Hydraulic properties and embolism in small-diameter roots of five temperate broad-leaved tree species with contrasting drought tolerance

? Context

It has been estimated that about half of a plant??s total hydraulic resistance is located belowground, but it is not well known how temperate tree species differ in root hydraulic properties and how these traits vary with the species?? drought tolerance.

? Aims

We examined root anatomical and hydraulic traits in five broad-leaved tree species with different drought tolerance, analyzed the relation between root anatomy and hydraulic conductivity and root embolism, and investigated the relation of these traits to the species?? drought tolerance.

? Methods

In small-diameter roots (2?C6?mm), we measured vessel diameters and vessel density, specific hydraulic conductivity, and the percental loss of conductivity (??native?? embolism) during summer in a mixed forest.

? Results

Specific conductivity was positively related to vessel diameter but not to vessel density. Drought-tolerant Fraxinus showed the smallest mean vessel diameters and drought-sensitive Fagus the largest. Specific conductivity was highly variable among different similar-sized roots of the same species with a few roots apparently functioning as ??high-conductivity roots??.

? Conclusion

The results show that coexisting tree species can differ largely in root hydraulic traits with more drought-sensitive trees apparently having larger mean vessel diameters in their roots than tolerant species. However, this difference was not related to the observed root conductivity losses due to embolism.     

Transpiration of silver Fir (Abies alba mill.) during and after drought in relation to soil properties in a Mediterranean mountain area

Context

Silver fir is declining and dying at its southern margin on the Mediterranean area, where climate is expected to be warmer and drier. At a regional scale, silver fir seems to be vulnerable to drought, and at a forest stand scale, tree death seems to be distributed according to soil water availability.

Aim

To understand the vulnerability of silver fir to drought, factors involved in the regulation of transpiration were assessed with respect to soil properties in order to document the spatial distribution of death rates.

Methods

Soil properties were characterized by electric resistivity measurements. Sap flow density and predawn needle water potential were recorded on sampled trees during several years, and crown specific transpiration was estimated. In addition, the vulnerability of coarse roots and branches to cavitation was quantified.

Results and conclusion

Trees growing on soils with a large water storage capacity were the most vulnerable to drought induced soil water deficits. Transpiration was down-regulated as soon as predawn water potential decreased. The vulnerability to cavitation was low, which protected the trees from run-away xylem embolism. Severe soil water deficits led to a rapid decrease of transpiration, which was still visible the following year. The drop-off in transpiration was mainly due to inner sap flow that almost ceased after the drought on all monitored trees. Our results suggest that root dynamic and the ability of roots to take up water were modified by soil water deficit over several years. Such a regulation scheme needs to be better documented and included in models to address silver fir forest responses to drought.     

Summer drought and canopy opening increase the strength of the oak seedlings–shrub spatial association

? Context

The knowledge of how shrub–seedling interactions vary with summer drought, canopy opening, and tree species is crucial for adapting forest management to climate change.

? Aims

The aim of this study was to assess variation in shrub–oak recruitment associations along a south–north drought climate gradient and between two levels of canopy cover in coastal dune forest communities in a climate change-adapted forest management perspective.

? Material and methods

Mapped data of associational patterns of seedlings of three oak species with interspecific pooled shrubs were analyzed using a bivariate pair correlation function in 10 (0.315 ha) regeneration plots located in forest and recent gap sites along the climate gradient. An index of association strength was calculated in each plot and plotted against a summer moisture index.

? Results

The association strength increased with increasing summer drought from wet south to dry north and from closed forests to gaps.

? Conclusion

Consistent with facilitation theory, our results suggest that climate change may shift associational patterns in coastal dune forest communities towards more positive associations, in particular in canopy gaps. In a perspective of climate change, foresters may need to conserve understory shrubs in gaps in order to promote oak species regeneration.     

An inconvenient truth about xylem resistance to embolism in the model species for refilling <Emphasis Type="Italic">Laurus nobilis</Emphasis> L.

Key message

Direct, non-invasive X-ray microtomography and optical technique observations applied in stems and leaves of intact seedlings revealed that laurel is highly resistant to drought-induced xylem embolism. Contrary to what has been brought forward, daily cycles of embolism formation and refilling are unlikely to occur in this species and to explain how it copes with drought.

Context

There has been considerable controversy regarding xylem embolism resistance for long-vesselled angiosperm species and particularly for the model species for refilling (Laurus nobilis L.).

Aims

The purpose of this study was to resolve the hydraulic properties of this species by documenting vulnerability curves of different organs in intact plants.

Methods

Here, we applied a direct, non-invasive method to visualize xylem embolism in stems and leaves of intact laurel seedlings up to 2-m tall using X-ray microtomography (microCT) observations and the optical vulnerability technique. These approaches were coupled with complementary centrifugation measurements performed on 1-m long branches sampled from adult trees and compared with additional microCT analyses carried out on 80-cm cut branches.

Results

Direct observations of embolism spread during desiccation of intact laurels revealed that 50% loss of xylem conductivity (Ψ₅₀) was reached at ??7.9?±?0.5 and ??8.4?±?0.3 MPa in stems and leaves, respectively, while the minimum xylem water potentials measured in the field were ??4.2 MPa during a moderate drought season. Those findings reveal that embolism formation is not routine in Laurus nobilis contrary to what has been previously reported. These Ψ₅₀ values were close to those based on the flow-centrifuge technique (??9.2?±?0.2 MPa), but at odds with microCT observations of cut branches (??4.0?±?0.5 MPa).

Conclusion

In summary, independent methods converge toward the same conclusion that laurel is highly resistant to xylem embolism regardless its development stage. Under typical growth conditions without extreme drought events, this species maintains positive hydraulic safety margin, while daily cycles of embolism formation and refilling are unlikely to occur in this species.

     

Soil compaction due to heavy forest traffic: measurements and simulations using an analytical soil compaction model

? Context

Models for predictions of soil compaction following forest traffic represent important decision tools for forest managers in order to choose the best management practices for preserving soil physical quality. In agricultural soil compaction research, analytical models are widely used for this purpose.

? Aims

Our objective was to assess the ability of an analytical model to predict forest soil compaction under forwarder traffic.

? Methods

We used the results from two experimental sites set up in north-eastern France in 2007 and 2008 to compare simulations using the SoilFlex model with observed bulk density following forwarder traffic.

? Results

The best model-based predictions were found when considering the mean initial soil conditions and an increased rebound parameter in the upper soil layers (0–10 cm) in comparison to the deeper layers (10–50 cm). The need to increase the rebound parameter in the soil surface layer to improve model accuracy was attributed to a large soil organic matter content in the uppermost layers of forest soils. For the site where initial soil mechanical parameters were measured as a function of soil bulk density and water content, the model performance was good, with a root mean square error (RMSE) of 0.06. The model performed poorer (RMSE of 0.11), especially for the surface soil layer, for the second site that was wetter at the time of traffic and where soil mechanical properties were not measured but estimated by means of pedo-transfer functions.

? Conclusions

SoilFlex was found to yield satisfactory predictions and could help forest managers estimate the risk of compaction and to select the most appropriate machinery for given soil conditions in order to preserve the soil from physical degradation during traffic in forest ecosystems. However, our results emphasise the need for research on soil mechanical properties of forest soils, in particular on the role of soil organic matter and roots on soil compressive properties.     

Coastal and interior Douglas-fir provenances differ in growth performance and response to drought episodes at adult age

Context

Since the 2003 drought and heat wave, Douglas-fir dieback has been reported in France in trees older than 30 years. Consequently, it is questioned whether selected Douglas-fir provenances are suited to the frequent and severe drought events which are forecast due to climate change.

Aims

Our objective was to contribute to the screening of variability in productivity and growth response to soil-water deficit of mature trees from provenances not currently used for plantation in France.

Methods

We sampled 22 provenances, including coastal and interior Douglas-fir, covering a wide part of its natural distribution, from Oregon to California for coastal provenances and from British Columbia to New Mexico for interior provenances. These provenances were planted at the mid 1970s in two provenance trials located in the south-west area of France. Variability of productivity, of wood density, and of radial growth in response to drought episodes among provenances was quantified and related to soil-water deficit computed by daily water balance calculations.

Results

Whatever the provenance, annual radial growth is highly dependent on local soil-water deficit (Felines R ² = 0.57, Le Treps R² = 0.49). Radial growth and wood properties exhibit large differences between provenances at 30 years old. Variability between provenances for all wood characteristics studied is mainly structured geographically. Coastal provenances perform best for productivity at 30 years old (619 cm²?±?59), and exhibit a small growth reduction in 2004, the second successive year of drought (?10.7 %?±?3.8). Surprisingly, the southern interior provenances from the driest environments in the natural range show a large growth reduction in 2004 (?30.5 %?±?5.2).

Conclusions

The provenances tested exhibited significant differences in growth performance and growth reduction induced by the soil-water deficit. The approach coupling retrospective analysis of radial growth on mature trees and water balance modelling is relevant for evaluating provenance adaptation to more frequent or severe drought episodes.     

Low soil temperatures increase carbon reserves in Picea mariana and Pinus contorta

? Context

Soil temperature can limit tree growth and function, but it is often unaddressed in understanding the successional status of trees.

? Aims

We tested how soil temperature affected carbon allocation strategies of two dominant co-occurring boreal conifer species, Pinus contorta and Picea mariana.

? Methods

We measured nonstructural carbon (NSC) concentrations, biomass, and photosynthesis of dormant and actively growing 2-year-old seedlings in response to three soil temperatures (5, 10, and 20 °C) under a common ambient air temperature.

? Results

For both species, variation in carbon reserves with soil temperature was more pronounced following seedling growth than during dormancy. For both species and all organ types (roots, needles, and stems), NSC concentrations were highest when seedlings were grown at 5 than 20 °C. Mass adjusted for NSC content was negatively correlated with NSC concentration for all organ types of both species. Soil temperature had a marginally significant effect on photosynthesis of pine; seedlings grown at 10 or 20 °C acquired more carbon than seedlings grown at 5 °C. Spruce seedlings photosynthesized more when grown at 20 °C than at 5 or 10 °C.

? Conclusion

Interspecific differences in allocation of carbon may underlie the responses of P. mariana and P. contorta to cold soils and consequently their successional status.     

The incorporation of extreme drought events improves models for beech persistence at its distribution limit

Context

Projections of species distribution models under future climate are usually based on long-term averages. However, singular extreme drought events presumably contribute to the shaping of distribution limits at the retreating low-elevation xeric limits.

Methods

The objectives of this study were to set up a distribution model based on extreme drought events (EDM), which uses sanitary logging information as a proxy of vitality response of beech, and compare it with the results of classical species distribution models (SDMs).

Results

Predictions of the EDM for 2025 were in agreement with those of the SDM, but EDM predicted a more serious decline in all regions of Hungary towards the end of the century.

Conclusion

These results suggest that the predicted increase in frequency and severity of drought events may further limit the distribution of beech in the future.     

Xylogenesis of Pinus pinaster under a Mediterranean climate

Context

The knowledge on cambial activity in water-limited environments, such as the Mediterranean, is still fragmentary. Dendrochronological studies have determined that spring precipitation plays an important part in determining tree-ring width and the properties of tracheids. However, the complex relation between cambial phenology and climate is still far from understood.

Aims

We studied the influence of climate, especially water stress, on maritime pine wood formation with the aim of determining the influence of drought on cambial activity.

Methods

A plantation of maritime pine (Pinus pinaster) was selected in the west coast of Portugal, to monitor cambial activity and wood formation using anatomical observations and band dendrometers. The trees were monitored weekly over 2 years (2010 and 2011).

Results

Xylem differentiation started earlier when warmer late winter temperatures were observed. Water stress triggered an earlier stop of wood formation and also the formation of tracheids with smaller lumen area. In both years a bimodal pattern of stem radial increment was registered by band dendrometers with two periods of increment: one in spring and another in autumn. The xylem anatomy study suggests that the autumnal increment period corresponded mostly to stem rehydration, since the differentiation of new xylem cells by the cambium was not observed.

Conclusion

Maritime pine cambial activity appears to be under a double climatic control: temperature influences cambial onset and water availability growth cessation.     

Growth response to climate and drought change along an aridity gradient in the southernmost Pinus nigra relict forests

Context

Tree populations at the rear edge of species distribution are sensitive to climate stress and drought. However, growth responses of these tree populations to those stressors may vary along climatic gradients.

Aims

To analyze growth responses to climate and drought using dendrochronology in rear-edge Pinus nigra populations located along an aridity gradient.

Methods

Tree-ring width chronologies were built for the twentieth century and related to monthly climatic variables, a drought index (Standardized Precipitation–Evapotranspiration Index), and two atmospheric circulation patterns (North Atlantic and Western Mediterranean Oscillations).

Results

Growth was enhanced by wet and cold previous autumns and warm late winters before tree-ring formation. The influence of the previous year conditions on growth increased during the past century. Growth was significantly related to North Atlantic and Western Mediterranean Oscillations in two out of five sites. The strongest responses of growth to the drought index were observed in the most xeric sites.

Conclusion

Dry conditions before tree-ring formation constrain growth in rear-edge P. nigra populations. The comparisons of climate-growth responses along aridity gradients allow characterizing the sensitivity of relict stands to climate warming.     

Effects of region, soil, land use, and terrain type on fuelwood properties of five tree/shrub species in the Sahelian and Sudanian ecozones of Mali

? Context

There is little information about inter- and intra-specific variation in fuelwood properties of trees/shrubs in West Africa.

? Aims

We studied variation in fuelwood properties of Balanites aegyptiaca, Combretum glutinosum, Guiera senegalensis, Piliostigma reticulatum, and Zizyphus mauritiana in the Sahelian and Sudanian ecozones of Mali.

? Methods

Trees were sampled on different soil (sandy, sandy/loam, or rocky), land use (parkland agroforest or woodland), and terrain (flat, temporarily flooded, or hill slope) types in five regions extending from the drier eastern to the more humid western parts of Mali. Basic density, volatile matter, fixed carbon, ash content, moisture content, gross calorific value, and fuel value index were measured for each tree and adjusted for tree age.

? Results

All fuelwood properties differed significantly among species, but there were significant species by region interactions. Most fuelwood properties of four species differed significantly among regions. Soil and/or land use type had significant effects on a few fuelwood properties of four species.

? Conclusion

In general, fuelwood production is recommended in all regions for G. senegalensis, in drier regions for B. aegyptiaca and C. glutinosum, in more humid regions for Z. mauritiana, and on rocky soils for all species. Fuelwood production of P. reticulatum is not recommended.     

Walnut (Juglans spp.) ecophysiology in response to environmental stresses and potential acclimation to climate change

? Context

Walnuts (Juglans spp.) are ecologically and commercially important trees, yet synthesis of past and current research findings on walnut ecophysiology is lacking, especially in terms of potential acclimation to climate change.

? Aims

This study aims to (1) investigate walnut ecophysiology by comparing its attributes to associated deciduous angiosperms, (2) address potential acclimation of walnut to climate change, and (3) identify areas for prioritization in future research.

? Results

There is considerable uncertainty regarding the magnitude of potential effects of climate change on walnut. Some studies tend to indicate walnut could be negatively impacted by climate change, while others do not. Walnut may be at a disadvantage due to its susceptibility to drought and frost injury in current growing regions given the projected increases in temperature and extreme climatic events. Other regions that are currently considered cold for walnut growth may see increased establishment and growth depending upon the rate of temperature increase and the frequency and severity of extreme climatic events.

? Conclusion

Research investigating a combination of environmental factors, such as temperature, carbon dioxide, ozone, water, and nitrogen is needed to (1) better project climate change effects on walnut and (2) develop management strategies for walnut acclimation and adaptation to climate change.     

Within-population variability in architectural traits and suitability to forestry conditions in Nothofagus nervosa (= N. alpina; Nothofagaceae)

? Context

There is evidence that Nothofagus nervosa (= N. alpina) is suitable for timber production in temperate regions due to its wood quality and fast growth. However, high intra-specific variability in the architectural traits of this species limits its usefulness.

? Aims

This study was aimed at evaluating intra-specific variability in the architecture of N. nervosa trees at the population level, with emphasis on traits related to the suitability of trees for timber production.

? Methods

The size of 13-year-old trees installed in an experimental field population and the extent of differentiation between trunk and branches were compared among families (trees derived from different mother trees).

? Results

The large majority of trees exhibited architectural features indicative of high suitability for timber production: clear differentiation between trunk and main branches and few occurrences of permanent trunk forks, apex deaths, and sylleptic branching. Trees belonging to different families differed in height and trunk diameter. Micro-environmental conditions had a very significant effect on most architectural traits, including the trunk differentiation.

? Conclusions

N. nervosa emerges as a suitable option for timber production in temperate regions. Trunk growth seems to be under both environmental and genetic control in this species.     

Carbon storage in biomass,litter, and soil of different plantations in a semiarid temperate region of northwest China

? Context

A large area of abandoned land in the semiarid temperate region of China has been converted into plantations over the past decades. However, little information is available about the ecosystem C storage in different plantations.

? Aim and methods

Our objective was to estimate the C storage in biomass, litter, and soil of four different plantations (monospecific stands of Larix gmelinii, Pinus tabuliformis, Picea crassifolia, and Populus simonii). Tree component biomass was estimated using allometric equations. The biomasses of understory vegetation and litter were determined by harvesting all the components. C fractions of plant, litter, and soil were measured.

? Results

The ecosystem C storage were as follows: Picea crassifolia (469 t C/ha)?>?Larix gmelinii (375 t C/ha), Populus simonii (330 t C/ha)?>?Pinus tabuliformis (281 t C/ha) (P?<?0.05), 59.5–91.1 % of which was in the soil. The highest tree and understory C storage were found in the plantation of Pinus tabuliformis (247 t/ha) and Larix gmelinii (1.2 t/ha) respectively. The difference in tree C fraction was significant among tree components (P?<?0.05), following the order: leaf?>?branch?>?trunk?>?root. The highest soil C (SC) was stored in Picea crassifolia plantation (411 t C/ha), while Populus simonii plantation had a higher SC sequestration rate than others.

? Conclusion

C storage and distribution varied among different plantation ecosystems. Coniferous forests had a higher live biomass and litter C storage. Broadleaf forests had considerable SC sequestration potential after 40 years establishment.     

Assessing changes in species distribution from sequential large-scale forest inventories

? Context

It is assumed that global change is already affecting the composition, structure and distribution of forest ecosystems; however, detailed evidences of altitudinal and latitudinal shifts are still scarce.

? Aims

To develop a method based on National Forest Inventory (NFI) to assess spatio-temporal changes in species distributions.

? Methods

We develop an approach based on universal kriging to compare species distribution models from the different NFI cycles and regardless of the differences in the sampling schemes used. Furthermore, a confidence interval approach is used to assess significant changes in species distribution. The approach is applied to some of the southernmost populations of Pinus sylvestris and Fagus sylvatica in the Western Pyrenees over the last 40 years.

? Results

An increase of the presence of the two species in the region was observed. Scots pine distribution has shifted about 1.5 km northwards over recent decades, whereas the European beech has extended its distribution southwards by about 2 km. Furthermore, the optimum altitude for both species has risen by about 200 m. As a result, the zone in which the two species coexist has been enlarged.

? Conclusions

This approach provides a useful tool to compare NFI data from different sampling schemes, quantifying and testing significant shifts in tree species distribution over recent decades across geographical gradients.     

Is the private forest sector adapting to climate change? A study of forest managers in north Wales

? Context

Two-thirds of Britain’s forest area is privately owned. Thus, understanding private forest owners and managers, and their attitudes to uncertainty and change, is essential for the success of climate change adaptation policies.

? Aim

The aims of this study are to (1) assess how beliefs in climate change in the private sector have influenced forest management practices; (2) identify constraints related to changes in species choice and silvicultural systems; (3) analyse the implications for implementing climate change policy in forestry.

? Method

Semi-structured interviews with key informants who provide advice to, or manage woodlands in, the private forest sector in north Wales.

? Results

Woodland managers and some advisers are not generally convinced of a need to adapt. They feel the future is uncertain, more usually in relation to tree disease than to climate change itself. Species choice is the principle focus of adaptation activities and reveals a deep divide in opinion. Commercial advisors look to new exotics but are inhibited by absence of markets, while small-scale owners rely more on native genetic diversity.

? Conclusions

Findings that are likely to apply widely include: the influential role of forest agents in forest management decisions including species choice; lack of confidence in climate change predictions, and in markets; more immediate concerns about tree pests and diseases; demand for leadership from the public sector, and for engagement amongst the private sector. Further research is needed across a wider area to test the variability in relationship between attitudes and behaviours, and local conditions including climate change predictions.     

Effect of selective logging on stand structure and tree species diversity in a subtropical evergreen broad-leaved forest

?Context

Selective logging followed by natural regeneration is rarely employed for restocking subtropical evergreen broad-leaved forests in East Asia compared with the use of clear-cutting.

?Aims

To clarify the succession of these forests, the effects of selective logging on stand structure, species diversity, and community similarity were studied in a mature and regenerating forest in Okinawa, Japan.

?Methods

Four study plots were established, and trees ≥1.2 m height were identified by species name, tree height, and diameter at breast height.

?Results

The results showed that the species composition of regenerating forest was similar to mature forest; however, the former had a greater species density and Shannon–Wiener index than the latter. Castanopsis sieboldii and Distylium racemosum, the predominant trees in the mature forest, continued to dominate the regenerating forest, with a broad layer distribution. High Sørensen and Jaccard community similarity indices for mature and regenerating forest indicated that the regeneration occurred in a progressive succession.

?Conclusion

The similar species composition and stand structure for both mature and regenerating forest, and the higher species diversity for the latter, provided no evidence of forest degeneration and suggested that the regenerating forest may develop into a stand similar to preselective logging forest.     

Disturbances in European beech water relation during an extreme drought

Context

In the context of a probable increase in intensity and frequency of extreme summer drought events, a better understanding of the key processes involved in water relations is needed to improve the theoretical foundations of predictive process-based models.

Aims

This paper aims to analyse how temperate deciduous trees cope with water shortage.

Methods

The exceptional summer drought of 2003 in Europe provided an opportunity to monitor stomatal conductance and twig water potential in European beech (Fagus sylvatica L.) at predawn and midday and to analyse variations with respect to leaf height within the canopy. By comparing our field measurements of twig water potential to values found in the literature, we confirmed the strong impact of soil water shortage on crown water relations.

Results

This paper shows that (1) the vertical gradient of stomatal conductance within the crown disappeared under extreme soil water depletion; (2) at maximum drought intensity, predawn twig water potential (ψ _pd) reached ?2.3 MPa at a height of 14 m in the crown and ?2.0 MPa at a height of 10 m. The significant differences in ψ _pd between the two measurement heights in the canopy may be due to night transpiration; (3) there was a close relationship between predawn twig water potential and relative extractable soil water; (4) as drought conditions intensified, there was a close relationship between canopy radiation interception and predawn water potential, as estimated daily from relative extractable soil water.     

Low light acclimation in five temperate broad-leaved tree species of different successional status: the significance of a shade canopy

?Context

Tree species differ largely in their capability to produce characteristic shade leaves with effective morphological and physiological acclimation to low light.

?Aims

By examining the sun/shade leaf differentiation in leaf morphology, foliar nitrogen and photosynthetic capacity in five temperate tree species of different successional status, we aimed at identifying those leaf traits that determine the development of a typical shade crown with low light-acclimated leaves.

?Methods

Leaf morphology, foliar N content, photosynthetic capacity (V _cmax, J _max and A _max) and leaf dark respiration (R _d) were measured in the canopies of 26 adult trees of Fraxinus, Acer, Carpinus, Tilia and Fagus species.

?Results

Six traits (the sun/shade leaf differentiation in specific leaf area, leaf size, A _max per leaf area or per mass, photosynthetic N use efficiency and R _d) were found to characterise best the degree of low light acclimation in shade leaves. All five species exhibited certain modifications in leaf morphology and/or physiology in response to low light; Fagus sylvatica showed the highest and Fraxinus excelsior the lowest shade leaf acclimation.

?Conclusions

Our results indicate that the five early/mid- to late-successional species have developed species-specific low light acclimation strategies in their shade crowns which differ in terms of the relative importance of leaf morphological and physiological acclimation.