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).     

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.     

Natural hybridisation between Quercus petraea (Matt.) Liebl. and Quercus pubescens Willd. within an Italian stand as revealed by microsatellite fingerprinting

Interspecific gene flow is frequently reported in the genus Quercus . However, interfertile oak species often seem to remain distinct, even within areas of sympatry. This study employed molecular markers to verify, at a fine scale, the presence of interspecific gene flow in a natural population of Quercus petraea and Quercus pubescens . Within a delimited area of 6 ha, all adult trees belonging to the studied oak complex and seeds from a subsample of such trees were collected and analysed using molecular microsatellite markers. A low interspecific genetic differentiation and a high level of interspecific genetic admixture suggested past hybridisation. Paternity inference of seeds allowed the estimation of pollination frequencies from the three groups of pollen donors ( Q. petraea, Q. pubescens , intermediate). We also assayed pollen viability and germinability of each species group. We observed natural hybridisation between Q. petraea and Q. pubescens, with a predominant component in the direction Q. petraea → Q. pubescens : Q. pubescens displayed a higher level of heterospecific pollination by Q. petraea (25.8%) and intermediate morphotypes (14.7%), compared to Q. petraea acting as pollen receptor (with less than 5% heterospecific pollinations). Intermediate 'mother trees' were pollinated in similar proportions by Q. petraea (23.1%), Q. pubescens (37.8%) and intermediate morphotypes (39.1%). The asymmetrical introgression observed for the studied generation may be caused, among other factors, by the relative abundance of trees from each species group in the studied area.     

Effects of liming on rhizosphere chemistry and growth of fine roots and of shoots of sessile oak (Quercus petraea)

Soil acidification can be detrimental to root growth and nutrient uptake, and liming may alleviate such acidification. In the following study, seedlings of sessile oak (Quercus petraea Liebl. M.) were grown in rhizotrons and subjected to liming (L) or gypsum (G) treatments and compared with the control (C). In order to study and interpret the impact of these calcium rich treatments on fine root development and tree growth, the following parameters were assessed: fine root biomass, fine root length, seedling development (height, diameter, leaves), seedling biomass, nutrient content of roots and seedlings, bulk soil and soil solution chemistry and rhizosphere soil chemistry. The results show that liming increased bulk soil pH, exchangeable Mg, Ca and the Ca/Al molar ratio, and decreased exchangeable Al, mainly in the A-horizon. Gypsum had a similar but smaller impact on exchangeable Al, Ca, H⁺ and the Ca/Al molar ratio in the A-horizon, but reacted with depth, so that exchangeable Mn, Mg and Ca were increased in the B-horizon. In the rhizosphere, the general pattern was determined by the treatment effects of the bulk soil. Most elements were more concentrated in the rhizosphere than in bulk soil, except for Ca which was less concentrated after liming or gypsum application. In the B-horizon rhizosphere pH was increased by the treatments (L > G,C) close to the root tips. Furthermore, the length of the zone with a positive root-induced pH increase was greater for the limed roots as compared with both the other treatments. Fine root growth was stimulated by liming (L > G,C) both in terms of biomass and length, whereas specific root length was not obviously affected apart from the indication of some stimulation after liming at the beginning. The live:dead ratio of fine roots was significantly higher in the limed rhizotrons as compared to the control (G not assessed), indicating lower mortality (higher longevity). Shoot growth showed greater lime-induced stimulation (L > G,C) as compared to root growth. As a result the shoot:root ratio was higher in the limed rhizotrons than in the control (L > G,C). Liming induced a higher allocation of P, S, Mg, Ca and K to the leaves, stem and twigs. Gypsum showed similar effects, but was only significant for S. Liming increased the foliar Ca/Al ratio by both increasing foliar Ca and decreasing foliar Al, whereas gypsum did not clearly improve foliar nutrition. This study suggests that a moderate application of lime can be successful in stimulating seedling growth, but that gypsum had no effect on seedling growth. It can be concluded that this lime-induced growth stimulation is directly related to the improved soil fertility status, and the alleviation of Al toxicity and acid stress, resulting in better foliar nutrition. The impact of liming on fine roots, as a consequence, was not limited to a stimulation of the total amount of fine roots, but also improved the root uptake performance. This revised version was published online in June 2006 with corrections to the Cover Date.     

Strength, diversity and plasticity of postmating reproductive barriers between two hybridizing oak species (Quercus robur L. and Quercus petraea (Matt) Liebl.)

Very little is known about the nature and strength of reproductive isolation (RI) in Quercus species, despite extensive research on the estimation and evolutionary significance of hybridization rates. We characterized postmating pre- and postzygotic RI between two hybridizing oak species, Quercus robur and Quercus petraea, using a large set of controlled crosses between different genotypes. Various traits potentially associated with reproductive barriers were quantified at several life history stages, from pollen-pistil interactions to seed set and progeny fitness-related traits. Results indicate strong intrinsic postmating prezygotic barriers, with significant barriers also at the postzygotic level, but relatively weaker extrinsic barriers on early hybrid fitness measures assessed in controlled conditions. Using general linear modelling of common garden data with clonal replicates, we showed that most traits exhibited important genotypic differences, as well as different levels of sensitivity to micro-environmental heterogeneity. These new findings suggest a large potential genetic diversity and plasticity of reproductive barriers and are confronted with hybridization evidence in these oak species.     

Tree rooting patterns and soil water relations of healthy and damaged stands of mature oak (Quercus robur L. and Quercus petraea [Matt.] Liebl.)

At three sites in northwestern Germany, which represent the centres of the present oak damage, root distribution and biomass beneath healthy and damaged trees of mature pedunculate oak (Quercus robur L.; Neuenburg site) and sessile oak (Q. petraea [Matt.] Liebl.; Lappwald and Sprakensehl sites) were investigated, and soil texture, bulk density, duration of waterlogging periods and the water available in the mineral soil were determined. For Neuenburg and Sprakensehl, the available soil water was related to leaf water parameters determined in a separate investigation. At the clayey and hydromorphic sites of Neuenburg and Lappwald, the measurements were performed in each one healthy and one damaged part of the site, which differed in the number of oaks with crown damage. In the damaged stand of Neuenburg, the clay content of the subsoil was higher than in the healthy stand, and the soil water availability was reduced especially in dry periods. Compared to healthy oaks of the healthy stand, the density of finest plus fine roots as well as the biomasses of finest roots were lower beneath damaged oaks of the damaged stand. With decreasing relative available soil water (actually available water in relation to water available at the saturation state), the relative leaf water content decreased in damaged, but not in healthy oaks. At Lappwald, similar differences in soil water availability between the healthy and the damaged stand were found, but had no effect on the distribution or biomass of the roots. At the sandy site (Sprakensehl), the available soil water decreased drastically during a dry period, and predawn leaf water potentials of both healthy and damaged oaks declined with decreasing relative available soil water. However, the damaged oaks were not inferior to the healthy ones with respect to root density and biomass. It is concluded that, in the damaged stand of Neuenburg, the high clay content of the subsoil, which results in prolonged periods of waterlogging, in sharp changes from waterlogging to drought and decreased water availability in dry periods, is the reason for the reduced biomass and density of roots of the pedunculate oak. Thus, in northwestern Germany, unfavourable soil water relations are considered as a factor contributing to crown damage of pedunculate oak at hydromorphic sites, but not to damage of sessile oak.     

The Description of Clones of Quercus robur L. and Q. petraea (Matt.) Liebl. with Microsatellites and AFLP in an Ancient Woodland

Abstract: In densely populated areas autochthonous Quercus robur L. (pedunculate oak) and Q. petraea (Matt.) Liebl. (sessile oak) (Fagaceae) populations have been maintained as ancient devastated woodlands. The continuous cutting, grazing and resprouting of such woodlands has enabled the development of clonal structures. For conservation purposes, an analysis of the actual number, size and spatial distribution of clones is necessary, especially when there is an interest in genetic variation of the population. This study describes for the first time - based on microsatellite and AFLP™ analysis - clones in an autochthonous mixed Q. robur and Q. petraea population that has been coppiced and grazed for several centuries. Based on six microsatellite loci and 69 polymorphic AFLP markers, only 14 unique genotypes were detected in a plot that consisted of 80 trees. Clones were observed for both Q. robur and Q. petraea. The largest clone diameters were observed for Q. robur, with distances up to 5.8 m. The observed clone sizes may indicate the age of the trees.     

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.     

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     

Photosynthesis of oak trees [Quercus petraea (Matt.) Liebl.] during drought under field conditions: diurnal course of net CO2 assimilation and photochemical efficiency of photosystem II

Adult trees of Quercus petraea were submitted to controlled water shortage in a natural stand near Nancy, France. Diurnal course of net CO₂ assimilation rate (A) was measured in situ together with chlorophyll a fluorescence determined on dark adapted leaves. In 1990, trees experienced a strong water stress, with predawn and midday leaf water potentials below –2·0 and –3·0 MPa, respectively. Diurnal course of A of well-watered trees exhibited sometimes important midday decreases in A related to high temperature and vapour pressure deficit. Decreases in initial (F_o) and maximal (F_m) fluorescence and sometimes in photochemical efficiency of photosystem II (F_v/F_m) were observed and probably revealed the onset of mechanisms for thermal de-excitation. These mechanisms were shown to be sensitive to dithiothreitol. All these effects were reversible and vanished almost completely overnight. Therefore, they may be considered as protective mechanisms adjusting activity of photosystem II to the electron requirement for photosynthesis. Water stress amplified these reactions: A was strongly decreased, showing important midday depression; diurnal reductions in F_m and F_v/F_m were enhanced. The same trends were observed during summer 1991, despite a less marked drought. These protective mechanisms seemed very effective, as no photoinhibitory damage to PS II could be detected in either water stressed or control trees.     

Fine root biomass in a beech (Fagus sylvatica L.) stand on Paiko Mountain,NW Greece

Abstract

Fine roots represent a small proportion of total plant biomass however they represent the most dynamic component of the root systems of woody plants. There is limited information on the beech fine root production in Mediterranean ecosystems and especially in Greece. We measured live, dead and total fine root biomass (d<2 mm) (LFRB, DFRB and TFRB, respectively) over a growing season in a beech (Fagus sylvatica L.) stand on Paiko mountain, NW Greece, in order to contribute to the generally scarce knowledge of the fine root biomass of beech stands. It was found that TFRB and LFRB increased from May to July and then decreased. LFRB decreased with soil depth while there was no pattern at the change of DFRB with soil depth.