Leaf morphological and genetic differentiation between <Emphasis Type="Italic">Quercus robur</Emphasis> L. and its closest relative,the drought-tolerant <Emphasis Type="Italic">Quercus pedunculiflora</Emphasis> K. Koch

• Introduction  

The study of phenotypic and genetic differentiation between incipient species or species that have recently diverged provides insights into the evolutionary history of species complexes and may contribute to our understanding of how species will evolve in contrasting environmental conditions.     

Altitudinal differentiation of <Emphasis Type="Italic">Quercus robur</Emphasis> in Bosnia and Herzegovina

Morphological variation based on eight measured and four derived traits was studied to establish whether there was significant variation between populations and to identify the influence of the altitude on morphological differentiation among 44 natural stands of common oak (Quercus robur L.) in Bosnia and Herzegovina, ranging from 82 to 860 m. The results point to significant intra- and inter-population differences. Elevation-related variation is less pronounced and determined largely by microclimatic factors. The results could provide management strategies for species reintroduction in the study area. There are no differences in leaf morphology between the different branches of individual trees. However, there are important variations between the altitudinal groups and closely linked to environmental factors in all traits. An important recommendation is to use seeds from oak with attention to their altitudinal origin.     

Oak (<Emphasis Type="Italic">Quercus robur</Emphasis> L.) regeneration as a response to natural dynamics of stands in European hemiboreal zone

The oak (Quercus robur L.) regeneration intensity was assessed in the core area of the Białowieża National Park (BNP) in Poland with respect to the selected ecological factors. The emphasis was placed on the response of oak regeneration to disturbances, including the large-scale dieback of spruce stands. Defining their effect could help predicting the role of oak in naturally developing lowland forest ecosystems in the European hemiboreal zone. The results of the study challenge the opinion that the ‘lime-oak-hornbeam forest’ is a ‘climax’ community, confirming a very poor regeneration represented by only two saplings taller than 0.5 m per hectare. By contrast, in spruce-dominated communities, from 49 to 848 taller saplings per hectare were found. The occurrence of saplings was associated with discontinuous canopy of late seral stage of stands, as well as with large gaps. Most of the best quality grown-up oak saplings developed in the immediate neighbourhood of spruce logs. The results of the research indicate that ‘lime-oak-hornbeam forest’ (Tilio-Carpinetum) should be rather perceived as a transient community, evolved from relict, culturally modified, oak woodlands. Because spruce had become a dominating species only after abandoning in mid-1800s the historical regime of anthropogenic disturbances (involving frequent forest fires), the observed phenomena related to the disintegration of spruce stands had probably no precedent over the last 500 years. To confirm whether the massive decline of spruce stands will finally result in the successful establishment of the new canopy oak generation, both in Białowieża and other forests of hemiboreal zone, further research is needed.     

Leaf-level responses to light in two co-occurring <Emphasis Type="Italic">Quercus</Emphasis> (<Emphasis Type="Italic">Quercus ilex</Emphasis> and <Emphasis Type="Italic">Quercus suber</Emphasis>): leaf structure,chemical composition and photosynthesis

We studied morphological, biochemical and physiological leaf acclimation to incident Photon-Photosynthetic-Flux-Density (PPFD) in Quercus ilex (holm oak) and Quercus suber (cork oak) at Mediterranean evergreen oak woodlands of southern Portugal. Specific leaf area (SLA) decreased exponentially with increasing PPFD in both species. Q. ilex had lower SLA values than Q. suber. Leaf nitrogen, cellulose and lignin concentration (leaf area-based) scaled positively with PPFD. Maximum rate of carboxylation (V_cmax), capacity for maximum photosynthetic electron transport (J_max), rate of triose-P utilization (V_TPU) and the rate of nonphotorespiratory light respiration (Rd) were also positively correlated with PPFD in both Quercus species, when expressed in leaf area but not on leaf mass basis. Q suber showed to have higher photosynthetic potential (V_cmax, J_max^m and V_TPU^m) and a higher nitrogen efficient nitrogen use than Q.ilex. Leaf chlorophyll concentration increased with decreasing PPFD, improving apparent quantum use efficiency (Φ) in both Quercus species. We concluded that, in Q.ilex and Q.suber, leaf structural plasticity is a stronger determinant for leaf acclimation to PPFD than biochemical and physiological plasticity.     

Introduced <Emphasis Type="Italic">Tuber aestivum</Emphasis> replacing introduced <Emphasis Type="Italic">Tuber melanosporum</Emphasis>: a case study

A Tuber melanosporum plantation established in 1994/1995 on Kibbutz Bar’am (in the Upper Galilee, Israel) gradually lost its T. melanosporum mycorrhiza. In 1999, T. aestivum inoculated seedlings were inadvertently introduced into the plantation to fill the gaps between trees. A single T. melanosporum fruit body was found in 1999. Although no truffles were found after 1999 and until 2009, in that year and in 2010, truffles were collected. Morphological and molecular analyses proved these to be T. aestivum. Thus, the intentionally introduced T. melanosporum mycorrhiza was replaced by that of another introduced mycorrhizal fungus, T. aestivum. Local oak species produced higher yields compared to introduced host species known to be good T. melanosporum plant symbionts. The yield was comparable to that reported for young commercial orchards, but the fruiting season was earlier than in Europe.     

Nitrogen budgeting and quality of exponentially fertilized <Emphasis Type="Italic">Quercus robur</Emphasis> seedlings in Ireland

Current policy in Ireland calls for increased quality and quantity of broadleaved seedling plantings. Simultaneously, concerns regarding environmental contamination necessitate development of nursery production methods that minimize impacts. Exponential (E) fertilization has potential for producing high-quality seedlings through nutrient loading, while simultaneously minimizing nitrogen (N) losses during culture. We investigated the effects of a wide range (0.10–1.0 g N seedling⁻¹ year⁻¹) of E fertilization rates on seedling morphology and nutrition of Quercus robur L., as well as soil N status and N budget during one growing season in a bareroot nursery in County Carlow, Ireland. The N budget showed significant N losses, especially in the highest fertilizer treatments, due to high precipitation. Seedlings receiving the 0.5 E rate were morphologically and nutritionally superior to all other treatments. Results indicate that it may be possible to produce 1-0 bareroot seedlings meeting target specifications in Ireland and other countries of similar climate and soils if further manipulations to seedling culture are implemented; however, potential environmental degradation via N leaching must be considered when N loading in wet years.     

Variation in the date of budburst in <Emphasis Type="Italic">Quercus robur</Emphasis> and <Emphasis Type="Italic">Q. petraea</Emphasis> across a range of provenances grown in Southern England

The widely observed advance in spring budburst across a range of temperate forest species due to climatic warming has received considerable attention. This change in forest phenology has important implications for the choice of species and provenances currently being planted, which need to be suited to both current and future climatic conditions. Using a provenance trial in the south of England, this study assessed variation in the timing of budburst across 23 different European provenances of oak (Quercus robur L. and Q. petraea (Matt.) Liebl.) over 8 years of observations. The order in which the different provenances reached budburst was related to provenance source latitude: the southern provenances were always earlier than those from more northerly latitudes. The statistical technique partial least squares regression was used to identify critical periods of both chilling and warming. A General Linear Model and three-dimensional temperature response surfaces were used to analyse the temporal trends in budburst. There was a negative correlation between the date of budburst and mean daily air temperature in both the chilling and warming periods for all provenances, which was statistically significant for a majority. Spring warming had a larger effect on budburst than winter chilling with a mean spring temperature-driven advance of 3.61 days/°C (standard error = 0.17 days/°C) and mean winter period temperature-driven advance of 0.99 days/°C (standard error = 0.17 days/°C). Surprisingly, there was no statistically significant interaction between mean air temperatures during the chilling and warming phases on the date of budburst.     

Temporal changes in the spatial pattern of leaf traits in a <Emphasis Type="Italic">Quercus robur</Emphasis> population

Background  

Plant resources in natural ecosystems are frequently spatially structured at a scale relevant to individuals. This spatial structure can be variable in time, and can even vary within a single growing season. Several leaf traits may show spatial structure at the same scale as light or soil resources. However, whether this spatial structure stays constant from one growing season to the next is still unknown.     

Phenotypic plasticity blurs ecotypic divergence in the response of <Emphasis Type="Italic">Quercus coccifera</Emphasis> and <Emphasis Type="Italic">Pinus halepensis</Emphasis> to water stress

The Mediterranean evergreen woody plants Quercus coccifera and Pinus halepensis grow in a range of environments where selection by drought, heat and high irradiance can drive genetic and phenotypic differentiation of populations. However, the role of these stresses in filtering out maladaptive genotypes remains unknown. We hypothesize that this filtering is an important process for woody Mediterranean species due to their low phenotypic plasticity reported in previous studies. We have studied the response of saplings of Q. coccifera and P. halepensis, originating from two contrasting populations (a rock outcrop and a garrigue formation), to water stress. Isozyme characterization of genetic diversity was done to determine whether populations were genetically distinct. Water response analysis was based on water relations, gas exchange, chlorophyll a fluorescence, pigment content, antioxidant status and morphological and structural parameters. Ecotypic differentiation was found for both Q. coccifera and P. halepensis populations, with a higher population isozyme similarity and a higher frequency of dominance of a few genotypes at the rock outcrop in both the species. P. halepensis exhibited small but significant differences between populations for plastic responses to water, with lower phenotypic plasticity in saplings from the rock outcrop. Although it was not found in Q. coccifera, this pattern suggests that ecotypic differentiation rendering stress-tolerant ecotypes involves a decreased plasticity. Phenotypic plasticity was not high but it explained over 75% of the total variability among individual plants. Thus, and although evidence for ecotypic divergence was found in both the species, saplings were plastic enough to blur ecotypic differentiation. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Spatial pattern of beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) and oak (<Emphasis Type="Italic">Quercus pubescens</Emphasis> Mill.) seedlings in natural pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) woodlands

Spatial pattern of recruitment is an important factor influencing population dynamics of plant communities. The spatial pattern is determined by seed dispersal and by the spatial variability of germination and initial survival. In the process of forest expansion following farmland abandonment, mid- and late-successional species are often dispersed in pioneer forests by birds. This could result in an aggregated spatial pattern of seeds that could influence the dynamics of these species in mixed pioneer forests. In the sub-Mediterranean area, mid- and late-successional species such as Quercus pubescens (downy oak) and Fagus sylvatica (European beech) are expected to replace pioneer Pinus species. Using a point sampling method we demonstrated that beech and oak seedlings (height <2 m) have a clumped distribution in the understorey of pine. This could result from an aggregated dispersal by jays (dispersal effect) or from preferential recruitment in particular habitats (habitat effect). To test these hypotheses we proposed a statistical analysis of spatial patterns of regeneration of beech and oak. Ground cover variables (i.e. cover by rock outcrops, herbs, box shrubs, mosses or pine) did not differ significantly around seedlings as compared with random sample plots. Likewise, clumped seedlings had growth similar to isolated seedlings, thus refuting the hypothesis of preferential location in the most favourable microsites. Aggregated dispersal seems to be involved in the process of regeneration. Since beech and oak seedlings have contrasting ecological demands, we discuss the implication of this pattern for the replacement dynamics of pine by these species.     

Successful <Emphasis Type="Italic">Agrobacterium</Emphasis>-mediated transformation of <Emphasis Type="Italic">Populus tomentosa</Emphasis> with apple <Emphasis Type="Italic">SPDS</Emphasis> gene

The problem of salinized soils has become one of the most serious constraints to agricultural and forest productivity. With the purpose of enhancing salt stress tolerance of Populus tomentosa, we transformed this tree species with spermidine synthase （SPDS） genes derived from an apple by an Agrobacterium-mediated method. Four transgenic clones were confu＇med by PCR and Southern blot analysis. As well, the expression of introduced SPDS genes was analyzed by real-time quantitative PCR.     

Influence of water-stress acclimation and <Emphasis Type="Italic">Tuber melanosporum</Emphasis> mycorrhization on <Emphasis Type="Italic">Quercus ilex</Emphasis> seedlings

Mycorrhizal and non-mycorrhizal holm oak (Quercus ilex L.) seedlings inoculated with black truffle (Tuber melanosporum) were grown under nursery conditions and subjected to drought hardening for 4 months in autumn and winter followed by irrigation for 10 days. Leaf water potential and stomatal conductance were monitored during the 4 months of drought. When the test was completed (March), measurements were made for each treatment (inoculated or non-inoculated), and watering regime (watered and water-stressed). Pressure–volume curves, osmotic potential at full turgor, osmotic potential at zero turgor and the tissue modulus of elasticity near full turgor were calculated. Mycorrhizal colonization and growth, and the content of the main mineral nutrients N, P, K, Ca and Mg were measured. Water stress affected plant growth, caused an elastic adjustment of the plant tissues, and decreased the P and K content, and inoculation improved the nitrogen content. Drought acclimation apparently achieved the goal of improving the drought tolerance of holm oak seedlings, without depressing ectomycorrhizal root colonization by T. melanosporum. José Antonio Rodríguez Barreal—Deceased     

Site quality of pedunculate oak (<Emphasis Type="Italic">Quercus robur</Emphasis> L.) stands in Galicia (northwest Spain)

Accurate estimates of forest productivity are required for sustainable forest management. Height of dominant and codominant trees at a reference age is often used as a measure of site productivity. Eight algebraic difference equations based on the models proposed by Sloboda, Bertalanffy-Richards, Korf, Hossfeld, and McDill-Amateis were tested on the transformed, longitudinal data structure that considers all possible growth intervals. Autocorrelation was modelled by expanding the error term as an autoregressive process according to this data structure. Generalized nonlinear least squares methods were used for model fitting. Several numerical and graphic analyses were used to compare the different candidate models. The relative error in dominant height prediction was used to select 60 years as the best reference age. An algebraic difference equation based on the model proposed by Korf provided the best compromise between biological and statistical aspects and produced the most adequate site index curves. This model is therefore recommended for height growth prediction and site classification of pedunculate oak stands in Galicia. The model is polymorphic and base-age invariant, having only one asymptote. Predictions of height for age intervals between t₁ and t₂ of more than 15 years should be considered with caution because of the associated critical error.     

<Emphasis Type="Italic">In vitro</Emphasis> morphogenic response of leaf sheath of <Emphasis Type="Italic">Phyllostachys bambusoides</Emphasis>

Nodal segments from secondary branches of saplings of Phyllostachys bambusoides were inoculated in MS medium to assess the in vitro morphogenic response of leaf sheath through the induction to callogenesis by Picloram(4-amino-3,5,6-trichloropicolinic acid) at different concentrations of carbohydrate under the same conditions with presence or absence of luminosity.In our experiment,secondary explants were kept in MS medium containing 8.0 mg·L-1 of Picloram for the callus formation.Calluses were transferred in MS medium supplemented with sucrose,fructose and glucose(control,2%,4% and 6%).Results show that Picloram induced the callogenesis in leaf sheath.The secondary embryogenesis was formed in yellow-globular callus.The sucrose as carbohydrate source in the absence of light was more efficient to induce rhizogenesis.Glucose was more efficiency in the presence of light.Callogenic induction and further embryogenesis evidenced the competence and determination of leaf sheath cells.     

At5g54160 gene encodes <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> 5-hydroxyconiferaldehyde <Emphasis Type="Italic">O</Emphasis>-methyltransferase

The function of an Arabidopsis thaliana gene, At5g54160 annotated as a caffeic acid O-methyltransferase CAOMT gene was characterized. The recombinant enzyme of this gene (AtOMT1) catalyzed the O-methylation of phenylpropanoid and flavonoid substrates. The specificity constants (k _cat/K _m) for 5-hydroxyconiferaldehyde (5-HCAld) and quercetin were both 0.11 μM⁻¹·min⁻¹. On the other hand, lignins of At5g54160-knockout Arabidopsis mutants lacked syringyl units. In addition, we showed that the gene silencing also resulted in significant accumulation of caffeyl alcohol (CaAlc). These results strongly suggested that At5g54160 gene is involved in syringyl lignin synthesis for the methylation of both 5-hydroxyconiferaldehyde and 3,4-dihydroxyphenyl compound(s). Part of this work was presented at the Annual Meeting of Japan Society for Bioscience, Biotechnology, and Agrochemistry, March 24–27, 2007     

Potential contribution of <Emphasis Type="Italic">Quercus durata</Emphasis> and <Emphasis Type="Italic">Adenostoma fasciculatum</Emphasis> supplemented with <Emphasis Type="Italic">Medicago sativa</Emphasis> on intake and digestibility in sheep and goats

A digestion trial was conducted to estimate the potential contribution of the chaparral browse Quercus durata (leather oak) and Adenostoma fasciculatum (chamise) on intake and nutrient digestion in goats and sheep. Four wether Kiko goats (avg. wt. 22.9 kg) and four wether Targhee sheep (avg. wt. 39.6 kg) were housed in metabolism cages. Alfalfa pellets were used as a dietary supplement. Digestibility of the shrubs was measured by difference and metabolizable energy (ME) and ME intake (MEI) were calculated from digestibility and plants intake values. Oak and chamise had low crude protein (CP) content (6.5 and 4.3% DM) and high content of extractable condensed tannins (17.8 and 9.8% DM). In vitro organic matter digestibility (OMD) and ME for oak and chamise were 35.6 and 33.1% and 5.0 and 4.7 MJ/kg DM, respectively. In vivo OMD (calculated by difference with alfalfa) and MEI, for browse were greater (P < 0.01) for goats fed chamise compared to sheep. In oak, OMD was similar for both animal species but MEI was greater (P < 0.01) for goats. Greater (P < 0.05) DM and CP intakes (per kg BW^0.75) were observed for goats fed either oak or chamise compared with sheep. When an energy source was given as supplement, the potential contribution of chamise and oak to total ME requirements for maintenance was about 58.3 and 60.4% in goats and about 17.1 and 32.9% in sheep, respectively. In consequence, metabolizable energy supplied by these shrubs may not be adequate for maintenance when, during grazing, they are consumed in high proportions with other chaparral plants that may have lower nutritional value than the alfalfa supplemented in this study. Under an appropriate supplementation program, goats could be more effective than sheep to control fuel load in California chaparral, as they consume more chamise and leather oak and obtain more nutrients from them.     

Insecticidal properties of <Emphasis Type="Italic">Thymus persicus</Emphasis> essential oil against <Emphasis Type="Italic">Tribolium castaneum</Emphasis> and <Emphasis Type="Italic">Sitophilus oryzae</Emphasis>

Red flour beetle Tribolium castaneum (Herbst) and rice weevil Sitophilus oryzae (L.) are considered to be the major insect pests in storage. Essential oils from aromatic plants are recognized as proper alternatives to fumigants. Thymus persicus (Ronniger ex Rech. f.) is one of these plants that have medicinal properties and is indigenous to Iran. The essential oil was obtained from aerial parts of the plant and analyzed by GC and GC–MS. Carvacrol (44.69%) and thymol (11.05%) were the major constituents of the oil extracted. In this experiment, fumigant toxicity of the essential oil was studied against T. castaneum, S. oryzae at 27 ± 1°C and 60 ± 5% RH in dark condition. The adult insects were exposed to the concentrations of 51.9, 111.1, 207.4 and 370.4 μl/l air to estimate median lethal time (LT₅₀) values. The fumigant toxicity was increased in response to increased essential oil concentrations. The LT₅₀ values at the lowest and the highest concentrations tested were ranged from 28.09 to 13.47 h for T. castaneum, and 3.86 to 2.30 h for S. oryzae. It was found that S. oryzae adults were much more susceptible to the oil than T. castaneum. After 24 h of exposure, the LC₅₀ values (95% fiducial limit) for T. castaneum and S. oryzae were estimated to be 236.9 (186.27–292.81) and 3.34 (2.62–4.28) μl/l air, respectively. These results suggest that T. persicus essential oil merits further study as potential fumigant for the management of these stored-product insects.     

Artificial regeneration with <Emphasis Type="BoldItalic">Quercus ilex</Emphasis> L. and <Emphasis Type="BoldItalic">Quercus suber</Emphasis> L. by direct seeding and planting in southern Spain

 Introduction   

The limited ability of Quercus species to regenerate naturally in Mediterranean forests has led to the development of various artificial regeneration methods; however, there is no general consensus as to what specific method is the best one for this purpose.     

<Emphasis Type="Italic">Quercus suber</Emphasis> forest and <Emphasis Type="Italic">Pinus</Emphasis> plantations show different post-fire resilience in Mediterranean north-western Africa

Key message

In the African rim of the Western Mediterranean Basin, cork oak forests and pine plantations coexist. Under similar fire regimes, cork oak forest is more resilient in terms of habitat structure (canopy, understory, and complexity of vegetation strata) than pine plantation. By contrast, both woodland types show similar resilience in plant species composition. Resilience in habitat structure varies between the two woodland types because of the resprouting and seeding strategies of cork oak and pine species, respectively. These differences can be relevant for the conservation of biodiversity of forested ecosystems in a future scenario of increased fire frequency and scale in the Mediterranean basin.

Context

Wildfires have major impacts on ecosystems globally. In fire-prone regions, plant species have developed adaptive traits (resprouting and seeding) to survive and persist due to long evolutionary coexistence with fire. In the African rim of the Western Mediterranean Basin, cork oak forest and pine plantation are the most frequently burnt woodlands. Both species have different strategies to respond fire: cork oak is a resprouter while pines are mostly seeders.

Aims

We have examined the hypothesis that pine plantations are less resilient in habitat structure (canopy, understory, diversity of vegetation strata) and plant composition than cork oak woodlands.

Methods

The habitat structure and plant species composition were measured in 30 burnt and 30 unburnt 700-m transects at 12 burnt sites from north-western Africa, where the two forest types can coexist. Habitat structure and plant species composition were compared between burnt and unburnt transects from cork oak and pine plantation woodlands with generalized linear mixed models and general linear models.

Results

The results showed significant interaction effect of fire and forest type, since cork oak forest was more resilient to fire than was pine plantation in habitat structure. By contrast, both forest types were resilient to fire in the composition of the plant communities, i.e., plant composition prior to fire did not change afterwards.

Conclusion

The higher structural resilience of cork oak forest compared to pine plantation is related to the resprouting and seeding strategies, respectively, of the dominant tree species. Differences in the responses to fire need to be considered in conservation planning for the maintenance of the Mediterranean biodiversity in a future scenario of changes in fire regime.

     

Interfertile oaks in an island environment. II. Limited hybridization between <Emphasis Type="Italic">Quercus alnifolia</Emphasis> Poech and <Emphasis Type="Italic">Q. coccifera</Emphasis> L. in a mixed stand

Hybridization and introgression between Quercus alnifolia Poech and Q. coccifera L. is studied by analyzing morphological traits, nuclear and chloroplast DNA markers. The study site is a mixed stand on Troodos Mountains (Cyprus), and the analyzed material includes both adult trees and progenies of specific mother trees. Multivariate analysis of morphological traits shows that the two species can be well distinguished using simple leaf morphometric parameters. A lower genetic diversity in Q. alnifolia than in Q. coccifera and a high interspecific differentiation between the two species are supported by an analysis of nuclear and chloroplast microsatellites. The intermediacy of the four designated hybrids is verified by both leaf morphometric and genetic data. Analysis of progeny arrays provides evidence that interspecific crossings are rare. This finding is further supported by limited introgression of chloroplast genomes. Reproductive barriers (e.g. asynchronous phenology, postzygotic incompatibilities) might account for this result. A directionality of interspecific gene flow is indicated by a genetic assignment analysis of effective pollen clouds with Q. alnifolia acting as pollen donor. Differences in flowering phenology and species distribution in the stand may have influenced the direction of gene flow and the genetic differentiation among effective pollen clouds of different mother trees within species.