Presence of triploids among oak species

BACKGROUND AND AIMS: Polyploids, organisms that have multiple sets of chromosomes, are common in certain plant and animal taxa. However, there are only a few reports of intraspecific ploidy variation within the genus Quercus. The aim of the study was to investigate the suspected ploidy level of two oaks that have unusual microsatellite banding patterns. METHODS: Polyploidy was investigated by using microsatellite analysis, stomata length measurements and nuclear DNA content estimation by flow cytometry. KEY RESULTS: Each putative triploid tree has patterns of microsatellite variation unexpected for diploid genomes, with up to three alleles at some loci, significantly longer stomata and 1.5 times more DNA per nucleus compared with diploids. CONCLUSIONS: To our knowledge, this report contains the first evidence for triploidy in Q. petraea and confirmation of this phenomenon in Q. robur. Regardless of the positive or negative aspects of the presence of triploid oaks in forest stands, it is of value to be able to screen for them. This study demonstrates that nuclear microsatellites and estimation of DNA content by flow cytometry can readily be used for this purpose.     

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.     

An evaluation of standard oak tree growth in Ruislip woods, West London

Tree-ring studies on dominant and co-dominant oak standards from four woodlands on the western fringes of London were studied in relation to known historical and meteorological conditions. Whilst previous studies suggested a decline in oak growth in recent decades, this was not apparent from this study. Major recruitment periods for the large oaks were determined. The sites had differing responses to known drought years. Coppicing of the undergrowth was found not to greatly influence the growth of the remaining standard trees on all occasions, although one woodland showed some growth release following recent coppicing. Records of lichens for each woodland were used to assess changing pollution levels at the sites.     

Climate change impacts on tree ranges: model intercomparison facilitates understanding and quantification of uncertainty

Model-based projections of shifts in tree species range due to climate change are becoming an important decision support tool for forest management. However, poorly evaluated sources of uncertainty require more scrutiny before relying heavily on models for decision-making. We evaluated uncertainty arising from differences in model formulations of tree response to climate change based on a rigorous intercomparison of projections of tree distributions in France. We compared eight models ranging from niche-based to process-based models. On average, models project large range contractions of temperate tree species in lowlands due to climate change. There was substantial disagreement between models for temperate broadleaf deciduous tree species, but differences in the capacity of models to account for rising CO(2) impacts explained much of the disagreement. There was good quantitative agreement among models concerning the range contractions for Scots pine. For the dominant Mediterranean tree species, Holm oak, all models foresee substantial range expansion.     

Transmission properties of an oak canopy in relation to photoperception

Abstract The spectra of incoming daylight and shadelight in a mature oak woodland were measured at intervals during the canopy cycle, and mean transmittance spectra were derived. Transmittance was spectrally neutral at ca. 0.55 during the light phase but, following leaf emergence, transmittance of PAR (400–700 nm) fell to ca. 0.1. Simultaneously, the red : far-red transmittance ratio fell to circa 0.6. Both showed little change during the summer and autumn until senescence, indicating that the optical properties of the canopy were surprisingly stable. There was no evidence that cloud cover influenced mean canopy transmittance, although transient sunflecks introduced great variability which, in combination with sampling bias, might explain previous contradictory reports. The red : far-red fluence rate ratio in the woodland showed a temporary increase in late summer, a result of a small increase in the red : far-red ratio of incoming daylight during this period. Reflectance and transmittance spectra and pigment content of sun and shade leaves were measured. Leaf transmittance spectra showed changes correlated with those of the canopy, and were related to changes in pigment content.     

Opposing effects of spring defoliation on late season oak caterpillars

ABSTRACT. 1. The pedunculate oak, Quercus rohur L., suffers high annual levels of spring defoliation in Wytham Woods. near Oxford.
2. This spring defoliation affects late season caterpillars through a variety of damage-induced changes in the leaves.
3. Diurnea fagella (D. & S.), one of the commonest late season caterpillars, shows reduced larval survival and pupal weight on regrowth foliage when compared to undamaged primary foliage.
4. D. fagella also suffer higher larval mortality on naturally damaged primary foliage than they do on undamaged foliage.
5. Despite this, the three commonest late season caterpillar species are more abundant on damaged trees than undamaged ones. and their distributions are biased towards damaged leaves within the canopy.
6. Other factors that may be more important than leaf damage in determining the distribution and abundance of late season caterpillars are discussed. D.fagella larvae spin their larval refuges more rapidly on damaged than undamaged foliage, and this may reduce mortality by natural enemies, or ameliorate adverse effects of weather.     

Differential activity of catalase and superoxide dismutase in seedlings and in vitro micropropagated oak (Quercus robur L.)

 Catalase (CAT) and superoxide dismutase (SOD) activity profiles were examined by native polyacrylamide gel electrophoresis in different tissues of seedlings and microcuttings of oak (Quercus robur L.) initiated from crown material (NL100A) and from basal epicormic shoots (NL100R), which differ in rooting ability. Two CAT isoforms were differentially active in seedlings and microcuttings; in particular, CAT-2 was activated in the basal callus of rooted microshoots. SOD isoenzymes, Mn-SOD and at least four Cu/Zn-SODs were found to be present, with Mn-SODs particularly active in microcuttings. No differences were found between the electrophoretic profiles of the two lines despite their different ontogenetic origin. The strong activity of CAT-2 in rooted microshoots indicates that this isoform is a protein specifically related to rooting. Received: 21 February 2000 / Revision received: 18 October 2000 / Accepted: 18 October 2000     

Factors underlying the late seasonal appearance of the lepidopterous leaf-mining guild on oak

Abstract. 1. Most lepidopterous leaf mining species found on the oak Quercus robur in Britain develop in summer. At this time of year, externally feeding caterpillars remove little leaf area since most of these free living folivores are predominantly spring feeders.
2. I forced Phyllonorycter harrisella (L.) miners to oviposit in spring, then exposed developing larvae to a wide range of leaf damage levels.
3. Leaf miner survivorship and mean female pupal weight were significantly greater in the experimental spring generation on undamaged oak leaves, this being when oak foliage is of the highest nutritional quality.
4. Leaf miner survivorship in all generations is correlated with natural leaf damage levels. Experimental leaf damage also reduces miner survivorship.
5. Leaf damage reduces miner survivorship by increasing the probability of larval death due to wound induced responses.
6. The role of asymmetrical competitive interactions between caterpillars and leaf-miners in determining the late seasonal appearance of miners is discussed. It is suggested that wound induced responses may play an important part in the structuring of phytophagous insect communities.     

The effects of increasing CO2 and temperature on oak leaf palatability and the implications for herbivorous insects

Rising levels of atmospheric CO₂ are expected to perturb forest ecosystems, although the extent to which specific ecological interactions will be modified is unclear. This research evaluates the effects of elevated CO₂ and temperature, alone and in combination, on the leaf nutritional quality of Pendunculate oak (Quercus robur L.), and the implications for herbiverous insect defoliators are discussed. A 3 °C temperature rise reduced leaf nutritional quality, by reducing foliar nitrogen concentration and increasing condensed tannin content. Doubling atmospheric CO₂ temporarily increased total phenolics, but also reduced leaf toughness. The nutritional quality of the second leaf flush (lammas growth) was considerably reduced at elevated CO₂. It is concluded that larval development of spring-feeding defoliators and hence adult fecundity may be adversely affected by increased temperatures.     

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.     

The potential for interspecific competition between two abundant defoliators on oak: leaf damage and habitat quality

Abstract. 1. The pedunculate oak, Quercus robur L., suffers high annual levels of spring defoliation in Wytham Woods, Oxon. The two major defoliators, Tortrix viridana L. and Operophtera brumata L., sometimes reach high enough densities to completely defoliate trees.
2. Experiments with larvae of both species enclosed on oak trees in Wytham Woods suggest considerable potential for intra- and interspecific competition between T. viridana and O. brumata.
3. T. viridana and O. brumata consume equivalent amounts of leaf tissue per unit caterpillar density. Despite this, O. brumata is the superior competitor, measured by mortality during the larval feeding period.
4. Evidence is presented that T. viridana larvae depend upon the protected microclimate of leaf rolls to maintain water balance, and that leaf damage may disrupt the construction of, and microclimate within, leaf rolls. O. brumata may therefore be competitively superior because it indirectly interferes with the water balance of T. viridana.
5. The importance of interspecific competition in the dynamics of T. viridana and O. brumata populations is discussed. Stochastic mortality factors acting on first instar larvae of both species probably make intense competition a rare event.     

Adaptive responses for seed and leaf phenology in natural populations of sessile oak along an altitudinal gradient

We assessed the adaptive potential of seed and leaf phenology in 10 natural populations of sessile oak (Quercus petraea) sampled along two altitudinal transects using common garden experiments. Population differentiation for both phenological traits was observed with high-altitude populations germinating and flushing later than low altitude ones. However, high genetic variation and heritability values were also maintained within populations, despite slightly decreasing for dates of leaf unfolding with increasing altitude. We suggest that biotic and abiotic fluctuating selection pressures within populations and high gene flow are the main mechanisms maintaining high genetic variation for these fitness related traits. Moreover, changes in selection intensity and/or selection pressures along the altitudinal gradient can explain the reduction in genetic variation observed for leaf phenology. We anticipate that the maintenance of high genetic variation will be a valuable resource for future adaptation of sessile oak populations undergoing an upslope shift caused by climate change.     

Morphological and molecular diversity among Italian populations of Quercus petraea (Fagaceae)

Quercus petraea (sessile oak) has a scattered distribution in southern and central Italy. The objective of this work was to evaluate the level and distribution of diversity in five Italian populations of Q. petraea by using morphological markers and hypervariable molecular markers such as microsatellites. Forty-eight morphological traits and six nuclear and three plastid loci were scored for each population. Evidence for differentiation in both sets of traits was found, but patterns of differentiation of morphological traits did not coincide with microsatellite differentiation. Morphological variation was correlated with ecological conditions at the site of origin. Analysis of molecular variance revealed significant genetic variation among populations (P < 0.001), both at the nuclear and plastid levels. There was a slight, but significant, correlation between nuclear genetic distance and geographic distance. The relatively high genetic diversity in the populations analysed indicates that the maintenance of their evolutionary potential is possible if population sizes are maintained or increased. Low levels of haplotype diversity found within the small southernmost population (Piano Costantino) indicates that genetic erosion may increase the extinction risk for this population.     

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).     

Water in the stems of sessile oak (Quercus petraea) assessed by computer tomography with concurrent measurements of sap velocity and ultrasound emission

Concurrent measurements of heat pulse velocity and ultrasound acoustic emission were performed on the trunks of adult Quercus petraea plants under different water stress conditions. On the trunk section of the plants the wood density was measured non-destructively using a mobile computer tomograph which measures the attenuation of a collimated beam of radiation which traverses the trunk. By scanning the trunk in different directions, it is possible to map wood density in the section. As wood density is strictly correlated with water content, this method allows evaluation of the water content in the trunk section and the water conditions in the different parts of the section. The computer-tomograph technique is non-invasive and is not influenced by climatic fluctuations. A close agreement was found between wood density and heat pulse velocity; the relationship between these two parameters and ultrasound acoustic emission is discussed. Trunks of sessile oak appear well suited as water storage reservoirs playing an important role in tree survival during extended periods of low soil water availability, especially in the context of global climatic change. Here the computer-tomograph methodology is described and suggestions arc made for further research development.     

Woodland area,species turnover and the conservation of bird assemblages in lowland England

A study over 4 years into the number of breeding bird species and species turnover (extinctions and colonisations) in relation to area was conducted in 35 woodlands, set in an intensively farmed landscape, in north-east Essex, UK. A total of 46 species was recorded. The number of species breeding increased with woodland area; the slope of the species–area relationship did not differ between years. Habitat diversity was the only other measured variable to influence species richness. Absolute species turnover was independent of woodland area but relative turnover declined with increase in woodland area. The numbers of territories of nine species were determined. For four summer visitors the number of woods occupied increased as the overall populations increased but, for the other species, changes in overall population size led to changes in numbers in occupied woods. Chaffinch Fringilla coelebs and Song Thrush Turdus philomelos were more associated with woodland edges, Nightingale Luscinia megarhynchos, Garden Warbler Sylvia borin, Chiffchaff Phylloscopus collybita and Willow Warbler P. trochilus with interiors. Several species showed an inverse relationship between population density and woodland area. Collections of small woods hold similar species richness to single large woods. While the acquisition of large woods for conservation purposes should be a priority, the extension of smaller woods to a size of about 10 ha would be highly beneficial to both the species richness and population stability of regional woodland bird assemblages.     

High nuclear genetic diversity, high levels of outcrossing and low differentiation among remnant populations of Quercus petraea at the margin of its range in Ireland

• Background and Aims Quercus petraea colonized Irelandafter the last glaciation from refugia on mainland Europe. Deforestation,however, beginning in Neolithic times, has resulted in small,scattered forest fragments, now covering less than 12 000 ha. • Methods Plastid (three fragments) and microsatellitevariation (13 loci) were characterized in seven Irish populationssampled along a north–south gradient. Using Bayesian approachesand Wright’s F-statistics, the effects of colonizationand fragmentation on the genetic structure and mating patternsof extant oak populations were investigated. • Key Results All populations possessed cytotypes commonto the Iberian Peninsula. Despite the distance from the refugialcore and the extensive deforestation in Ireland, nuclear geneticvariation was high and comparable to mainland Europe. Low populationdifferentiation was observed within Ireland and populationsshowed no evidence for isolation by distance. As expected ofa marker with an effective population size of one-quarter relativeto the nuclear genome, plastid variation indicated higher differentiation.Individual inbreeding coefficients indicated high levels ofoutcrossing. • Conclusions Consistent with a large effective populationsize in the historical migrant gene pool and/or with high geneflow among populations, high within-population diversity andlow population differentiation was observed within Ireland.It is proposed that native Q. petraea populations in Irelandshare a common phylogeographic history and that the presentgenetic structure does not reflect founder effects.