessile ( Q. petraea) and six pedunculate ( Q. robur) autochthonous Flemish oak populations was investigated with AFLP markers. One sessile and one pedunculate oak population were additionally screened for detailed leaf characteristics using an image analysis system. Principal coordinate analysis on the AFLP data classified the oaks in two main groups, according to their taxonomic status. No species-specific AFLP markers were found using four primer combinations, but marker frequency differences up to 71% were recorded between both species. Analysis of the genetic structure showed that the divergence between species, as observed by ordination, was significant. Both species revealed similar diversity levels. A smaller though significant differentiation was also revealed for both species among populations within species. Molecular and morphology based approaches showed a high degree of consistency. Screening of 60 AFLP primer combinations using a bulking strategy did not allow identifying species-specific markers, which supports the conclusions reached in previous studies. The distribution of genetic variability at the species and at the population level is discussed.     

Pollen dispersal inferred from paternity analysis in a mixed oak stand of Quercus robur L. and Q. petraea (Matt.) Liebl.

Paternity analysis was used to determine the spatial distribution of male parents of 984 offspring collected from 13 identified mother trees in a natural stand of 5.76 ha and comprising 296 adult trees of Quercus petraea and Q . robur . For seven of the 13 maternal progeny arrays sampled, we found an excess of nearby matings and a preferential direction of pollination. For the remaining progeny arrays, no departure from random distribution of male parents was detected. A common trend among all families was a high percentage (averaging 65% for Q. robur and 69% for Q. petraea ) of offspring that were pollinated by male parents from outside the study site. By pooling the data over all families, the average pollen dispersal curve within the stand was inferred and fitted to a negative exponential distribution. This model extrapolated for distances over the spatial scale of the study stand was insufficient to explain the high level of gene flow detected by the paternity analysis, suggesting a substantial level of long-distance pollination events. The genetic composition of the pollen pools received by each maternal tree was compared and showed significant differentiation that could be attributed to differences in male reproductive success. By contrast, no significant differentiation between the pollen clouds from outside and inside the study stand was detected, suggesting genetic homogeneity between the surrounding forest and the study stand.     

Morphological and physiological reactions of young deciduous trees (Quercus robur L., Q. petraea [Matt.] Liebl., Fagus sylvatica L.) to waterlogging

One-year-old seedlings of Quercus robur L., Q. petraea (Matt.) Liebl. and Fagus sylvatica L. were cultivated in lysimeters and subjected to waterlogging for 17 weeks, interrupted by a five-week drainage period during summer. The growth of Q. robur was less affected by waterlogging than that of Q. petraea and Fagus. Waterlogging resulted in the formation of adventitious roots in Q. robur and Q. petraea, but not in Fagus. In contrast to Fagus, Q. robur and, to a lesser extent, Q. petraea were able to generate roots even below the water table. The hydraulic conductance of the excised root systems, the stomatal conductance and, in Fagus, the leaf water potential and the leaf-mass related hydraulic conductance were decreased by waterlogging. The decrease in the hydraulic conductance was largest in Fagus, and smallest in Q. robur. The roots of Fagus responded to anaerobic conditions with an increase in ethanol concentration. The measurements of nitrate reductase activities in roots and leaves provided no indications of a persistent contribution of NO₃ ⁻ metabolism to the alleviation of waterlogging-induced stress. It is concluded that Q. robur and, to a lesser extent, Q. petraea can tolerate waterlogging periods better than Fagus due to a different pattern of root formation, and to a better adjustment of leaf biomass production to the hydraulic conductivity of the root system. This revised version was published online in June 2006 with corrections to the Cover Date.     

Epicormic ontogeny in Quercus petraea constrains the highly plausible control of epicormic sprouting by water and carbohydrates

Background and Aims

There is increasing evidence that suppressed bud burst and thus epicormic shoot emergence (sprouting) are controlled by water–carbohydrate supplies to entire trees and buds. This direct evidence is still lacking for oak. In other respects, recent studies focused on sessile oak, Quercus petraea, have confirmed the important constraints of sprouting by epicormic ontogeny. The main objective of this paper was thus to provide provisional confirmation of the water–carbohydrate control and direct evidence of the ontogenic constraints by bringing together results already published in separate studies on water status and distribution of carbohydrates, and on accompanying vegetation and epicormics, which also quantify epicormic ontogeny.

Methods

This paper analyses results gained from a sessile oak experiment in which part of the site was free from fairly tall, dense accompanying vegetation. This experiment was initially focused on stand water status and more recently on the carbohydrate distribution of dominant trees. External observations of the epicormic composition and internal observations with X-ray computer tomography were undertaken on 60 and six trees, respectively.

Key Results

Sprouting was more intense in the part of the stand free from accompanying vegetation and on upper trunk segments. A clear effect of epicormic ontogeny was demonstrated as well: the more epicormics a trunk segment bears, the more chances it had to bear sprouts.

Conclusions

These results indirectly infer water–carbohydrate control and show direct evidence of constraints by epicormic ontogeny. These results have far-reaching consequences related to the quantification of all functions fulfilled by any type of epicormic structure in any part of the tree.     

The effects of liming and gypsum applications on a sessile oak (Quercus petraea (M.) Liebl.) stand at La Croix-Scaille (French Ardennes) I. Site characteristics, soil chemistry and aerial biomass

In a former 45 to 50 year old sessile oak ( Quercus petraea (M.) Liebl.) coppice mixed with birch (Betula pubescens Ehrh.) and rowan (Sorbus aucuparia L.) on a poor acidic forest soil at la Croix-Scaille in the French Ardennes, several liming amendments were applied in 1990 and 1994. Data on soil and soil solution composition, as well as stand growth and foliar composition were collected between 1994 and 1997. All treatments, containing 1.4 t ha-1 equivalent of CaO supplied as lime, gypsum or a mixture of the two, resulted in an increase of cation exchange capacity and base saturation down to 15 cm and for CaSO4 treatments down to 30 to 45 cm, increases of soil pH and Ca concentration at the surface and a decrease of Al concentration in the soil and soil solution in the surface layers. No negative effects like increased nitrate or cation leaching were observed. Although Mg nutrition was not improved by the treatments (not containing Mg), a relative and maintained gain of radial increment of sessile oak in the order of 40% for both lime and gypsum applied, was observed immediately from the first year on, after the application (1991).     

Long-term changes in wood density and radial growth of Quercus petraea Liebl. in northern France since the middle of the nineteenth century

Long-term changes in sessile oak (Quercus petraea Liebl.) growth and wood density were studied using cores collected from 99 even-aged high forest stands between 56 and 187 years old, located in northeastern and north-central France. Growth and density trends were tested by analysis of variance and covariance. Two models were applied to two samples, sample A and sample B (sample B being a sub-sample with limited cambial age and calendar date ranges). Model 1 showed a significant increase in radial growth: +35%, +87% and +66% in earlywood width, latewood width and ring width, respectively, from 1811 to 1993 for sample A. Consequently, there was a positive trend in latewood ratio (+14%). A slight decrease in wood density was found: -3.3% and -5.4% for earlywood and latewood density, respectively. Despite an increase in latewood percentage, mean ring density showed a -2.0% decrease. Model 1 applied to a biomass indicator (density2ring width) showed a 62% increase from 10.4 to 16.8 kg m^-3 between 1811 and 1993 for sample A. Results for sample B were slightly different: the increase in latewood ratio was not detected. Model 2 showed a change with time in the positive hyperbolic relationship between mean density and ring width. The results are discussed. The decrease in wood density cannot be explained by N atmospheric deposition or by long-term changes in average temperature. Increasing atmospheric CO₂ levels cannot be invoked owing to the present lack of studies. Finally, hypotheses concerning long-term changes in wood anatomical characteristics are proposed.     

Realized gene flow within mixed stands of Quercus robur L. and Q. petraea (Matt.) L. revealed at the stage of naturally established seedling

The estimates of contemporary gene flow assessed based on naturally established seedlings provide information much needed for understanding the abilities of forest tree populations to persist under global changes through migration and/or adaptation facilitated by gene exchange among populations. Here, we investigated pollen‐ and seed‐mediated gene flow in two mixed‐oak forest stands (consisting of Quercus robur L. and Q. petraea [Matt.] Liebl.). The gene flow parameters were estimated based on microsatellite multilocus genotypes of seedlings and adults and their spatial locations within the sample plots using models that attempt to reconstruct the genealogy of the seedling cohorts. Pollen and seed dispersal were modelled using the standard seedling neighbourhood model and a modification—the 2‐component seedling neighbourhood model, with the later allowing separation of the dispersal process into local and long‐distance components. The 2‐component model fitted the data substantially better than the standard model and provided estimates of mean seed and pollen dispersal distances accounting for long‐distance propagule dispersal. The mean distance of effective pollen dispersal was found to be 298 and 463 m, depending on the stand, while the mean distance of effective seed dispersal was only 8.8 and 15.6 m, which is consistent with wind pollination and primarily seed dispersal by gravity in Quercus. Some differences observed between the two stands could be attributed to the differences in the stand structure of the adult populations and the existing understory vegetation. Such a mixture of relatively limited seed dispersal with occasional long distance gene flow seems to be an efficient strategy for colonizing new habitats with subsequent local adaptation, while maintaining genetic diversity within populations.     

Porosity and available water of temporarily waterlogged soils in a Quercus robur (L.) declining stand

Pedunculate oak (Quercus robur L.) is particularly sensitive to decline in clayey soils presenting a high-perched temporary water table. These soils induce two successive constraints in one-year cycle: water excess (and hypoxy) in winter and early spring, and water shortage in summer (water stress being more restrictive to oak). We determined the porosity and water properties of temporarily waterlogged clayey soils supporting forest stands of decliningQuercus robur trees in a 101yr-old oak stand in Belgium (50°06N, 4°16E). Roots unevenly colonized the soil down to 1.6m: fine roots (diameter<5mm) were mostly distributed on the surface horizons (0–0.3 m) and around 1.3m deep, despite dense clayey horizons appearing at 0.35m depth. Clay content below this depth reached 46–51. Illite and vermiculite were the dominant clay minerals. The clayey horizons exhibited marked shrink–swell properties: bulk density at 30kPa increased from 1.41 to 1.88gcm^–3 from the surface to 2m depth. There was also evidence of hypoxic conditions caused by water saturation of pore space in the rooting zone from October to mid-April. Extractable water (EW), calculated between moisture tensions of 5 and 1600kPa was 152.8mm. The level of perched temporary water table strongly depended on the seasonal rhythm of water uptake by trees and on the shrink–swell behaviour of soil.     

Genetic variation at allozyme and RAPD loci in sessile oak Quercus petraea (Matt.) Liebl.: the role of history and geography

The nuclear genetic variation within and among 21 populations of sessile oak was estimated at 31 RAPD loci in conjunction with previous estimates of variation at eight allozyme loci. The aim of the study was to assess the relative role of isolation-by-distance and postglacial history on patterns of nuclear variation. Because of its small effective population size and maternal transmission, the chloroplast genome is a good marker of population history. Both kinds of nuclear variation (RAPD and allozyme) were therefore compared, first, to the geographical distances among populations and, secondly, to chloroplast DNA restriction polymorphism in the same populations. Multiple Mantel tests were used for this purpose. Although RAPDs revealed less genetic diversity than allozymes, levels of genetic differentiation ( G _ST) were identical. The standard genetic distance calculated at all RAPD loci was correlated with geographical distances but not with the genetic distance calculated from chloroplast DNA data. Conversely, allozyme variation was correlated with chloroplast DNA variation, but not with geography. Possibly, divergent selection at two allozyme loci during the glacial period could explain this pattern. Because of its greater number of loci assayed, RAPDs probably provided a less biased picture of the relative role of geography and history.