Scarce population genetic differentiation but substantial spatiotemporal phenotypic variation of water-use efficiency in <Emphasis Type="Italic">Pinus sylvestris</Emphasis> at its western distribution range

Water and carbon fluxes in forests are largely related to leaf gas exchange physiology varying across spatiotemporal scales and modulated by plant responses to environmental cues. We quantified the relevance of genetic and phenotypic variation of intrinsic water-use efficiency (WUE_i, ratio of net photosynthesis to stomatal conductance of water) in Pinus sylvestris L. growing in the Iberian Peninsula as inferred from tree-ring carbon isotopes. Inter-population genetic variation, evaluated in a provenance trial comprising Spanish and German populations, was low and relevant only at continental scale. In contrast, phenotypic variation, evaluated in natural stands (at spatial level) and by tree-ring chronologies (at temporal inter-annual level), was important and ten- and threefold larger than the population genetic variance, respectively. These results points to preponderance of plastic responses dominating variability in WUE_i for this species. Spatial phenotypic variation in WUE_i correlated negatively with soil depth (r?=???0.66; p?<?0.01), while temporal phenotypic variation was mainly driven by summer precipitation. At the spatial level, WUE_i could be scaled-up to ecosystem-level WUE derived from remote sensing data by accounting for soil water-holding capacity (r?=?0.63; p?<?0.01). This outcome demonstrates a direct influence of the variation of leaf-level WUE_i on ecosystem water and carbon balance differentiation. Our findings highlight the contrasting importance of genetic variation (negligible) and plastic responses in WUE_i (large, with changes of up to 33% among sites) on determining carbon and water budgets at stand and ecosystem scales in a widespread conifer such as Pinus sylvestris.     

Response of Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescens Willd.) to soil and atmospheric water deficits under Mediterranean mountain climate

The physiological responses to water deficits of Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescens Willd.) were studied under Mediterranean mountain climate. Minimum leaf water potentials were ?3.2 MPa for oak and ?2.1 MPa for pine, with higher predawn values for pubescent oak. Relative sap flow declined in both species when vapour pressure deficit (D) went above ca. 1.2 kPa, but stomatal control was stronger for pine during the 2003 summer drought. P. sylvestris plant hydraulic conductance on a half-total leaf area basis (k _L,s?1) was 1.2–2.6 times higher than the values shown by Q. pubescens, and it showed a considerably steeper decrease during summer. Leaf-level gas exchange was positively related to k _L,s?1 in both species. Scots pine was more vulnerable to xylem embolism and closed stomata to prevent substantial conductivity losses. The results of this study confirm that pubescent oak is more resistant to extreme drought events.     

Species selection in hardwoods research: variations in baseline physiological responses of select temperate hardwood tree species

Drought periods are becoming more extreme worldwide and the ability of plants to contribute towards atmospheric flux is being compromised. Properly functioning stomata provide an exit for water that has been absorbed by the roots, funneled into various cell parts, and eventually released into the atmosphere via transpiration. By observing the effects that weather conditions such as climate change may have on stomatal density, distribution, and functioning, it may be possible to elucidate a portion of the mechanisms trees use to survive longer periods of water stress. This study analyzed stomatal density (SD), stomatal conductance (gs ), CO2 assimilation (A), instantaneous water-use efficiency (WUEi ), and transpiration (E) rates in six native tree species in the Midwestern USA and showed that trees within the same ecotype followed similar trends, but that trees within the same family did not when exposed to identical greenhouse conditions. Naturally drought tolerant tree species demonstrated lower g s and higher WUEi , while intolerant species had higher SD. This study showed negative or no correlation between SD and g s , A, E, and WUEi and positive correlations between E and A and gs and E.     

水分胁迫对四倍体刺槐苗生长和光合作用的影响

刺槐是我国三北地区的主要防护造林树种,尤其是西北干旱、半干旱地区有广泛的种植;但由于当地气候条件的制约,刺槐难于获得生长所需的足够水分[1],生长受到严重抑制,在黄土高原出现了大面积的"小老树"和低劣残次林[2-4],其生态、经济效益都很低.     

Growth,biomass allocation,and water use efficiency of 31 apple cultivars grown under two water regimes

Plant growth, biomass allocation, carbon isotope composition (δ¹³C), and water use efficiency (WUE) of 31 cultivars of apple (Malus domestica Borkh.) grown under two water regimes were measured. Drought-stressed plants showed significant declines in tree height, trunk diameter, biomass production, and total leaf area, the extent to which depended upon cultivar. Also, gas exchange rates, instantaneous and long-term efficiencies (WUE_I and WUE_L, respectively), and values for δ¹³C differed among cultivars and watering regimes. Variations in WUE_I were mainly due to changes in stomatal conductance (g _s) under drought condition. ‘Qinguan’ and ‘Golden Delicious’ had greater trunk diameter, tree height, and had higher biomass production and WUE_L under drought stress, implying that they are more suitable for arid and semi-arid regions. Moreover, WUE_L was significantly and positively correlated with δ¹³C under two watering regimes, which suggests a potential for evaluating water use efficiency of Malus by measuring carbon isotope composition.     

Drought effects on water use efficiency, freezing tolerance and survival of Eucalyptus globulus and Eucalyptus globulus × nitens cuttings

Genetic improvement of Eucalyptus genotypes for drought and frost resistance is essential for successful intensive management of commercial plantations. Understanding the physiological mechanisms that relate water use and frost resistance for highly deployed genotypes may allow for better prediction of their future performance, genetic selection and seedling management for site specific purposes. We studied whether instantaneous water use efficiency (WUE _i ) may serve as drought, freezing and photoinhibition tolerance predictor by studying its response on six E. globulus clones (Eg1–Eg6) and four E. globulus × E. nitens hybrid seedlings (Egn1–Egn4) under drought and irrigated (control) treatments. Net photosynthesis (A) and transpiration (E) were studied using a gas exchange system in order to calculate WUE _i (A/E). Simultaneous chlorophyll a fluorescence measurements were performed to assess the non photochemical quenching components. Frost tolerance of plants under control and drought treatments were evaluated by measuring temperatures that exert 50% photoinactivation of photosystem II. Finally, drought tolerance was evaluated by plant survival within each genotype after rehydration. Our results showed significant genotype variability in the rate of soil and xylem water potential decrease during drought. While most of the genotypes reached ?4.0 MPa in about 35 days of drought, genotypes Eg6 and Egn4 required 56 days of drought to reach this xylem water potential. WUE _i exhibited significant differences among genotypes and irrigation treatments. Genotypes Eg5 and Egn4 increased their WUE _i between 70 and 80% after drought. This was associated with a more conservative control of water loss at the stomatal level combined with maintenance of relatively higher rates of net photosynthesis than the other genotypes under drought conditions. Plants exposed to drought were more freezing tolerant than control plants, having in average 3°C lower LT₅₀ than well irrigated ones. There was no a clear correlation between WUE _i and drought tolerance or drought-induced photoinhibition, however WUE _i was inversely correlated with LT₅₀. Our results suggest that WUE _i is not suitable by itself to select drought tolerant genotypes, but may provide evidence for discarding drought sensitive genotypes. In addition, it could provide valuable information to select for freezing tolerance.     

Browsing Damage on Pine (Pinus sylvestris and P. contorta) by a migrating moose (Alces alces) Population in Winter: Relation to Habitat Composition and Road Barriers

In Scandinavia, moose (Alces alces L.) sometimes cause severe browsing damage to economically-important pine. Moose-vehicle accidents have spurred construction of fences along roads, and these may interfere with moose migration between summer and winter ranges, or the road alone may be a barrier. If this happens and moose build up along roads, landowners may suffer economically. Therefore, this study investigated whether roads, fences or other factors influence the use of young pine stands by moose. Eighty stands along roads in northern Sweden were evaluated in which individually-browsed branches were counted on 9972 pines. Moose browsing was not significantly related to birch (Betula pendula Roth, B. pubescens Ehrh.) density, nor did it differ between pines (Pinus contorta Douglas or P. sylvestris L.). However, increased pine density, site productivity and proximity to a highway were associated with increased browsing. Further large-scale studies are needed to understand moose habitat selection and the effects of roads.     

Influence of drought on tree rings and tracheid features of Pinus nigra and Pinus sylvestris in a mesic Mediterranean forest

We investigated the influence of climate on the ring width and xylem anatomy of two co-occurring pines (Pinus nigra Arn. and P. sylvestris L.) in the mountains of east-central Spain in order to test their utility for dendroclimatic reconstructions. We developed chronologies of ring width, mean lumen diameter and mean cell-wall thickness (in the earlywood, latewood, and the total annual ring) and the number of cells between 1960 and 2006. Drought, expressed as the standardized precipitation-evapotranspiration index (SPEI), was the main climatic driver of tree radial growth, although trees were also sensitive to temperature (negative effect in previous autumn and current summer) and precipitation (with a general positive effect). P. sylvestris response was stronger to climate of the current year, whereas the effect of previous-year climate was more important for P. nigra. Warm and dry summers reduced ring width, tracheid lumen, and wall thickness in both species, whereas warm winter-spring temperatures had the opposite effect, primarily for P. sylvestris. Previous-year or early-season conditions mainly affected earlywood features, whereas latewood was more responsive to summer climate. Overall, climate appeared to be a stronger limiting factor for P. sylvestris. During periods of drought, cell-wall thickness was reduced while lumen width increased in the latewood of P. sylvestris. This could compromise its hydraulic safety against drought-induced cavitation as our site was close to the southern and dry edge of the species distribution area. Our results suggest that anatomical variables record different and stronger climate information than ring width variables, especially in P. sylvestris. Reconstruction models for SPEI at the 3-month scale were developed for July–August and September–October using principal components regression. The best models included anatomical and width variables of both pine species suggesting that tracheid chronologies can be useful for drought reconstructions especially at mesic sites or with species that encode a mixed drought and temperature-precipitation signal.     

Intraspecific variation in drought response of Populus cathayana grown under ambient and enhanced UV-B radiation

? The effects of drought, enhanced UV-B radiation and their combination on plant growth and physiological traits were investigated in a greenhouse experiment in two populations of Populus cathayana Rehder originating from high and low altitude in south-west China.

? In both populations, drought significantly decreased biomass accumulation and gas exchange parameters, including net CO₂ assimilation rate (A), stomatal conductance (g _s ), transpiration rate (E) and photosynthetic nitrogen use efficiency (PNUE). However, instantaneous water use efficiency (WUE _i ), transpiration efficiency (WUE _T ), carbon isotope composition (δ¹³C) and nitrogen (N) content, as well as the accumulation of soluble protein, UV-absorbing compounds and abscisic acid (ABA) significantly increased in response to drought. On the other hand, cuttings from both populations, when kept under enhanced UV-B radiation, showed very similar changes, as under drought, in all above-mentioned parameters.

? Compared with the low altitude population, the high altitude population was more tolerant to drought and enhanced UV-B, as indicated by the higher level of biomass accumulation, gas exchange, water-use efficiency, ABA concentration and UV-absorbing compounds.

? After one growing season of exposure to different UV-B levels and watering regimes, the decrease in biomass accumulation and gas exchange, induced by drought, was more pronounced under the combination of UV-B and drought. Significant interactions between drought and UV-B were observed in WUE _i , WUE_T, δ¹³C, soluble protein, UV-absorbing compounds, ABA and in the leaf and stem N, as well as in the leaf and stem C:N ratio.

? Our results showed that UV-B acts as an important signal allowing P. cathayana seedlings to respond to drought and that the combination of drought and UV-B may cause synergistically detrimental effects on plant growth.

     

Effect of cleaning methods in mixed pine–deciduous stands on moose damage to Scots pines in southern Finland

Abstract

The effect of different cleaning treatments on moose (Alces alces L.) damage to Scots pines (Pinus sylvestris L.) was examined using a generalized linear mixed model (GLMM) with a binomial response (logit-link function). The treatments were the following: no cleaning, point-cleaning of broadleaves (mainly birch, Betula pendula Roth and B. pubescens Ehrh.) within a radius of 1 m from the pine, point-cleaning delayed 2 years, topping of competing broadleaves (main stem cut at a height of 1.3 m) and total cleaning of broadleaves. A randomized complete block design with three replicates was established in three planted pine stands (mean height of pines 1.5 m). Measurements taken four growing seasons later (mean height of pines 3.5 m) showed that all cleaning treatments reduced moose damage, although the difference was not significant (p=0.078) in the case of topping. The proportion of damaged pines was predicted as a function of the characteristics of pines and broadleaves in two untreated, planted pine stands with a height of 3 m. Within these untreated stands the number of damaged pines increased in plots with a higher number of broadleaves and taller broadleaves. In conclusion, the results show that on relatively fertile growing sites of pine cleaning treatments reduced moose damage on pine. It is recommended that cleaning is performed before broadleaved trees occur as overgrowth above pines.     

Foliar Carbon Isotope Composition (δ 13C) and Water Use Efficiency of Different Populus deltoids Clones Under Water Stress

Foliar carbon isotope composition (δ ¹³C), total dry biomass, and long-term water use efficiency (WUE_L) of 12 Populus deltoids clones were studied under water stress in a greenhouse. Total dry biomass of clones decreased greatly, while δ ¹³C increased. Single-element variance analysis in the same water treatment indicated that WUE_L difference among clones was significant. Clones J₂, J₆, J₇, J₈, and J₉ were excellent with high WUE_L. Extremely significant δ ¹³C differences among water treatments and clones were revealed by two-element variance analysis. Water proved to be the primary factor affecting δ ¹³C under water stress. It showed that there was a good positive correlation between δ ¹³C and WUE_L in the same water treatment, and that a high WUE_L always coincided with a high δ ¹³C. δ ¹³C might be a reliable indirect index to estimate WUE_L among P. deltoids clones. Translated from Scientia Silvae Sinicae, 2005, 41(1) (in Chinese)     

One-hundred-year foliage comparison of Pinus ponderosa and Pinus sylvestris under dry growing conditions in Brandenburg,Germany

Abstract

The dynamics of six different needle parameters of Pinus sylvestris L. and Pinus ponderosa Dougl. ex P. & C. Laws. were examined retrospectively for a 112-year-old mixed stand in Brandenburg, Germany, using the needle trace method. Similarities were found in needle production, needle loss and needle density. However, needle age, needle retention and total number of needles revealed significant differences between the tree species, with higher values for P. ponderosa. Pinus ponderosa yielded approximately twice as much mean whole-crown needle dry mass as P. sylvestris. Furthermore, different branching systems could be detected between the species, with both pines following “Corner's rule”. The results suggest that under identical growing conditions, P. ponderosa exhibits more efficient water use and can therefore maintain a bigger crown (as the basis for increased growth) than P. sylvestris.     

Development of Pinus sylvestris Main Stems Following Three Different Precommercial Thinning Methods in a Mixed Stand

Reducing competition by point cleaning may be an attractive alternative to conventional precommercial thinning in forestry. In this study, survival and development of main stems of Scots pine (Pinus sylvestris L.) following point cleaning were examined in a mixed stand of mainly Scots pine and birch (Betula pendula Roth and Betula pubescens Ehrh.) in northern Sweden. Treatments studied were: no precommercial thinning, conventional precommercial thinning, point cleaning of all secondary stems, and point cleaning of only the secondary stems that were higher than the main stem, within a radius of 0.8 m from the main stem, respectively. A randomised complete block design with three replicates was used. Measurements taken 13 growing seasons after these treatments showed that conventional precommercial thinning was the best treatment for retaining Scots pines as main stems. However, conventional precommercial thinning resulted in the poorest timber quality of the remaining Scots pines in terms of stem taper, thickest branch diameter and relative live crown height. In conclusion, point cleaning has to be followed by a complementary cleaning operation if the objective is to retain Scots pines as main stems in such mixed stands.     

Impact of solar activity on the growth of pine trees: case study

In this work, we focus on the analysis of data for the ring width of two pine tree species: samples of six maritime pines (Pinus pinaster), grown in the north of Portugal, and one sample of Scots pine (Pinus sylvestris), grown in the south of Slovakia. A negative impact of solar activity (SA) was observed on the growth of these pine trees with particularly strong impact in the case of the maritime pines. The width of the annual rings was generally smaller in the years of maximum SA; furthermore, it was found that it is latewood width that is affected whereas earlywood width is not affected; as a corollary, the percentage of latewood also shows a significative negative correlation with SA.     

Using a retrospective dynamic competition index to reconstruct forest succession

Understanding forest dynamics and stand structures is crucial for predicting forest succession. However, many forests have been altered due to century-long land-use practices, which complicates the reconstruction of past and current successional trajectories. For a better understanding of successional processes, we suggest studying the intra- and interspecific competition among single trees across time. We introduce a tree-ring based competition index to reconstruct the competitive dynamics of individual trees over time. This new retrospective dynamic competition index combines a temporal and a spatial component by calculating the yearly ratio between the basal area increments (bai) of the neighbouring trees and the subject tree. The new index is applied to mixed Scots pine (Pinus sylvestris L.) and pubescent oak (Quercus pubescens Willd.) stands in the inner-Alpine dry-valley Valais, for which a change in species composition is hypothesised. The aim is to analyse current stand structures in terms of recent changes in the competitive interactions at the single tree level and to relate these competitive dynamics to land-use change and increasing drought due to climate change. On five plots, the positions of 456 trees were recorded and increment cores were taken to derive bai data. The individual dynamic competition index curves were aggregated in clusters, which define typical patterns of competitive dynamics in both tree species. A large percentage of the trees (87% in oak, 70% in pine) were clustered into a group of trees with constant competition at a relatively low level. However, a smaller group of pines (20%) had recently faced increasing competition. In addition, stand structure analyses indicated a change towards a higher proportion of oak. This change in the competitive ability between oak and pine was found to be related to drought, in that oak had a competitive advantage in dry years. Furthermore, the high proportion of dead branches in pines with decreasing competitive abilities indicated increasing competition for light as a consequence of natural development towards a later successional stage that favours the more shade-tolerant oak. The new retrospective dynamic competition index proved to be promising in studying forest succession. The tree-ring based method allows us to identify changes in the competitive ability of single trees with a high temporal resolution and without repeated assessments.     

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.

     

Can we use shelterwoods in Mediterranean pine forests to promote oak seedling development?

The use of shelterwoods to favour the development of natural or underplanted seedlings is common in temperate forests but rare in the pine forests of the Mediterranean area. Our aim was to assess the use of shelterwoods in Aleppo pine (Pinus halepensis) woodlands in southern France to promote the survival and growth of two co-occurring oak species: the deciduous Quercus pubescens and the evergreen Quercus ilex.Twelve Aleppo pine stands were selected and differentially thinned to create a light shelterwood (basal area = 10 m²/ha, irradiance 52%), a medium shelterwood (basal area = 19 m²/ha, irradiance 33%) and a dense shelterwood (basal area = 32 m²/ha, irradiance 13%). A total of 1248 sowing points, half composed of Q. pubescens and half of Q. ilex, were then set up in these three conditions. Seedling survival and growth were monitored for 3 years. Plant stress was assessed by measuring predawn leaf potential and photosynthetic performance through the F_v/F_m ratio. Soil moisture was also recorded at two depths during two growing seasons.Survival was high for both species in all three conditions due to three consecutive wet years. The lowest survival was recorded for Q. pubescens in the dense shelterwoods. Growth in diameter and height increased from the dense to the light shelterwoods. Shrubs developed more strongly in the light shelterwood, and increasing shrub cover enhanced height growth. Photosynthetic performance was lowest for Q. pubescens in dense shelterwoods and highest in light shelterwoods, whereas the reverse was true for Q. ilex. The lowest predawn potentials were recorded in the dense shelterwoods even though higher soil water content values were measured in this treatment during the summer drought.We show that light shelterwoods were more beneficial to growth than denser ones, indicating control mainly by light availability during the 3 years of the study. However, as lower soil moisture at 30-50 cm depth and faster understorey development were also recorded in this condition, more extended observation is needed to determine whether this benefit persists in subsequent years.     

Evaluating the effect of plant water availability on inner alpine coniferous trees based on sap flow measurements

Climate simulations anticipate an increase in mean summer temperature with synchronous decrease in summer precipitation during the course of the current century in Central Europe. As a consequence, transpiration of forest trees and stands may be altered along with soil water availability. In this study, the effect of reduced plant water availability to conifers was investigated into an open Pinus sylvestris forest (Erico-Pinetum typicum; P. sylvestris 60 %, Picea abies 20 %; and Larix decidua 20 %) within the inner alpine dry valley of the Inn River in Tyrol, Austria. For reducing plant water availability, we installed a transparent roof construction above the forest floor to prevent precipitation to reach the soil. The roofed area covered 240 m² and included 10 trees. A respective number of 11 trees served as controls in the absence of any manipulation. Roofing significantly reduced plant water availability as indicated in lower predawn needle water potentials. Sap flow density (Q _s) was 63, 47, and 24 % lower in roofed P. sylvestris, P. abies, and L. decidua trees, respectively, as compared to control trees. Our findings suggest that P. sylvestris and P. abies behaves “isohydric” as they close their stomata relatively early under conditions of reduced plant water availability and thus stabilize their water relations, whereas L. decidua behaves “anisohydric” and maintains high transpiration rates.     

Evaluation of forest growth simulators with NFI permanent sample plot data from Finland

Tree-level and stand-level forest growth simulators and their combination were evaluated using data from a large network of permanent sample plots of the National Forest Inventory covering the whole of Southern Finland. The simulators were built up with the SIMO framework. The evaluation was carried out both at the stand-level and separately for Scots pine (Pinus sylvestris), Norway spruce (Picea abies), silver birch (Betula pendula) and white birch (Betula pubescens) strata within the plots. Effects of different factors, e.g. age, soil type, stand density and geographical location on the results were also analysed.     

Reconstructing forest history from archaeological data: A case study in the Duero basin assessing the origin of controversial forests and the loss of tree populations of great biogeographical interest

The study of more than 700 charcoals from the Vaccaean settlement of Pintia (Padilla del Duero, Valladolid) provides new local and detailed data about the Iron Age II forest composition in Continental-Mediterranean inland Spain. Pinus gr. sylvestris/nigra (from which an unexpectedly high number of charcoals were identified), together with Mediterranean pines (also highly represented), Juniperus L. and evergreen and deciduous Quercus, were the main taxa living in the area, revealing two different marked biogeographical elements in the late-Holocene landscapes of the Duero basin: Mediterranean and Eurosiberian. On one hand, the results shed new light on the assessment of the origin of Mediterranean pinewoods in the area. On the other hand, the charcoal assemblage of Pintia reveals a more important presence of the current Mediterranean highland pines in the lowlands of the Central Iberian Peninsula in the late-Holocene suggesting the existence of interglacial refugia. Finally, this work highlights palaeoecological studies as valuable tools to support decisions on forest management related to conservation and restoration of our natural heritage.