Analysing branching pattern in plantations of young red oak trees (Quercus rubra L., Fagaceae)

Branching patterns of the growth units of monocyclic or bicyclic annual shoots on the main axis of 5-year-old red oaks were studied in a plantation in south-western France. For each growth unit, the production of axillary structures associated with each node was described in the form of a sequence. For a given category of growth units, homogeneous zones (i.e. zones in which composition in terms of type of axillary production does not change substantially) were identified on such sequences using a dedicated statistical model called a hidden semi-Markov chain. For instance, on the first growth unit of bicyclic annual shoots, a zone with 1-year-delayed branches was found systematically below a zone with buds and one-cycle-delayed branches. Branching patterns shown by the growth unit of monocyclic annual shoots and on the second growth unit of bicyclic annual shoots were very similar. Branches with a 1-year delay in development tended to be polycyclic at the top of the growth unit and monocyclic lower down. The number of nodes shown by the branched zone of the growth unit of monocyclic annual shoots was stable, irrespective of the total number of nodes of the growth unit. In contrast, the second growth unit of bicyclic annual shoots exhibited a correlation between the number of nodes in the branching zone and the total number of nodes. The contribution made by this method to understanding plant functioning is discussed.     

Microsatellite markers for the Indian golden silkmoth, Antheraea assama (Saturniidae: Lepidoptera)

Antheraea assama, an economically important and scientifically unexplored Indian wild silkmoth, is unique among saturniid moths. For this species, a total of 87 microsatellite markers was derived from 35 000 expressed sequence tags and a microsatellite‐enriched sub‐genomic library. Forty individuals collected from Tura and West Garo Hills region of Northeast India were screened for each of these loci. Ten loci from expressed sequence tags and one from genomic library were found to be polymorphic. These microsatellite markers will be useful resources for population genetic studies of A. assama and other closely related species of saturniids. This is the first report on development of microsatellite markers for any saturniid species.     

Population dynamics and growth patterns for a cohort of northern red oak (Quercus rubra) seedlings

Summary I studied the survival and development of a 1986 cohort of northern red oak (Quercus rubra L.) seedlings growing under a variety of overstory and microsite conditions in a northern hardwood forest dominated by northern red oak, red maple (Acer rubrum L.) paper birch (Betula papyrifera Marsh.), and scattered white pine (Pinus strobus L.). Fifty naturally regenerating seedlings of oak were randomly selected in each of three canopy classes: no overstory, partial overstory, and complete overstory. Growth and mortality were measured for six years. Seedling height growth decreased with overstory density, with less growth evident with even a partial overstory. Seedling survival also declined with overstory density and depended on microtopography to a lesser extent. After six years, 92% of the seedlings survived in the open, compared to 54% under the partial overstory, and 36% under the complete overstory. The open environment, in which woody and herbaceous regrowth formed a low canopy reducing light intensities to about 50% of full sunlight, provided a favorable site for the growth and survival of northern red oak.     

Radial growth and climate responses of white oak (Quercus alba) and northern red oak (Quercus rubra) at the northern distribution limit of white oak in Quebec, Canada

Aim The objectives of this study were: (1) to compare radial growth patterns between white oak (Quercus alba L.) and northern red oak (Quercus rubra L.) growing at the northern distribution limit of white oak; and (2) to assess if the radial growth of white oak at its northern distribution limit is controlled by cold temperature. Location The study was conducted in three regions of the Ottawa valley in southern Québec. All stands selected were located at the northern limit of distribution of Q. alba. Methods Twelve mixed red and white oak stands were sampled and increment cores were extracted for radial growth analyses. For each oak species, 12 chronologies were derived from tree‐ring measurement (residual chronologies). Principal components analysis and redundancy analysis were used to highlight the difference between radial growth in both species and to determine their radial growth–climate association. Results There was little difference between the radial growth of each species; Q. alba, however, exhibits more year‐to‐year variation in growth than Q. rubra. More than 65% of the variance in radial growth was shared among sites and species. Both species showed a similar response to climate, which suggested that the limit of distribution of Q. alba might not be determined by effects on growth. Both species had a classic response to climate and drought in the early growing season. Main conclusions The northern distribution limit of Q. alba does not appear to be directly controlled by effects on growth processes as indicated by the similarities in radial growth and response to climate between the two species. The location of the stands on southern aspects suggested that cold temperature could have been a major factor controlling the distribution limit of Q. alba. However, it is speculated that stands growing on southern aspects may be more prone to forest fires or to drought, which would favour the maintenance and establishment of oaks, and of Q. alba in particular. Models relating the northern distribution limits of species to broad climate parameters like annual mean temperature will need to be reviewed to incorporate more biologically relevant information. Such assessments will in turn provide better estimates of the effect of climate changes on species distribution.     

Callus formation from root protoplasts of Quercus rubra L. (red oak)

Root protoplasts of Quercus rubra L. were isolated from 12 day old seedlings with an enzyme mixture containing Cellulase R1O + Rhozyme HP150 + Macerozyme R1O, supplemented with cysteine and bovine serum albumin.Protoplasts were purified by a Ficoll density gradient centrifugation and cultured at low density in a liquid medium. The modified woody plant medium, containing 2.2 M benzyladenine + 1.8; M zeatin + 5.3 M -naphthaleneacetic acid + 2.2 M dichlorophenoxyacetic acid, allowed sustained divisions and formation of microcalluses.Protoplast — derived microcallus developed into green and compact callus when transferred to an agarose solidified medium, supplemented with casein hydrolysate and indole 3-acetic acid (devoid of 2,4 dichlorophenoxyacetic acid) and placed under low illumination.Abbreviations IAA indole-3-acetic acid - IBA indole-3-butyric acid - BA benzyladenine - BSA bovine serum albumin - 2,4-D 2,4-dichlorophenoxyacetic acid - FW fresh weight - GA₃ gibberellic acid - MES 2-(N-morpholino) ethanesulfonic acid - MS medium Murashige and Skoog medium (1962) - NAA -naphthaleneacetic acid - WPM woody plant medium (Lloyd and Mc Cown, 1981)     

Pyrosequencing of the northern red oak (Quercus rubra L.) chloroplast genome reveals high quality polymorphisms for population management

Given the low intraspecific chloroplast diversity detected in northern red oak (Quercus rubra L.), more powerful genetic tools are necessary to accurately characterize Q. rubra chloroplast diversity and structure. We report the sequencing, assembly, and annotation of the chloroplast genome of northern red oak via pyrosequencing and a combination of de novo and reference-guided assembly (RGA). Chloroplast DNA from 16 individuals was separated into four MID-tagged pools for a Genome Sequencer 20 quarter-run (Roche Life Sciences, Indianapolis, IN, USA). A four-step assembly method was used to generate the Q. rubra chloroplast consensus sequence: (1) reads were assembled de novo into contigs, (2) de novo contigs were aligned to a reference genome and merged to produce a consensus sequence, (3) the consensus sequence was aligned to the reference sequence and gaps between contigs were filled with reference sequence to generate a "pseudoreference", and (4) reads were mapped to the pseudoreference using RGA to generate the draft chloroplast genome. One hundred percent of the pseudoreference sequence was covered with a minimum coverage of 2× and an average coverage of 43.75×. The 161,304-bp Q. rubra chloroplast genome draft sequence contained 137 genes and one rps19 pseudogene. The sequence was compared to that of Quercus robur and Q. nigra with 951 and 186 insertion/deletion or SNP polymorphisms detected, respectively. A total of 51 intraspecific polymorphisms were detected among four northern red oak individuals. The fully sequenced and annotated Q. rubra chloroplast genome containing locations of interspecific and intraspecific polymorphisms will be essential for studying population differentiation, phylogeography, and evolutionary history of this species as well as meeting management goals such as monitoring reintroduced populations, tracking wood products, and certifying seed lots and forests.     

Hydraulic conductivity of red oak (Quercus rubra L.) leaf tissue does not respond to light

The permeability of leaf tissue to water has been reported to increase under illumination, a response reputed to involve aquaporins. We studied this ‘light response’ in red oak (Quercus rubra L.), the species in which the phenomenon was first detected during measurements of leaf hydraulic conductance with the high‐pressure flow meter (HPFM). In our HPFM measurements, we found that pre‐conditioning leaves in darkness was not sufficient to bring them to their minimum conductance, which was attained only after an hour of submersion and pressurization. However, pre‐conditioning leaves under anoxic conditions resulted in an immediate reduction in conductance. Leaves light‐ and dark‐acclimated while on the tree showed no differences in the time course of HPFM measurement under illumination. We also studied the effect of light level and anoxia on rehydration kinetics, finding that anoxia slowed rehydration, but light had no effect either in the lab (rehydration under low light, high humidity) or on the tree (acclimation under high light, 10 min of dark prior to rehydration). We conclude that the declines in conductance observed in the HPFM must involve a resistance downstream of the extracellular air space, and that in red oak the hydraulic conductivity of leaf tissue is insensitive to light.     

Microsatellite markers derived from Quercus mongolica var. crispula (Fagaceae) inner bark expressed sequence tags

In reforestation programs the genetic composition and diversity of populations that could be used as sources of planting material needs to be carefully considered to maximize the chances of successful establishment. For such purposes genetic analyses that include the identification of functional genes are required. In this study, we constructed a cDNA library from inner bark of Quercus mongolica (which is widely distributed in Japan) and collected 3385 ESTs. After constructing 2140 unigenes, 274 microsatellites were found within them. The most frequent microsatellite had AG motif (48%) and the next most common was AAG motif (12%). There were no CG repeats in the unigenes. In total, 20 EST-SSR markers were developed, polymorphisms of which were described by using eight individuals from eight populations over the species' distributional range. The number of alleles per locus (Na) and observed heterozygosity (H(o)) ranged from 2 to 12, and from 0.25 to 1.00, respectively. Cross-species amplification was successful for 19 loci in eight individuals of Q. serrata and for 20 loci in eight individuals of Q. dentata, with values of Na and H(o) comparable to those of Q. mongolica. The EST-SSR markers characterized in this study should facilitate the analysis of genetic diversity in future studies.     

Geometric morphometric analyses of leaf variation in four oak species of the subgenus Quercus (Fagaceae)

Abstract

This paper defines a protocol for the study of leaf morphology and morphometric relationships among Quercus frainetto, Quercus robur, Quercus petraea and Quercus pubescens. Leaf shapes were investigated in an attempt to characterize and discriminate among these four species. Landmark and outline data were compared in order to ascertain their relative efficacy for describing variation in leaf morphology. MANOVA was performed to assess intra‐specific differentiation and to compare the distribution of variance in individuals, populations and species. A strong correlation was found between landmark and outline datasets of the four species: both methods proved capable of discriminating among the four species. Finally, a Two‐Block Partial Least‐Squares analysis was performed for each species in order to assess the correlation between intra‐specific shape variation and environmental factors.     

Effect of water availability on the phenotypic expression of herbivore resistance in northern red oak seedlings (Quercus rubra L.)

The phenotypic expression of quantitative characters is a function of the individual's genotype and the environment in which it is measured. In a previous reciprocal transplant study, we found that patterns of genetic differences in resistance to herbivores among adjacent subpopulations of northern red oak (Quercus rubra L.), were consistent with a local adaptation hypothesis. The goal of this study was to determine if variation in water availability may have been a mechanism responsible for these previously observed patterns. In 1989 a common garden study was initiated using acorns from maternal trees occupying either a north- or south-facing slope microhabitat in an oak-hickory forest in east central Missouri, USA. The seedlings were grown under one of two water treatments, irrigated or natural. In 1992, we utilized this experiment to examine the quantitative character of the percentage of leaf area damaged by herbivores, which is a measure of the phenotypically expressed level of resistance. Specifically, we made three predictions: (1) because northern red oak seem to grow best in mesic environments, seedlings receiving more water should show greater resistance to herbivores; (2) if the subpopulations from north- and south-facing slope microhabitats are genetically differentiated with respect to the quantitative character of resistance to herbivores, then there will be a significant effect of maternal slope microhabitat on the percentage of leaf area damaged; and (3) if the pattern of resistance to herbivores found among subpopulations reflects local adaptation to moisture levels in their own microhabitat, then we would expect to find a significant maternal slope microhabitat by water treatment interaction, with north-facing slope seedlings incurring less damage in the wetter (irrigated) treatment and south-facing slope seedlings incurring less damage in the drier (natural) treatment. Our data supported the first two predictions: seedlings in the irrigated treatment showed a significantly lower percentage of leaf area damage than those in the natural treatment, and the percentage of leaf area damaged was significantly lower on seedlings from maternal plants occupying the north-facing slope microhabitat. However, we found no significant interaction between maternal slope microhabitat and water treatment. These findings demonstrate that northern red oak supbopulations respond phenotypically to water availability, but this factor does not appear to be the underlying mechanism behind the previously observed local adaptation expressed as resistance to herbivores.     

Microsatellite DNA markers for a grasshopper: Prionotropis hystrix rhodanica (Orthoptera,Pamphagidae)

Microsatellite loci were isolated from Prionotropis hystrix rhodanica, an endangered grasshopper species, endemic to a steppic area southeast of France. Six polymorphic loci were obtained from an enriched partial genomic libray. A high genetic diversity was observed at all loci, with an observed number of alleles ranging from seven to 29, and an observed heterozygosity between 0.71 and 0.90. Test of cross‐amplifications on two closely related taxa are presented.     

The effect of heat waves,elevated [CO2] and low soil water availability on northern red oak (Quercus rubra L.) seedlings

The frequency and intensity of heat waves are predicted to increase. This study investigates whether heat waves would have the same impact as a constant increase in temperature with the same heat sum, and whether there would be any interactive effects of elevated [CO₂] and soil moisture content. We grew Quercus rubra seedlings in treatment chambers maintained at either ambient or elevated [CO₂] (380 or 700 μmol CO₂ mol^?1) with temperature treatments of ambient, ambient +3 °C, moderate heat wave (+6 °C every other week) or severe heat wave (+12 °C every fourth week) temperatures. Averaged over a 4‐week period, and the entire growing season, the three elevated temperature treatments had the same average temperature and heat sum. Half the seedlings were watered to a soil water content near field capacity, half to about 50% of this value. Foliar gas exchange measurements were performed morning and afternoon (9:00 and 15:00 hours) before, during and after an applied heat wave in August 2010. Biomass accumulation was measured after five heat wave cycles. Under ambient [CO₂] and well‐watered conditions, biomass accumulation was highest in the +3 °C treatment, intermediate in the +6 °C heat wave and lowest in the +12 °C heat wave treatment. This response was mitigated by elevated [CO₂]. Low soil moisture significantly decreased net photosynthesis (A_net) and biomass in all [CO₂] and temperature treatments. The +12 °C heat wave reduced afternoon A_net by 23% in ambient [CO₂]. Although this reduction was relatively greater under elevated [CO₂], A_net values during this heat wave were still 34% higher than under ambient [CO₂]. We concluded that heat waves affected biomass growth differently than the same amount of heat applied uniformly over the growing season, and that the plant response to heat waves also depends on [CO₂] and soil moisture conditions.     

Preliminary investigations on somatic embryogenesis from leaf discs of red oak (Quercus rubra L.)

Somatic embryos were obtained from leaf discs of juvenile red oak plants. Basal inductive nutrient medium was a modified Murashige and Skoog solution enriched with 500 mg L^–1 casein hydrolysate, 100 mg L^–1 polyvinylpyrrolidone, 5.4 M naphthaleneacetic acid and 0.09 M benzyladenine. Embryogenesis was obtained only from leaf discs in the presence of light and increased when the adaxial surface of the explants (with midrib or main veins present) was in contact with the medium. Large variation was observed in all experiments. Recurrent embryogenesis was observed at the base of embryo clusters with callus present; conversely, embryogenic potential was rapidly lost by subculturing full calli. Maturation, germination and development of isolated somatic embryos were obtained. However, the vast majority of embryos did not have viable apical bud meristems and on only a few occasions were shoots produced.Abbreviations BA N⁶-benzyladenine - CH casein hydrolysate - 2iP isopentenyladenine - NAA naphthaleneacetic acid - 2.4-D 2.4-dichlorophenoxyacetic acid - GA gibberellic acid - PVP polyvinylpyrrolidone     

Effect of a red oak species gradient on genetic structure and diversity of Quercus castanea (Fagaceae) in Mexico

Incipient reproductive barriers are a common characteristic of oak species. Disruption of these barriers promotes changes in diversity and genetic structure of the species involved. Quercus castanea is a red oak with wide geographic distribution in Mexico, which presents atypically high morphological variability when it occurs in sympatry with other red oak species, suggesting that hybridization may explain the observed variation. We tested if the genetic structure and diversity levels of Q. castanea are related to the number of red oak species growing in sympatry. In total, 14 microsatellite (SSRs) primers (six nSSRs and eight cpSSRs) were used in 120 Q. castanea individuals (20/site) belonging to six populations, where the number of red oak species associated varied from zero to five. Results showed a positive and significant relationship between the genetic diversity of Q. castanea and the number of red oak species growing in sympatry, regardless of the marker type or the parameter of genetic diversity analyzed. Also, we found a higher genetic differentiation of Q. castanea populations using cpSSRs in comparison with nSSRs. Our results suggest that temperate forests with high red oaks species richness co-dominated by Q. castanea promote the increase in this species genetic diversity. From a conservation perspective, high genetic diversity levels of foundation species such as Q. castanea may have positive cascade effects extending to other species in the community.     

Microsatellite markers for Spergularia media (L.) C. Presl. (Caryophyllaceae) and their cross-species transferability

We developed 11 polymorphic microsatellite markers for Spergularia media (Caryophyllaceae), a perennial halophyte of coastal salt meadows and continental areas of western Eurasia. The number of alleles per locus observed in a single population of 20 individuals from the German North Sea coast ranged from 3 to 20. Observed and expected heterozygosities ranged from 0.200 to 0.850 and from 0.278 to 0.936 respectively. Observed heterozygosities were lower than expected heterozygosities at all loci, presumably as a consequence of inbreeding. All markers cross-amplified in the closely related S. salina. Of these, nine were polymorphic in the heterologous species. Only two primer pairs generated PCR products in S. diandra.     

Clonal population structure and genetic variation in sand-shinnery oak,Quercus havardii (Fagaceae)

We investigated clonal population structure and genetic variation in Quercus havardii (sand-shinnery oak), a deciduous rhizomatous shrub that dominates vegetation by forming uninterrupted expanses of ground cover over sandy deposits on the plains of western Texas, western Oklahoma, and eastern New Mexico. Isozyme electrophoresis (15 loci coding 11 enzymes) was used to recognize and map clones arrayed in a 2000-m transect (50-m sample intervals) and a 200 × 190 m grid (10-m sample intervals). Ninety-four clones were discovered, 38 in the transect and 56 in the grid, resulting in an estimated density of ～15 clones per hectare. Clones varied greatly in size (～100-7000 m), shape, and degree of fragmentation. The larger clones possessed massive interiors free of intergrowth by other clones, while the smaller clones varied in degree of intergrowth. The population maintained substantial levels of genetic variation (P = 60%, A = 2.5, H(exp) = 0.289) comparable to values obtained for other Quercus spp. and for other long-lived perennials. The population was outcrossing as evidenced by conformance of most loci to Hardy-Weinberg expected genotype proportions, although exceptions indicated a limited degree of population substructuring. These data indicate that despite apparent reproduction primarily through vegetative means, Q. havardii possesses conventional attributes of a sexual population.     

Genetic evidence for hybridization in red oaks (Quercus sect. Lobatae, Fagaceae)

? Premise of the study: Hybridization is pervasive in many plant taxa, with consequences for species taxonomy, local adaptation, and management. Oaks (Quercus spp.) are thought to hybridize readily yet retain distinct traits, drawing into question the biological species concept for such taxa, but the true extent of gene flow is controversial. Genetic data are beginning to shed new light on this issue, but red oaks (section Lobatae), an important component of North American forests, have largely been neglected. Moreover, gene flow estimates may be sensitive to the choice of life stage, marker type, or genetic structure statistic. ? Methods: We coupled genetic structure data with parentage analyses for two mixed-species stands in North Carolina. Genetic structure analyses of adults (including F(ST), R(ST), G'(ST), and structure) reflect long-term patterns of gene flow, while the percentage of seedlings with parents of two different species reflect current levels of gene flow. ? Key results: Genetic structure analyses revealed low differentiation in microsatellite allele frequencies between co-occurring species, suggesting past gene flow. However, methods differed in their sensitivity to differentiation, indicating a need for caution when drawing conclusions from a single method. Parentage analyses identifed >20% of seedlings as potential hybrids. The species examined exhibit distinct morphologies, suggesting selection against intermediate phenotypes. ? Conclusions: Our results suggest that hybridization between co-occurring red oaks occurs, but that selection may limit introgression, especially at functional loci. However, by providing a source of genetic variation, hybridization could influence the response of oaks and other hybridizing taxa to environmental change.     

Branch growth and leaf numbers of red maple (Acer rubrum L.) and red oak (Quercus rubra L.): response to defoliation

Summary Branch growth and leaf formation from terminal and from lateral buds of red maple (Acer rubrum L.) and red oak (Quercus rubra L.) were measured in response to simulated insect defoliation. A single large branch representative of the crown of each tree was used for enumeration of growth and of bud numbers throughout three successive years of 0, 50, 75, and 100% leaf removal for the entire tree. Leaf number per tree for both species after the last year of defoliation was reduced in direct proportion to the severity of defoliation, in comparison to the predefoliation status of the trees. Bud number per tree for red maple, but not for red oak, was also reduced in proportion to severity of defoliation.Averaged over all defoliation treatments, defoliation reduced branch growth more than leaf production. Furthermore, the reduction in branch growth and leaf production was greater in red oak than in red maple. Three years of successive defoliation reduced the mean lateral plus terminal branch growth by 40% in red oak and by 23% in red maple, while leaf number was reduced 22% in red oak and remained unchanged in red maple. In red maple, 100% defoliation caused greater branch death than the 50 or 75% defoliation treatments, and the amount of death was greater after each successive year of defoliation. In contrast to red maple, undefoliated red oak incurred a substantial amount of branch death throughout the study which was little affected by defoliation treatment.     

High genetic diversity and stable Pleistocene distributional ranges in the widespread Mexican red oak Quercus castanea Née (1801) (Fagaceae)

The Mexican highlands are areas of high biological complexity where taxa of Nearctic and Neotropical origin and different population histories are found. To gain a more detailed view of the evolution of the biota in these regions, it is necessary to evaluate the effects of historical tectonic and climate events on species. Here, we analyzed the phylogeographic structure, historical demographic processes, and the contemporary period, Last Glacial Maximum (LGM) and Last Interglacial (LIG) ecological niche models of Quercus castanea, to infer the historical population dynamics of this oak distributed in the Mexican highlands. A total of 36 populations of Q. castanea were genotyped with seven chloroplast microsatellite loci in four recognized biogeographic provinces of Mexico: the Sierra Madre Occidental (western mountain range), the Central Plateau, the Trans‐Mexican Volcanic Belt (TMVB, mountain range crossing central Mexico from west to east) and the Sierra Madre del Sur (SMS, southern mountain range). We obtained standard statistics of genetic diversity and structure and tested for signals of historical demographic expansions. A total of 90 haplotypes were identified, and 29 of these haplotypes were restricted to single populations. The within‐population genetic diversity was high (mean h_S = 0.72), and among‐population genetic differentiation showed a strong phylogeographic structure (N_ST = 0.630 > G_ST = 0.266; p < .001). Signals of demographic expansion were identified in the TMVB and the SMS. The ecological niche models suggested a considerable percentage of stable distribution area for the species during the LGM and connectivity between the TMVB and the SMS. High genetic diversity, strong phylogeographic structure, and ecological niche models suggest in situ permanence of Q. castanea populations with large effective population sizes. The complex geological and climatic histories of the TMVB help to explain the origin and maintenance of a large proportion of the genetic diversity in this oak species.