Genetic variation for carbon isotope composition in Juglans regia L.: relationships with growth, phenology and climate of origin

? Among the traits of breeding interest for the common walnut tree Juglans regia L., characteristics such as timing of budbreak and leaf fall, water-use efficiency and growth performance are regarded as being of utmost relevance in Mediterranean conditions.

? The authors evaluated intraspecific variation in δ¹³C (carbon isotope composition, surrogate of intrinsic water-use efficiency, WUE_i) for 22 J. regia families grown in a progeny test under supplementary irrigation, and investigated whether such variation correlated with climatic indicators of native habitats. The genetic relationships between δ¹³C, growth and phenology were also assessed during two consecutive years.

? Overall, the most water-use-efficient families (i.e. with higher δ¹³C), which originated mainly from drought-prone provenance regions which have a high vapour pressure deficit and low rainfall, exhibited less height growth and smaller DBH. Using a stepwise regression procedure, δ¹³C was included as the main explanatory variable of genotypic variation in growth traits, together with growing season duration (for DBH in both years) and flushing (for height in 2007).

? It was concluded that WUE_i is largely unconnected to phenology effects in the explanation of growth performance for J. regia, therefore suggesting the opportunity of simultaneously selecting for low WUE_i and extended growing period to maximise productivity in non-water-limited environments.

     

Intraspecific variation in growth and allocation patterns in seedlings of Pinus pinaster Ait. submitted to contrasting watering regimes: can water availability explain regional variation?

? Interpopulation variation in key functional traits of Pinus pinaster Ait. is well recognized. However, the relative importance of drought tolerance to explain this regional variation in the species remains elusive.

? Here, we raise the question whether water availability constitutes a likely driver of regional variation in biomass allocation, growth and morphological traits of ten populations that cover the distribution range of P. pinaster. We carried out an experiment where seedlings of five families per population were submitted to two contrasting watering treatments.

? The effects of water availability and population were significant for relative diameter and height growth rate, biomass allocation and number of lateral stems and dwarf shoots. Total dry mass significantly differed between watering treatments but it did not among populations. Populations could be clustered into four main groups. Root mass fraction explained most of the variation and significantly correlated to altitude but not to aridity.

? The geographical pattern of genetic variation found in morphology and biomass allocation did not translate into population differences in drought tolerance or phenotypic plasticity to water availability, indicating that water availability is not a likely driver of the regional variation observed in the studied traits of P. pinaster at the seedling stage.

     

Genetic correlations between pulpwood and solid-wood selection and objective traits in Eucalyptus globulus

? If selective breeding is to be successful, significant genetic variation must be present in the traits targeted for improvement (i.e. “objective traits”).

? This study aimed to quantify genetic variation in Eucalyptus globulus pulpwood and sawn-timber objective traits (rotation-age whole-tree volume, survival, whole-tree basic density, sawn-board Janka hardness and sawn-board internal checking) and estimate additive genetic correlations between these and inexpensively-assessed “selection traits”.

? Significant genetic variation was identified in all objective traits at the subrace and/or family within subrace level.

? Selection-age diameter at breast height (1.3 m, DBH) was strongly genetically correlated with rotation-age volume (0.78) and survival (0.82). Subrace and additive genetic correlations of selectionage Pilodyn penetration with rotation-age 12-×-12-mm-sample basic density (?0.70 and ?0.75 respectively) and whole-tree basic density (?0.83 and ?0.91 respectively) were also strong.

? No significant subrace or additive genetic correlation between wood-sample gross shrinkage and sawn-board internal checking was detected. However, subrace and additive genetic correlations of sawn-board Janka hardness with Pilodyn penetration (?0.75 and ?0.58 respectively) and sample gross shrinkage (?0.77 and ?0.73 respectively) were significantly different from zero.

? These findings suggest that genetic improvement of the examined objective traits is possible through selective breeding, although none of the assessed selection traits were strongly correlated with internal checking.

     

Fecundity and larval development of Monochamus galloprovincialis (Coleoptera Cerambycidae) in experimental breeding

?The study aimed at clarifying basic life traits (fecundity, mortality and development) of Monochamus galloprovincialis, the vector of Bursaphelenchus xylophilus, the causative agent of the Pine Wilt Disease, in Portugal.

? Mating and oviposition experiments were conducted on P. sylvestris logs under laboratory conditions during two years. Larval development was followed outdoors.

? Fecundity was high, with 138.2 eggs per female.

? For the first time, the number of larval instars (4) in M. galloprovincialis was recognized and instars were described.

? Informations are provided on the shape and size of the larval galleries. Boring of galleries in sapwood and heartwood started at the third instar. All instars could overwinter but adults emerged simultaneously resulting in a univoltine life cycle for the majority of the individuals.

? An obligatory diapause in the forth instar was assumed. However, 8.1% of the insects had a two year development.

? Important differences in mortality were noted between the two experimental years, which could be explained by differences on the size of the breeding logs.

? There is a high biological proximity between M. galloprovincialis populations in France and those in Portugal and Northern Europe. However, a higher fecundity and longevity was observed in France, as well as a higher percentage of insects with a two year development, compared with portuguese populations.

     

Does ontogeny modulate irradiance-elicited plasticity of leaf traits in saplings of rain-forest tree species? A test with Dicorynia guianensis and Tachigali melinonii (Fabaceae, Caesalpinioideae)

? Irradiance elicits a large plasticity in leaf traits, but little is known about the modulation of this plasticity by ontogeny. Interactive effects of relative irradiance and ontogeny were assessed on leaf traits for two tropical rainforest tree species: Dicorynia guianensis Amshoff and Tachigali melinonii (Harms) Barneby (Fabaceae, Caesalpinioideae).

? Eleven morphological and physiological leaf traits, relative to photosynthetic performance, were measured on saplings at three different architectural development stages (ASD 1, 2 and 3) and used to derive composite traits like photosynthetic N-use efficiency. Measurements were made along a natural irradiance gradient.

? The effect of ASD was very visible and differed between the two species. For Dicorynia guianensis, only leaf mass-per-area (LMA) significantly increased with ASDs whereas for Tachigali melinonii, almost all traits were affected by ASD: LMA, leaf N content and photosynthetic capacity increased from ASD 1 to ASD 3. Photosynthetic N-use-efficiency was not affected by ASD in any species.

? Leaf traits were severely modulated by irradiance, whereas the degree of plasticity was very similar among ASDs. Only few interactions were detected between irradiance and ASD, for leaf thickness, carbon content, and the ratio Chl/N in T. melinonii and for photosynthetic capacity in D. guianensis.

? We conclude that ontogenic development and irradiance-elicited plasticity modulated leaf traits, with almost no interaction, i.e., the degree of irradiance-elicited plasticity was stable across development stages and independent of ontogeny in these two species, at least in the early stages of development assessed here.

     

Epicormics in 13-year-old Quercus petraea: small effect of provenance and large influence of branches and growth unit limits

? The provenance effect on epicormics is poorly documented and restricted to epicormic shoots. Our objective was to characterize the relationship between epicormics and provenance on 13-year-old Quercus petraea trees, taking into account their growth traits.

? On the growth units (GU) studied, epicormics were essentially isolated buds (95%) and they were mainly located at branch bases, secondarily on GU limits and marginally on lateral position. Provenance effect was very small on the number of epicormics and slightly larger on the number of branches and GU limits. On the contrary, the growth situation-independently from provenance-proved to have a larger effect on the 2 latter growth traits.

? Altogether, these results clearly show the predominant effect of morphological traits and thus of ontogeny. This opens important research perspectives: how growth conditions and genetics (at progeny and individual levels) influence height growth, setting and fate of axillary buds on the one hand and fate of epicormic buds inserted directly on the stem, or at branch bases on the other hand.

     

Genetic correlations among juvenile wood quality and growth traits and implications for selection strategy in Pinus radiata D. Don

? Juvenile wood quality in Pinus radiata is affected by factors such as low density, stiffness, and high microfibril angle, spiral grain, and shrinkage. Adverse genetic correlations between growth and wood quality traits remain as one of the main constraints in radiata pine advanced generation selection breeding program.

? Juvenile wood property data for this study were available from two progeny tests aged 7 and 6 y. We estimated the genetic correlations between stiffness, density, microfibril angle, spiral grain, shrinkage in the juvenile core and DBH growth in radiata pine, and) to evaluated various selection scenarios to deal with multiple objective traits.

? Negative genetic correlations were found for modulus of elasticity (MoE) and density with microfibril angle, spiral grain, shrinkage, and DBH. We observed low to moderate unfavourable genetic correlations between all wood quality traits and DBH growth.

? These low to moderate genetic correlations suggest that there may be some genotypes which have high DBH growth performance while also having high wood stiffness and density, and that the adverse correlation between DBH and MoE may not entirely prohibit the improvement of both traits. Results indicate that, in the short term, the optimal strategy is index selection using economic weights for breeding objective traits (MAI and stiffness) in radiata pine.

? In the long-term, simultaneously purging of the adverse genetic correlation and optimizing index selection may be the best selection strategy in multiple-trait selection breeding programs with adverse genetic correlations.

     

Impact of poplar water status on leaf-beetle (Chrysomela populi) survival and feeding

? The aim of this study is to assess the impact of drought on insect-tree relationships. Survival and feeding performances of leaf-beetles, Chrysomela populi put in cages with leaves of host-plants submitted to different levels of water stress were compared.

? Ninety 1-year-old poplar (Populus × euramericana) cuttings were grown in a greenhouse, and distributed into 3 groups submitted to different water treatments during six weeks. Adult C. populi were then encaged for four days with these cuttings, together with the five terminal leaves of a twig. Beetle survival and consumed leaf areas were then determined.

? Predawn leaf water potential values indicated that the drought treatments induced different levels of plant water stress. Plant height growth and total leaf numbers were reduced by plant stress. Beetle survival and leaf areas eaten were also reduced proportionally to plant water deficit.

? In conclusion, this experiment suggests that drought has a negative impact on C. populi survival and feeding.

     

Genetic variation in productivity, leaf traits and carbon isotope discrimination in hybrid poplars cultivated on contrasting sites

? We examined the relationships between productivity, leaf traits and carbon isotope discrimination in bulk leaf matter (Δ₁) and in phloem sap (Δ_s) from more than 5-year-old trees belonging to Populus deltoides × P. nigra and Populus trichocarpa × P. deltoides; trees were grown in alluvial and non alluvial sites in a commercial poplar plantation.

? On both sites, a large genetic variability was evidenced for all variables. The genotypic ranking remained stable between years for all variables, while it differed between sites. Δ₁ scaled positively with Δ_s and neither Δ₁ nor Δ_s were correlated with productivity. A significant genotype by site interaction was evident for all variables. The non alluvial site resulted in lower productivity, and in thicker/denser leaves with lower nitrogen and carbon contents. Noteworthy, the genotypic ranking for Δ₁ measured at the alluvial site was similar to that previously established in a glasshouse.

? As observed in previous studies from younger trees, there is a potential to select genotypes, combining high productivity and high water-use efficiency, for growth in moderately drought-prone areas.

     

Diversity of water use efficiency among Quercus robur genotypes: contribution of related leaf traits

? Previously, a large intra-specific diversity and a tight genetic control have been shown for Δ¹³C (carbon isotope discrimination) in a pedunculate oak (Quercus robur L.) family, which is an estimator for intrinsic water use efficiency (W _i), a complex trait defined as the ratio of net CO₂ assimilation rate (A) to stomatal conductance for water vapour (g _s ).

? In the present study, twelve genotypes with extreme phenotypic values of Δ¹³C were selected within this family to (i) asses the stability of genotype differences across contrasting environments and for different measures W _i; (ii) quantify the relationship between Δ¹³C and W_i within this family; (iii) identify which leaf traits drive the diversity in W _i observed in this family.

? Genetic variability of Δ¹³C and W _i was largely independent from different temporal integration scales and their correlation was found to be strong (R ² = 88% for leaf sugars) within this family.

? Weak correlations between measures of W _i with estimators of photosynthetic capacity, suggest a minor role of the latter in the diversity of W _i.

? However, the tight correlation between g _s and Δ¹³C as well as W _i, and the related genotypic variation in stomatal density, suggest that the genotypic diversity in W _i within this pedunculate oak family might be due to differences in g _s .

     

Compared effects of competition by grasses (Graminoids) and broom (Cytisus scoparius) on growth and functional traits of beech saplings (Fagus sylvatica)

? The effect of two weeds (grasses and broom, Cytisus scoparius) competition on the growth and functional traits of European beech saplings (Fagus sylvatica) was investigated in an experimental plantation in the French Massif central.

? We hypothesized that grasses would have a much more harmful effect than broom on beech growth through strong competition for soil water and nitrogen.

? A randomized block design was used with three separate blocks, each possessing three types of vegetation; grasses (mainly Agrostis capillaris, Holcus mollis and H. lanatus), broom, and bare soil as a control.

? Two years after plantation beech sapling growth was significantly greater in bare soil than with competition from grasses, and was intermediate on broom. Growth was positively correlated with beech leaf nitrogen content (N_m) and xylem water potential (Ψ_x). N and Ψ_x were dependent, with nitrogen uptake probably decreasing with increasing drought. There was no clear correlation between growth and light transmitted 10 cm below the apex of the saplings.

? Beech leaf mass on an area basis (LMA) was correlated with the three resource variables (water, nitrogen and light) and with growth, confirming that LMA is a sensitive functional trait integrating the degree of stress experienced by the saplings.

? The results suggest that broom is less competitive than grasses and, if properly managed on a sufficiently well-watered site, can generate satisfactory conditions for beech establishment and growth.

     

Ectomycorrhizal fungus (Paxillus involutus) and hydrogels affect performance of Populus euphratica exposed to drought stress

? Mycorrhizal fungi and hydrogels (water-absorbing polymers) can improve water availability for trees. The combination of both factors for plant performance under water limitation has not yet been studied.

? To investigate the influence of the ectomycorrhizal fungus Paxillus involutus, hydrogel and the combination of both factors, a drought-sensitive poplar, Populus euphratica, was examined in this study.

? After 16 weeks of inoculation, no ectomycorrhizas were found. Nevertheless, P. involutus-inoculated poplars displayed increased concentrations of soluble sugars and osmolality, leading to an improved water status. Growth was diminished compared with non-inoculated P. euphratica. The presence of hydrogels in the rooting medium resulted in increased biomass and higher plant water content and decreased the osmolality of plant tissues. Drought markedly decreased water contents in rooting medium and plants, and leaf chlorophyll fluorescence, and stimulated the root growth, concentrations of soluble sugars and osmolality in plants. Under drought conditions, P. euphratica exhibited osmoregulation by accumulation of low-molecular-weight carbohydrates.

? These data indicate that adding hydrogels to soils may improve the plant performance. The reasons for improved osmoregulation by fungi and hydrogels were probably related to their stimulating influence on the formation of soluble carbohydrates under drought conditions.

     

Genetic variation in architectural seedling traits of Patagonian cypress natural populations from the extremes of a precipitation range

? Species distributed along wide environmental ranges are expected to be either plastic or adapted to local optima. The elucidation of which of these alternatives prevails, is crucial in planning breeding and conservation strategies for not yet domesticated species. Austrocedrus chilensis (Cupressaceae) is the most commercially important conifer of the temperate forests of Argentina and the target of a domestication program. A steep precipitation gradient characterizes its Argentinean range.

? Variation within and differentiation among four natural populations of this Patagonian cypress representative of two contrasting precipitation regimes (>1 300 and 330 mm per y) were assessed by analyzing several morpho/architectural traits in one-year-old seedlings grown in a greenhouse.

? Progenies from one of the two xeric populations did not differ from those corresponding to both humid-site populations. The two most variable populations in terms of additive genetic variance belonged to contrasting precipitation regimes. Differentiation among populations was low as measured by each and every variable (average Q _st = 0.088). Morphological traits related to the main axis size would have a dubious adaptive meaning.

? The results suggest that the Patagonian cypress would have not evolved genetic pools adapted to local optima, and thus it appears to be a phenotypically plastic species, at least regarding growth at juvenile age.

     

Effects of genetic entry and competition by neighbouring trees on growth and wood properties of cloned Norway spruce (Picea abies)

? The aim of this work was to study both the effects of genetic entry and competition by neighbouring trees on growth, yield, wood density traits and fibre properties of 20 Norway spruce clones grown in an experimental trial located in southern Finland. The material included 10 Finnish clones, 2 Russian clones and 8 provenance-hybrids clones, the latter ones representing crosses between Finnish and foreign parent trees.

? We found that, compared to growth and yield, wood density traits and fibre properties showed, on average, lower phenotypic variations. Moreover, significant differences could be observed among the clones regardless of the trait (p < 0.05). Conversely, on average, no clear differences could be found between Finnish, Russian and provenance-hybrids clones in most of the studied traits.

? The marked differences among the clones, and lack of clear differences among the provenances suggested that any ranking, regarding different traits, should be based on individual clones. The phenotypic correlations between different traits ranged from moderate to strong suggesting that selection based on one trait would affect other traits. Regardless of clone, the growth of trees (e.g. breast height diameter) decreased and the wood density increased with increasing competition by neighbouring trees, which was expressed by competition index.

     

Integrating functional diversity into tropical forest plantation designs to study ecosystem processes

? The elucidation of relationships between biodiversity and ecosystem processes has been limited by the definition of metrics of biodiversity and their integration into experimental design. Functional trait screening can strengthen the performance of these designs.

? We suggest the use of Rao’s quadratic entropy to measure both functional diversity and phylogenetic diversity of species mixtures proposed for an experimental design, and demonstrate how they can provide complementary information.

? We also present an index assessing the statistical performance of these independent variables in different experimental designs. Measurement of independent variables as continuous vs. discrete variables reduces statistical performance, but improves the model by quantifying species differences masked by group assignments.

? To illustrate these advances, we present an example from a tropical forest tree community in which we screened 38 species for nine functional traits. The proposed TropiDEP design is based on the relative orthogonality of two multivariate trait axes defined using principal component analysis.

? We propose that independent variables describing functional diversity might be grouped to calculate independent variables describing suites of different traits with potentially different effects on particular ecosystem processes. In other systems these axes may differ from those reported here, yet the methods of analysis integrating functional and phylogenetic diversity into experimental design could be universal.

     

Seasonal changes in carbohydrates, cyclitols, and water relations of 3 field grown Eucalyptus species from contrasting taxonomy on a common site

? Alterations in plant chemistry underpin a suite of physiological adaptations to arid conditions. Qualitative and quantitative differences in leaf chemistry are found in the genus Eucalyptus correlating with physiological adaptation to aridity.

? Here we investigate seasonal water relations of three field grown eucalypt species grown at a common site known to differ in their ability to accumulate the cyclic sugar alcohol, quercitol.

? We show that quercitol contributes significantly to osmotic relations in field grown trees of Eucalyptus melliodora but is present only in trace amounts in E. rubida and E. obliqua.

? Measured concentrations of quercitol account for the difference in osmotic potentials between species and can be interpreted as a mechanism for adaptation to low water availability.

     

Large-scale structural variation and long-term growth dynamics of Juniperus thurifera trees in a managed woodland in Soria, central Spain

? Juniper woodlands have a great conservational and productive importance in Spain. However, basic traits of their structural variation and growth dynamics are unknown.

? To characterize the structural variation and assess both height and radial growth patterns, we sampled and measured stem cross-sections every 0.5 m in height from 107 Spanish juniper (Juniperus thurifera L.) trees, which were selected based on a 350 m regular grid over a whole woodland of 3 300 ha in area. We used dendroecological techniques and spatial analysis to study tree structure and growth.

? Structural traits such as bole diameter and height were mainly determined by tree life-related parameters such as tree age and growth rate variation, while abiotic factors such as altitude played a minor role in determining structural variation. Over the last 300 years, juniper establishment has been continuous in time but discontinuous in space. Large-scale spatial heterogeneity of tree establishment, and the presence of an early growth suppression and abrupt growth changes in junipers > 100 years old are consequences of an intense management that almost ceased in the late 19th century.

? Abrupt growth releases and suppressions were synchronic among different age classes, and coincided with documentary records on livestock decrease and key historical changes in land use. Growth patterns suggest that juniper is a brow sing-tolerant species that is able to survive large periods of intense browsing pressure.

     

Productivity of temperate broad-leaved forest stands differing in tree species diversity

? Understanding the effects of tree species diversity on biomass and production of forests is fundamental for carbon sequestration strategies, particularly in the perspective of the current climate change. However, the diversity-productivity relationship in old-growth forests is not well understood.

? We quantified biomass and above-ground production in nine forest stands with increasing tree species diversity from monocultures of beech to stands consisting of up to five deciduous tree species (Fagus sylvatica, Fraxinus excelsior, Tilia spp., Carpinus betulus, Acer spp.) to examine (a) if mixed stands are more productive than monospecific stands, (b) how tree species differ in the productivity of stem wood, leaves and fruits, and (c) if beech productivity increases with tree diversity due to lower intraspecific competition and complementary resource use.

? Total above-ground biomass and wood production decreased with increasing tree species diversity. In Fagus and Fraxinus, the basal area-related wood productivity exceeded those of the co-occurring tree species, while Tilia had the highest leaf productivity. Fagus trees showed no elevated production per basal area in the mixed stands.

? We found no evidence of complementary resource use associated with biomass production. We conclude that above-ground productivity of old-growth temperate deciduous forests depend more on tree species-specific traits than on tree diversity itself.

     

Modulation of leaf physiology by age and in response to abiotic constraints in young cuttings of two Populus deltoides × P. nigra genotypes

? It is of importance, when comparing physiological responses of leaves to environmental constraints among different genotypes, to take into account any effect related to leaf position and age within the canopy that might interfere with the response to the constraints.

? To document such effects, photosynthetic capacity and tolerance to heat and to oxidation were measured on leaves from the top to the bottom of three-month-old single-stem rooted cuttings of Populus deltoides × P. nigra genotypes, ‘Dorskamp’ and ‘Luisa_Avanzo’, thus taking into account a gradient of ages from youngest and still expanding (top) to oldest and fully expanded (bottom) leaves.

? All recorded traits were tightly linked to the age of the leaves. Photosynthetic capacity gradually increased during leaf expansion, in parallel with chlorophyll content and relative nitrogen allocation to RuBisCO and to bioenergetics. On the contrary, dark respiration gradually decreased during leaf expansion until a minimum value was reached at maturity. Compared to expanding leaves, young mature leaves were characterized by a lower sensitivity to heat and a higher one to oxidations generated by methyl-viologen.

? Leaf characteristics appeared to vary along the stem to a larger extent than between the two genotypes that display largely different productivities in plantations.

     

Methodology to assess both the efficacy and ecotoxicology of preservative-treated and modified wood

? Wood used in outdoor conditions out of ground contact is susceptible to weathering, inducing both fungal decay and leaching of components to the environment.

? This paper presents a methodology to determine these two parameters for untreated, preservative-treated and modified wood. Therefore, the wood was first leached and subsequently exposed to fungal decay of the most prominent wood-rotting fungi. The crustacean Daphnia magna was exposed to the leachates to provide information on their impact on the environment.

? Combining both parameters reveals that preservative-treated wood and modified wood are capable of protecting the wood adequately for application under use class 3 conditions without posing a threat to the environment.

? This proves the suitability of the concept of combining efficacy and ecotoxicology for the evaluation of new types of wood treatments.