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 .

     

Photosynthetic temperature adaptation of Pinus cembra within the timberline ecotone of the Central Austrian Alps

? Temperature is suggested to determine the upper limit of tree life. Therefore, future climate warming may be of importance for tree distribution within the European Alps, where low temperatures limit carbon metabolism.

? We focused on the effects of air and soil temperature on net photosynthesis (P _n) of Pinus cembra an evergreen climax species of the timberline ecotone of the Central Austrian Alps. Light response and temperature response curves were estimated along an altitudinal gradient ranging from the forest limit up to the krummholz limit in both summer and fall.

? In general, P _n was significantly lower in fall as compared to summer. Nevertheless, independent from season mean P _n values tended to increase with elevation and were positively correlated with root zone temperatures. The specific leaf area by contrast declined with increasing elevation. Furthermore, the temperature optimum of net photosynthesis declined with increasing elevation and was positively correlated with the mean maximum air temperature of the 10 days prior the date of measurement.

? Thus, our findings appear to reflect a long-term adaptation of the photosynthetic apparatus of Pinus cembra to the general temperature conditions with respect to elevation combined with a short term acclimation to the prevailing temperature regime.

     

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.

     

Tree-ring Δ13C reveals the impact of past forest management on water-use efficiency in a Mediterranean oak coppice in Tuscany (Italy)

? The effects of thinning and heavy stand density reduction was investigated in Turkey oak (Quercus cerris L.) forests of central Italy, to evaluate the physiological responses and the growth status of trees that survived a past coppice cut and thinning to convert the stand to high-forest.

? The working hypothesis was that a strong decrease in stand density would cause a decreasing in canopy-intrinsic water-use efficiency (measured as the ratio of CO₂ assimilation to stomatal conductance, A/g), thus an increase in tree-ring carbon isotopic discrimination (Δ¹³C).

? The tree-ring Δ¹³C of the remaining trees (“survivors”) was found to have significantly (P < 0.05) raised between year two and year seven since the coppice stand was thinned (high-forest conversion thinnings). This effect was mostly caused by a large decrease in tree-rings Δ¹³C at control site which was characterized by high density and competition by trees. An increase in survivors tree-rings Δ¹³C probably indicates an improved water availability, possibly induced by a decrease in competition and in stand density or a decrease in the precipitation intercepted by the canopy (i.e., a stronger increase in g over A since a decrease in A is highly unlikely). A change in foliar nitrogen, foliar Δ¹³C and content in chlorophylls was also recorded seven years after thinning.

? Thinnings carried out to convert old abandoned coppices into high-forest stands induce short-term stimulation of Turkey oak growth by increasing light and water availability. We were able to make a detailed reconstruction of the impact of past silvicultural treatment on the stand using a tree-ring wood Δ¹³C time-series.

     

Physiological variation among western redcedar (Thuja plicata Donn ex D. Don) populations in response to short-term drought

? Variation in the ability of western redcedar (Thuja plicata Donn ex D. Don) populations to withstand water stress may exist because this species is found in coastal and interior biogeoclimatic subzones representing the full range of precipitation regimes in British Columbia, Canada.

? Seven western redcedar populations from locations in British Columbia, representing a wide range of habitat types, were assessed for their gas exchange and water relations response to controlled drought.

? Before drought, population variation occurred in stomatal conductance, net CO₂ assimilation rate and intrinsic water use efficiency and the relative water content at turgor loss point. During drought, populations had different responses of net CO₂ assimilation to decreasing predawn shoot water potential. After drought, populations differed in stomatal conductance and intrinsic water use efficiency, plus osmotic potential at turgor loss point, osmotic potential at saturation and apparent cuticular transpiration. Western redcedar populations from drier-inland habitats had a lower osmotic potential at turgor loss point, lower relative water content at turgor loss point and lower apparent cuticular transpiration in response to drought than populations from coastal origin with temperate maritime habitat.

? Reduction of cuticular water loss and adjustments of cellular water relations in response to drought was found to occur among seven western redcedar populations originating along a precipitation gradient while; there were minimal population differences in the gas exchange response to drought.

     

Extending sapwood — Leaf area relationships from stems to roots in Coast Douglas-fir

? Studies of allometric relationships between leaf area and the cross-sectional area (CSA) of sapwood in the stem have shed light on the structural and functional relationships between water-conducting and photosynthetic tissues.

? The purpose of this study was to test whether sapwood-leaf area relationships could be extended from stems to roots in coast Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco). Twelve trees were felled, their stumps were excavated, and the CSA of sapwood and heartwood were estimated for individual roots, entire root systems, and stem section.

? Root sapwood CSA was greater than sapwood CSA throughout the stem, and the ratio of leaf area to sapwood CSA (A _l :A _s ) was accordingly lower for root sapwood. The relationship between sapwood CSA and leaf area was more variable in roots and at groundline compared to crown base. Root A _l :A _s decreased with relative tree height (tree height/mean stand height).

? The strong allometric relationship between leaf area and the CSA of sapwood in the stem generally holds when extended to roots. The greater CSA of sapwood in roots versus stems may reflect differences in their roles in supporting the tree.

     

Drought susceptibility and recovery of transplanted Quercus rubra seedlings in relation to root system morphology

? Transplant shock, implicated by depressed seedling physiological status associated with moisture stress immediately following planting, limits early plantation establishment. Large root volume (Rv) has potential to alleviate transplant shock because of higher root growth potential and greater access to soil water.

? We investigated impacts of drought and transplant Rv on photosynthetic assimilation (A), transpiration (E), stomatal conductance (g _s ), predawn leaf xylem water potential (Ψ_L), and growth of northern red oak (Quercus rubra L.) seedlings to explain mechanisms associated with susceptibility to transplant shock. One year-old barerooot seedlings were graded into four Rv categories and either well watered or subjected to drought consisting of low, medium, or high moisture stress by discontinuing irrigation at 22-day intervals for 3 months. Thereafter, all treatments were re-watered to examine recovery.

? Transplant shock was signified by reduced A, E, g _s, and Ψ_L, which generally increased with increasing moisture stress and Rv. Physiological status improved during recovery, though stress was still evident in seedlings exposed to medium or high moisture stress and in larger Rv seedlings. Growth declined with increasing moisture stress but was generally similar among Rv treatments, likely reflecting greater A at the whole plant level and/or reliance upon stored reserves in large Rv seedlings.

? The most effective drought avoidance mechanisms were root growth, stomatal regulation, reduced leaf area, and higher growth allocation to roots relative to shoots. Our results suggest that large initial Rv does not enhance drought avoidance during the first season after transplant in northern red oak seedlings.

     

Effects of atmospheric and climate change at the timberline of the Central European Alps

? This review considers potential effects of atmospheric change and climate warming within the timberline ecotone of the Central European Alps. After focusing on the impacts of ozone (O₃) and rising atmospheric CO₂ concentration, effects of climate warming on the carbon and water balance of timberline trees and forests will be outlined towards conclusions about changes in tree growth and treeline dynamics.

? Presently, ambient ground-level O₃ concentrations do not exert crucial stress on adult conifers at the timberline of the Central European Alps. In response to elevated atmospheric CO₂ Larix decidua showed growth increase, whereas no such response was found in Pinus uncinata. Overall climate warming appears as the factor responsible for the observed growth stimulation of timberline trees.

? Increased seedling re-establishment in the Central European Alps however, resulted from invasion into potential habitats rather than upward migration due to climate change, although seedlings will only reach tree size upon successful coupling with the atmosphere and thus loosing the beneficial microclimate of low stature vegetation.

? In conclusion, future climate extremes are more likely than the gradual temperature increase to control treeline dynamics in the Central European Alps.

     

Current uptake of 15N-labeled ammonium and nitrate in flooded and non-flooded black spruce and tamarack seedlings

? We investigated the effects of flooding for three weeks on physiological responses and uptake of NH ₄ ⁺ and NO ₃ ^? by black spruce (Picea mariana (Mill.) BSP.) and tamarack (Larix laricina (Du Roi) K. Koch) seedlings fertilized with labeled (¹⁵NH₄)₂SO₄ or K¹⁵NO₃ in a growth chamber experiment.

? Flooding reduced photosynthesis (A), transpiration (E), water use efficiency (WUE), and current uptake of NH ₄ ⁺ and NO ₃ ^? in both species.

? Under flooding, there were no significant differences between the two species in uptake of either NH ₄ ⁺ or NO ₃ ^? at the whole-plant level but black spruce had higher translocation of NH ₄ ⁺ to the shoots than did tamarack.

? Under non-flooded conditions, black spruce seedlings exhibited higher uptake of both NH ₄ ⁺ and NO ₃ ^? than did tamarack and demonstrated preferential uptake of NH ₄ ⁺ (19. 7 mg g^?1dw) over NO ₃ ^? (12.3 mg g^?1dw after three weeks). In contrast, non-flooded tamarack seedlings had equal uptake of NH ₄ ⁺ (4.96 mg g^?1dw) and NO ₃ ^? (4.97 mg g^?1dw).

? We hypothesize that the ability of tamarack to equally exploit both ¹⁵NH ₄ ⁺ and ¹⁵NO ₃ ^? would confer an advantage over black spruce, when faced with limitations in the availability of different forms of soil nitrogen.

     

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.

     

Do artificial and natural defoliation have similar effects on physiology of Eucalyptus globulus Labill. seedlings?

? Artificial defoliation is often used to simulate defoliation by herbivory and is usually considered a good indication of a plant’s response to a given type of damage. However, the findings of studies directly comparing the two defoliation types are inconsistent.

? Here, the short term effects of artificial and insect defoliation by larvae of Paropsisterna agricola on growth, biomass allocation and photosynthetic capacity of Eucalyptus globulus seedlings were compared in a glasshouse experiment. The artificial defoliation was carried out to closely resemble the spatial patterns observed for insect defoliation.

? Height and diameter increments were reduced as a result of insect defoliation, whereas artificial defoliation had no significant effect on height. Increased photosynthetic capacity was observed in response to both treatments, but the magnitude of this increase was larger in insect-than in artificially-defoliated seedlings. Significant reductions in foliar carbohydrate content and total biomass were noticeable in artificially-defoliated seedlings. Although the foliar carbohydrate levels also decreased across the crown zones following insect defoliation treatment, seedlings allocated a large amount of their biomass in the branches of the damaged zone.

? Despite our best endeavours to simulate insect defoliation in the artificial treatment, the latter may not reflect accurately the full strength of the effects. However, artificial and insect defoliation were similar in their direction of the responses they caused in E. globulus seedlings.

     

Genetic structure of Tunisian natural carob tree (Ceratonia siliqua L.) populations inferred from RAPD markers

? Seven RAPD markers were used to assess the genetic diversity and structure of ten Tunisian natural Ceratonia siliqua L. populations from different geographic and bioclimatic zones.

? The species maintain a high diversity within population as estimated by the percentage of polymorphic loci and Shannon’s index (P% = 76.31, $\bar H'_{pop} = 0.569$ ). The range of variation between populations was large. Populations from the upper semi-arid bioclimates, with more continuous distribution area showed the highest level of variation.

? A high genetic differentiation among populations (Φ_ST = 0.250 and $\bar G_{ST} = 0.347$ ), as a result of population isolation was revealed. Nevertheless, the genetic structure is in accordance with bioclimate indicating that ecological factors also should influence differentiation. Populations from the sub-humid, upper semi-arid and mean semi-arid zones clustered together and were distinct from those of the lower semi-arid ones.

? Conservation strategy should be made according to the level of polymorphism within population and bioclimate.

     

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.

     

Sap flow measurement by a single thermal dissipation probe: exploring the transient regime

? In thermal dissipation method the stationary regime is used to estimate sap flow density from the temperature difference between a heating probe and a non-heating one.

? A new approach, based on the non-stationary regime of a single heating probe, is investigated on an olive tree branch. Probe temperature is monitored for different flow densities and for heating and cooling modes. Measurement sequences of 10 min using 5 s time steps are made just after the heating is switched on or off.

? Results show that temperature variation has a quasi-exponential phase whose duration decreases for increasing flow densities. Thermal indices using temperature of the probe measured at initial, final, and intermediate times t _i were developed. Relationships between sap flow density and thermal indices were found to be linear for t_i = 10 s?60 s with better coefficients of determination in cooling mode (R ² = 0.94–0.96) than in heating (R ² = 0.83–0.93). Validation tests confirmed the robustness of the single probe method with lowest error obtained for t _i = 20 s.

? Single probe method seems to be a promising technique to measure tree sap flow. Its application could be considered under cooling mode using t _i = 20 s and relatively short measurement sequences e.g. 5–10 min.

     

Inoculation of seven pine species or hybrid seedlings with Korean isolates of pinewood nematode under greenhouse conditions

? Seedlings of seven pine species or hybrids, Pinus densiflora, Pinus thunbergii, Pinus virginiana, Pinus echinata, Pinus koraiensis, Pinus thunbergii × Pinus massoniana and Pinus × rigitaeda, were inoculated in a greenhouse with 100, 1 000 and 10 000 pathogenic nematodes, Bursaphelenchus xylophilus, to compare initial symptom development and to evaluate susceptibility to pinewood nematodes.

? Genetic types were significantly different in the starting day of needle discoloration (χ² = 43.48; P < 0.001) and the period from nematode inoculation to seedling death (χ² = 32.85; P > 0.001), but treatment groups were not statistically different in the days from inoculation to seedling death in any of the pine species.

? There was no relationship between seedling mortality and the number of nematodes recovered from infected seedlings. Only eight of the nematode-inoculated seedlings remained alive by day 83 after inoculation, including four seedlings each of P. echinata and P. × rigitaeda grown from bulked seed collections.

? Seedlings from the bulk lots were likely more genetically diverse than those from the single-source lots. A broader genetic representation of pine species or hybrids is suggested for screening for planting stocks resistant to B. xylophilus.

     

Effects of organic residue management and legume cover on growth of pine seedlings, nutrient leaching and soil properties

? The short-term effect of organic residue management on the growth and nutrition of Pinus pinaster Ait. seedlings, and on nutrient leaching and chemical properties of an acid soil was assessed through a lysimeter experiment. Treatments included absence, placement on the soil surface, and incorporation into the soil (with and without legume cover cropping) of organic residues (forest floor litter or forest floor litter plus harvest residues).

? Residues placed on the soil surface enhanced seedling growth. Organic residues reduced nutrient losses (NO ₃ ^? , Ca and Mg) and resulted in nutrient accumulation in the soil. Harvest residues positively affected K seedling nutrition status and enhanced K soil accumulation.

? Legume cover cropping reduced soil nutrient losses (N, Ca, Mg and K) during the early stage of seedling growth; it also improved seedling nutrition status (N and P), but without any effect on growth.

? Harvest residues plus forest floor litter placed on the soil surface was the most appropriate management to both reduce nutrient losses through leaching and increase height of seedlings at the end of the experimental period (two years).

     

Mesophyll conductance to CO2 and leaf morphological characteristics under drought stress during Quercus ilex L. resprouting

? Quercus ilex L., the dominant species in Mediterranean forests and one with a great capacity for resprouting after disturbances, is threatened by the expected increase in fire frequency and drought associated with climate change.

? The aim of this study was to determine the contribution of photosynthesis limitants, especially mesophyll conductance (g _mes) during this species’ resprouting and under summer drought.

? Resprouts showed 5.3-fold increased g _mes and 3.8-fold increased stomatal conductance (g _s) at mid-day with respect to leaves of undisturbed individuals. With increased drought, structural changes (decreased density and increased thickness) in resprouts contributed to the observed higher photosynthesis and increased g _mes. However, g _mes only partially depended on leaf structure, and was also under physiological control. Resprouts also showed lower non-stomatal limitations (around 50% higher carboxylation velocity (V _c,max) and capacity for ribulose-1,5-bisphosphate regeneration (J _max)). A significant contribution of g _mes to leaf carbon isotope discrimination values was observed.

? g _mes exhibits a dominant role in photosynthesis limitation in Q. ilex and is regulated by factors other than morphology. During resprouting after disturbances, greater capacity to withstand drought, as evidenced by higher g _mes, g _s and lower non-stomatal limitants, enables increased photosynthesis and rapid growth.

     

Frost damage in Pinus sylvestris L. stems assessed by chlorophyll fluorescence in cortical bark chlorenchyma

? Several techniques were used to investigate the threshold minimum temperature inducing severe frost damage during winter in Scots pine (Pinus sylvestris L.) needles and stems from southern provenances in the Iberian Peninsula.

? Chlorophyll (Chl) fluorescence, electrolyte leakage (EL), visual scoring (VS), and the normalized difference vegetation index (NDVI) were compared.

? Chl fluorescence, and in particular the maximum potential photosystem II (PSII) efficiency — estimated through the dark-acclimated F_V/F_M ratio-, was found to be a simple, non-destructive indicator of freezing tolerance in needles, as expected.

? Moreover, the existence of a Chl-containing tissue, the cortical bark chlorenchyma, allowed us the use of Chl fluorescence as a new non-destructive indicator of frost damage in live tissues of Pinus sylvestris L. stems.

? Freezing tolerance values of southern provenances are compared with those found by other authors in the northern distribution limit of the species.

     

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.

     

The effect of brown-rot decay on water adsorption and chemical composition of Scots pine heartwood

? The effect of brown-rot (Coniophora puteana) decay on the water adsorption capacity and concentration of extractives of Scots pine (Pinus sylvestris L.) heartwood were studied by comparing corresponding properties of decayed and undecayed wood samples.

? The samples derived from 39 felled trees having a large between-tree variation in the extractive concentrations, and subsequently in the mass loss in the decay test. The water adsorption capacity, expressed as equilibrium moisture content (EMC), was measured at a high relative humidity (RH ～100%, 21 °C).

? In contrast to the widely held belief, the water adsorption capacity of brow-rotted heartwood appeared to be significantly higher than that of undecayed heartwood.

? The chemical composition of heartwood was changed radically by the fungus: the concentration of stilbenes, resin acids and free fatty acids decreased, while the concentration of soluble sugars increased as a result of decay. In addition, fungal sugars were found in the decayed samples. The concentration of total phenolics increased, which obviously reflected chemical changes in cell wall constituents other than extractives.

? As a conclusion, the information concerning the hygroscopicity of brown-rotted wood might be valuable e.g. when carrying out repairs on buildings damaged by advanced decay.