Radial mechanical properties of high-temperature dried Norway spruce (Picea abies) wood

Abstract

Wood drying experiments were conducted in which the temperature and the drying rate were controlled independently. The mechanical properties of dried wood in radial loading were analysed in relation to drying parameters. Mass loss, due to thermal degradation of the wood structural components, occurred predominantly in slow high-temperature drying processes. Despite the higher mass loss, slowly dried wood showed similar radial strength and stiffness to rapidly dried wood. The formation of irreversible hydrogen bonds (hornification) within the wood structure may compete with the effects of mass loss on the radial mechanical behaviour of wood. However, both the mass loss and the hornification resulted in wood specimens with lower hygroscopicity. Application of slow high-temperature drying to reduce microscopic cell-wall damage, caused by anisotropic shrinkage of cell-wall layers, did not seem to affect the radial mechanical properties of wood. The effects of stress relaxation within the wood cell wall on the mechanical behaviour of wood may be offset by the degradation of structural components along with drying. Radial mechanical properties may be improved by rapid high-temperature drying up to high final dryness.     

Hydraulic and mechanical properties of young Norway spruce clones related to growth and wood structure

Stem segments of eight five-year-old Norway spruce (Picea abies (L.) Karst.) clones differing in growth characteristics were tested for maximum specific hydraulic conductivity (k(s100)), vulnerability to cavitation and behavior under mechanical stress. The vulnerability of the clones to cavitation was assessed by measuring the applied air pressure required to cause 12 and 50% loss of conductivity (Psi(12), Psi(50)) and the percent loss of conductivity at 4 MPa applied air pressure (PLC(4MPa)). The bending strength and stiffness and the axial compression strength and stiffness of the same stem segments were measured to characterize wood mechanical properties. Growth ring width, wood density, latewood percentage, lumen diameter, cell wall thickness, tracheid length and pit dimensions of earlywood cells, spiral grain and microfibril angles were examined to identify structure-function relationships. High k(s100) was strongly and positively related to spiral grain angle, which corresponded positively to tracheid length and pit dimensions. Spiral grain may reduce flow resistance of the bordered pits of the first earlywood tracheids, which are characterized by rounded tips and an equal distribution of pits along the entire length. Wood density was unrelated to hydraulic vulnerability parameters. Traits associated with higher hydraulic vulnerability were long tracheids, high latewood percentage and thick earlywood cell walls. The positive relationship between earlywood cell wall thickness and vulnerability to cavitation suggest that air seeding through the margo of bordered pits may occur in earlywood. There was a positive phenotypic and genotypic relationship between k(s100) and PLC(4MPa), and both parameters were positively related to tree growth rate. Variability in mechanical properties depended mostly on wood density, but also on the amount of compression wood. Accordingly, hydraulic conductivity and mechanical strength or stiffness showed no tradeoff.     

Stand responses to initial spacing in Norway spruce plantations in Norway

Abstract

Using data from nine spacing experiments of Norway spruce (Picea abies (L.) Karst.) in Norway, covering wide ranges of site index and initial spacing, this study evaluated stand basal area and volume responses to initial spacing and examined whether these responses varied by stand ages or site quality. We developed nonlinear regression models that described the standing basal area or volume responses to initial spacing along with site index and stand ages. The results show that closer spacings produced higher standing basal area and volume than wider spacings. The response curves are highly nonlinear in younger stands and become nearly linear in mid-rotation stands, indicating stronger responses at younger ages and weaker responses as age increases. Furthermore, for young stands, spacing effects are stronger at closer than at wider spacings. The basal area and volume responses to initial spacing tend to be similar across site indices. However, the interaction of site index and stand age on spacing responses makes it difficult to isolate the effect of site index on spacing responses. Mortality is higher and begins earlier at closer spacings than wider. The mean diameter of the largest 100, 400, 600, and 800 trees ha^?1 increased with spacing in three out of the nine experiments. Dominant height did not vary by initial spacing for any of the experiments. The findings suggest that the extra volume production in stands of closer initial spacing is restricted to early stand development.     

Fertilization effects on mean stomatal conductance are mediated through changes in the hydraulic attributes of mature Norway spruce trees

Stomatal conductance was quantified with sap flux sensors and whole-tree chambers in mature Norway spruce (Picea abies (L.) Karst.) trees after 3 years of exposure to elevated CO(2) concentration ([CO(2)]) in a 13-year nutrient optimization experiment. The long-term nutrient optimization treatment increased tree height by 3.7 m (67%) and basal diameter by 8 cm (68%); the short-term elevated [CO(2)] exposure had no effect on tree size or allometry. Nighttime transpiration was estimated as approximately 7% of daily transpiration in unchambered trees; accounting for the effect of nighttime flux on the processing of sap flux signals increased estimated daily water uptake by approximately 30%. Crown averaged stomatal conductance (g(s)) was described by a Jarvis-type model. The addition of a stomatal response time constant (tau) and total capacitance of stored water (C(tot)) improved the fit of the model. Model estimates for C(tot) scaled with sapwood volume of the bole in fertilized trees. Hydraulic support-defined as a lumped variable of leaf-specific hydraulic conductivity and water potential gradient (K(l)DeltaPsi) -was estimated from height, sapwood-to-leaf area ratio (A(s):A(l)) and changes in tracheid dimensions. Hydraulic support explained 55% of the variation in g(s) at reference conditions for trees across nutrient and [CO(2)] treatments. Removal of approximately 50% of A(l) from three trees yielded results suggesting that stomatal compensation (i.e., an increase in g(s)) after pruning scales inversely with K(l)DeltaPsi, indicating that the higher the potential hydraulic support after pruning, the less complete the stomatal compensation for the increase in A(s):A(l).     

Intra-annual variations in climate influence growth and wood density of Norway spruce

Intra-annual radial growth variations of two Norway spruce trees (Picea abies (L.) Karst.) were monitored over 4 years, at four heights up the stem, by means of point-dendrometers. The trees were then felled and radial wood samples were cut from the radii that had been monitored by the dendrometers and analyzed for density. From the radial growth measurements recorded by the dendrometers, we related positions within the rings to dates, thus making possible investigation of the relationships between changes within the rings in wood density and fluctuations in climate or growth rate. Radial growth started in early April and ended, with large intra-annual differences, in August or September. Short-term variations in growth rate were related to fluctuations in climate parameters and soil water reserves. The sensitivity of radial growth to climate decreased with stem height. Wood density responded strongly to drought events, and a dry period in June 1996 induced false-ring formation. Wood density was relatively independent of growth rate and climatic conditions during the first part of the growing season, but increased with decreasing radial growth rate later in the growing season.     

Hydraulic and anatomical properties of light bands in Norway spruce compression wood

Compression wood (CW), which is formed on the underside of conifer branches, exhibits a lower specific hydraulic conductivity (k(s)) compared with normal wood. However, the first-formed tracheids of an annual ring on the underside of a conifer branch often share several properties with normal tracheids, e.g., thin cell walls and angular cross sections. These first-formed tracheids appear bright when observed by the naked eye and are therefore called light bands (LB). In this study, hydraulic and related anatomical properties of LBs were characterized and compared with typical CW and opposite wood (OW). Measurements were made on branches of Norway spruce (Picea abies (L.) Karst.). Specific hydraulic conductivity was measured with fine cannulas connected to microlitre syringes. Micro- and ultrastructural analysis were performed on transverse and radial longitudinal sections by light and scanning electron microscopy. Xylem areas containing both typical CW and LBs had a k(s) 51.5% that of OW (7.95 +/- 0.97 m(2) s(-1) MPa(-1) x 10(-4)), whereas k(s) of pure CW was only 26.7% that of OW. The k(s) of LBs (6.38 +/- 0.97 m(2) s(-1) MPa(-1) x 10(-4); 80.3% of OW) was estimated from these k(s) values because the cannulas were too wide to measure the k(s) of LBs directly. Mean lumen area of first-formed tracheids on the underside of branches was 65.7% that of first-formed tracheids in OW and about three times that of CW. Light-band tracheids exhibited a bordered pit frequency of 42.7 +/- 1.3 pits mm(-1), which was three times that in CW and 1.6 times that in OW. Bordered pit apertures in LB tracheids (9.15 +/- 0.60 microm(2)) were 1.7 times wider than those in CW and similar in aperture to those in OW. The high k(s) of LBs was correlated with their wide tracheid lumina, high pit frequency and wide pit apertures. We therefore suggest that LBs have a primarily hydraulic function within the mechanically optimized CW region. This might be important for supplying water to living tissues on the underside of branches, as well as to other distal areas along water transport pathways following the spiral grain of wood.     

Importance of soil extractable phosphorus distribution for mature Norway spruce nutrition and productivity

Phosphorus is an essential nutrient for forest growth. In this study, we assessed the impact of soil extractable phosphorus using two simple extraction methods on nutrition and productivity of Norway spruce in sixteen mature forest stands on different bedrocks and soils in Bavaria, Southern Germany. Representative trees were sampled for needles, twigs, branches, stem bark, and stem wood. Total phosphorus content in the tree parts and soil phosphorus stock extractable with citric acid and sodium bicarbonate up to a soil depth of 80 cm were determined. We found that easily soil extractable phosphorus is a suitable indicator for estimating phosphorus uptake and stand productivity in Norway spruce. In contrast, organic layer phosphorus showed no significant correlation with aboveground biomass phosphorus contents. In the biomass, the highest phosphorus contents were measured in young needles and twigs, but the highest correlation with soil phosphorus was detected for phosphorus contents in needles and bark. The stock of phosphorus extracted by citric acid down to 40 cm soil depth revealed the best correlation with phosphorus in needles and bark. Therefore, as a supplemental or alternative method to needle analysis, our study suggests the use of phosphorus contents in stem bark to evaluate tree phosphorus nutrition. These results highlight the suitability of the citric acid soil extraction method to characterize plant available phosphorus in Norway spruce ecosystems.     

Automated image processing for quantification of blue-stain discolouration of Norway spruce wood

Bioincising is a promising method for enhancing liquid uptake (e.g. preservatives or wood-modification agents) in refractory wood. Incubation with the white-rot fungus, Physisporinus vitreus, which selectively degrades pit membranes, results in deeper and more homogeneous penetration of liquids. Conventional methods of assessing the degree of fungal discolouration of wood after treatment with preservatives (e.g. European standard EN 152) are partly based on a subjective rating scale, which gives a rough value of the surface colonisation by blue-stain fungi. Hence, an automated image processing (AIP) procedure was developed for standardised quantification of the segmentation thresholds of discolouration and tested against manual segmentation analysis. Using the red filter in the AIP method revealed high correlation (R ² 0.95) and allowed for more user friendly and objective determination of blue staining of wood.     

Effects of thinning and fertilization on wood properties and economic returns for Norway spruce

The effects of silviculture on wood and tracheid properties, and economic returns of Norway spruce (Picea abies (L.) Karst.) were investigated in two case studies, one comparing different thinning intensity in southeastern Finland and the other considering the effects of optimal nutrient addition in northern Sweden. Models for predicting the wood and tracheid properties of Norway spruce were integrated into a distance-independent process-based growth and yield model. Increasing the thinning intensity resulted in a lower mean wood density, tracheid length, and latewood proportion in harvested wood. Wood density and tracheid length of harvested pulpwood slightly decreased in later thinnings and final cuts. Thinning regimes with high early growing stock and decreasing later growing stock were most profitable. Nutrient addition accelerated volume growth and increased the value growth. The increase in volume growth due to nutrient addition more than offset the economic influence of the loss in wood density and tracheid length.     

Comparison of methods to quantify loss of hydraulic conductivity in Norway spruce

? Percent loss of hydraulic conductivity (PLC) is an important measure in plant water relations, but the available methods differ and results have rarely been compared.

? We compared PLC measured in Norway spruce (Picea abies) by quantifying conductivity before and after removing emboli, either by flushing with high water pressure or by infiltration under a partial vacuum, with relative water loss and staining conductive xylem to test advantages and possible problems of commonly applied methods.

? Infiltration removed nearly all gas from sections of drought-stressed and unstressed twigs, and flushing and infiltration efficiently removed emboli. Infiltration appears less prone to producing clogged xylem elements than high pressure flushing. Relative water loss (RWL) and the proportion of xylem stained with phloxine B (PSX) was also highly correlated with PLC, the latter can be quantified by image analysis, and also shows the pattern of xylem dysfunction. Loss of conductivity was more common in the inner annual rings, in early wood within an annual ring, and in compression wood, though pattern differed substantially between branches.

? Advantages and potential problems of these methods are discussed and it is suggested that RWL or PSX may be used as proxy measures for PLC in species when the correlations have been established.

     

Effects of experimental drought on the fine root system of mature Norway spruce

Norway spruce (Picea abies (L.) Karst.) is an economically important, but relatively drought-sensitive tree species that might suffer from increasing drought intensities and frequencies, which are predicted to occur in parts of central Europe under future climatic change. In a throughfall exclusion experiment using sub-canopy roofs, we tested the hypothesis that enhanced drought leads to an increase in fine root mortality, and also to a higher, subsequent fine root growth. Fine root production and mortality were assessed using two independent approaches, sequential soil coring (organic layer) and direct observations in minirhizotrons (organic layer plus upper mineral soil). Six weeks of throughfall exclusion resulted in mild drought stress, i.e. a reduction in average soil moisture from 20 to 12 vol.% during the treatment. Based on the sequential coring data, experimental drought did not result in significant changes in fine root biomass during the 6-week treatment period, but caused an increase in fine root mortality by 61% in the 6 weeks following the drought treatment. Remarkably, fine root production showed a synchronous increase in this period, which more than compensated for the loss due to increased mortality. The minirhizotron data confirmed that the drought treatment increased fine root loss in the organic layer. Based on this method, however, root loss occurred during the drought period and was not compensated by increased root production. The mild drought stress was mainly restricted to the organic layer and did not significantly influence fine root dynamics in the mineral soil. We calculated that the drought event resulted in an extra input of about 28 g C m⁻² and 1.1 g N m⁻² to the soil due to increased fine root mortality. We conclude that even periods of mild drought significantly increase fine root mortality and the associated input of root-derived C to the soil organic matter pool in temperate Norway spruce forests.     

Occurrence of Heterobasidion basidiocarps on cull pieces of Norway spruce left on cutting areas and in mature spruce stands

Fruiting of Heterobasidion on cull pieces and stumps of Norway spruce was investigated in cutting areas and mature spruce stands located in southern Finland. Cull pieces of variable size and showing butt rot were left on three clear‐cut areas and in one thinned stand. Additionally, a part of the cull pieces was transported to mature forest sites with closed canopy. During the succeeding 3–4 years the cull pieces were investigated annually for sporocarps of Heterobasidion, and the area of actively sporulating pore layer of each sporocarp was measured. Root bases of spruce stumps in the logging areas were excavated and sporocarps found on the stumps also measured. At the onset of the experiment, Heterobasidion spp. were isolated from 76% of the cull pieces showing butt rot; 85% of the isolates were identified as H. parviporum and 15% as H. annosum s.s. During the following 3–4 years sporocarps were found on 20% of the 1938 cull pieces where Heterobasidion butt rot was initially detected visually. Sporocarp formation was promoted by advancement of butt rot, increasing cull piece diameter and end‐to‐end ground contact, but restricted by the colonization of the cull piece by Stereum sanguinolentum. Between‐site differences were significant but could not be explained by differences in tree cover. At the end of the investigation period the average sporulating area of Heterobasidion sporocarp per metre of cull piece was higher than the average sporulating area per stump at three of four logging sites. Hence, leaving cull pieces containing Heterobasidion butt rot at logging areas in southern Finland can considerably increase local production of Heterobasidion spores.     

Hydraulic and stomatal adjustment of Norway spruce trees to environmental stress

A study of how the water conducting systems of 30-50-year-old Norway spruce (Picea abies (L.) Karst.) trees growing at three sites adjust to shade and waterlogging indicated that water relations characteristics varied with the life histories of the trees. Xylem was more efficient at conducting water and stomata were more sensitive to atmospheric evaporative demand in trees subjected to favorable growth conditions (control trees) than in trees growing in shade or waterlogged conditions. At the same soil water availability, shade-grown trees suffered more severely from water deficit than control trees. Under conditions of high atmospheric vapor pressure deficit, foliage of shade-grown trees exhibited low water potentials, as a result of low hydraulic conductance of the vascular system and inefficient stomatal control. Because of the increased internal resistance to water flow, more negative leaf water potentials (Psi(x)) must be reached to provide an adequate water supply to the foliage. It is concluded that dynamic water stress is one of the main causes of the continuing growth retardation in suppressed Norway spruce trees after their release from the overstory. Trees growing in waterlogged soil (bog-grown trees) were characterized by weak stomatal control, resulting in large water losses from the foliage. Although bog-grown trees exhibited uneconomical water use, they possessed mechanisms (e.g., osmotic adjustment) that allowed leaves to tolerate low Psi(x) while stomata remained open. Under conditions of sufficient soil water availability and moderate atmospheric vapor pressure deficit, soil-to-leaf conductance was highest in bog-grown trees (1.45 +/- 0.06 mmol m(-2) s(-1) MPa(-1)), followed by control and shade-grown trees (1.04 +/- 0.04 and 0.77 +/- 0.05 mmol m(-2) s(-1) MPa(-1), respectively). The lowest soil-to-leaf conductance (0.45 +/- 0.04 mmol m(-2) s(-1) MPa(-1)) was recorded in control trees at high atmospheric evaporative demand, and was probably caused by tracheid cavitation.     

Growth recovery of mature Norway spruce and European beech from chronic O<Subscript>3</Subscript> stress

Elevated O₃ levels can strongly impair the health and vitality of forest ecosystems. Free-air exposure systems reveal that forest tree and stand growth can be reduced strongly under chronic O₃ stress. Detailed knowledge of the effect of O₃ exposure on photosynthesis, carbon sequestration, allometry and growth during chronic stress is available. However, knowledge of growth response after O₃ reduction is scarce. Here, we analyse the growth of mature Norway spruce (Picea abies (L.) Karst.) and European beech (Fagus sylvatica L.) in the free-air O₃ fumigation experiment at Kranzberg Forest. We compare tree growth over a 9-year period (2008–2016) after exposure to O₃ (2000–2007). During 2?×?O₃ exposure, the annual basal area growth of Norway spruce and European beech decreased by 24 and 32%, respectively. After cessation of 2?×?O₃ exposure, the annual basal area growth of Norway spruce and European beech not only recovered but exceeded the growth of the trees in the control condition by 14 and 24%, respectively. The growth resilience and resistance of trees previously exposed to 2?×?O₃ towards drought stress and late frost was hardly lower than that of the trees in the control condition. The capacity for growth recovery even after long-term chronic O₃ stress emphasizes the strong beneficial effect of air pollution control on the health of forest ecosystems and on the global land carbon sink.     

Occurrence of microorganisms in the wood of Norway spruce trees from polluted sites

Bacteria, yeasts and other fungi were isolated from stem and root wood of Norway spruee trees with disease symptoms and were identified in order to evaluate a possible mierobial involvement in the widespread tree decline in Central Europe.     

Control of tracheid cross-sectional dimensions in Norway spruce and Scots pine wood raw material

Abstract

Wood properties, including tracheid cross-sectional dimensions, show a large degree of variation. To improve the properties of products made from wood, different methods to control variation have been developed. This study aims to determine the theoretical efficiency of three control strategies: the fractionation of pulped tracheids into earlywood and latewood, the separation of juvenile and mature wood, and sorting of logs according to tree size. The efficiency of each method was studied by first constructing virtual trees from measured tracheid cross-sectional dimensions, then simulating the efficiency of above-mentioned methods. The tracheid dimension data include Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.). The simulations show that separation into earlywood and latewood classes has the highest theoretical efficiency and yields the lowest variances in raw material. Classification into juvenile and mature wood groups is the second most efficient method, and the sorting of logs according to the size class of the tree is the least efficient method. It was also concluded that the variation in cell-wall thickness and radial diameter mainly originates from differences between earlywood and latewood, whereas the variation in tangential diameter mainly originates from differences between mature and juvenile wood.     

Interactions between soil scarification and Norway spruce seedling types

Interactions between scarification treatments and seedling types of Norway spruce were examined at two different locations in southern Sweden. The scarification treatments were not scarified control, mounding and soil inversion and the seedling types were a 10 week-old containerized seedling (mini seedling), a 2 year-old containerized seedling and a 2 year-old hybrid seedling, grown as a containerized seedling the first year and as a bare-root seedling the second year. Site preparation intensity had a greater effect on the mini seedlings. The results indicated that mini seedlings established faster in the soil inversion treatment compared to the larger seedling types. At the same seedling age, the mini seedlings had faster or similar growth rate as the containerized and the hybrid seedlings regardless of scarification treatment. Although mini seedlings grew as well as or even better than larger seedlings if they were successfully established, they were more sensitive to their planting environment and proper handling was critical. Problems with frost heaving and competing vegetation have to be taken into consideration when choosing site preparation method and seedling type.     

Comparison of colonization capacity by asexual spores of Heterobasidion species in Norway spruce wood

Two species of the tree pathogenic fungus Heterobasidion spp. exist in Sweden, Heterobasidion annosum s.s. and Heterobasidion parviporum. Both species are known to infect Norway spruce (Picea abies). The aim of the study was to examine the interspecific competition between H. annosum s.s. and H. parviporum as well as their colonization rate in fresh Norway spruce wood. Equal amount of conidiospores from each species was sprayed together on 30 fresh, previously uninfected, Norway spruce billets. After incubation in a greenhouse, the proportion of Heterobasidion spp. colonies belonging to each species was recorded. Of the 196 colonies isolated from the upper part of billets, 195 were H. parviporum. All isolated colonies further down in the billets were H. parviporum. To study the colonization rate, H. annosum s.s. and H. parviporum were sprayed alone on 30 spruce billets each, incubated and growth recorded both vertically and horizontally. H. parviporum grew further down in the billets (p = 0.008) and covered a larger area (p < 0.001) than H. annosum s.s. While H. annosum s.s. and H. parviporum both infected fresh Norway spruce wood H. parviporum outgrew and outcompeted H. annosum s.s during the early colonization stage.     

Influence of nitrogen and phosphorous availability and ozone stress on Norway spruce seedlings

Four-year-old Norway spruce (Picea abies L. (Karst.)) seedlings were exposed to ambient and elevated (1.5 x ambient in 1997 and 1.6 x ambient in 1998) ozone concentrations [O3] and three nitrogen (N) and two phosphorus (P) availabilities: "optimal" values (control); 70% of the control N and P values (LN and LP); and 150% of the control N value (HN). Treatments were applied in an open-field ozone fumigation facility during the 1997 and 1998 growing seasons. Effects on growth, mineral and pigment concentrations, stomatal conductance and ultrastructure of needles were studied. The HN treatment increased growth significantly, whereas elevated [O3] had a slight or variable impact on growth and biomass allocation in all N treatments. Although there were no significant effects of the LP treatment on plant growth during the second year, there was a reduction in 1-year-old shoot dry mass in the elevated O3 + LP treatment at the end of the experiment. There were no significant treatment effects on mineral concentrations of current-year and 1-year-old needles at the final harvest. In response to the HN treatment, chlorophyll a and b and carotenoid concentrations increased significantly in current-year needles. Chlorophyll a/b ratio decreased in response to elevated [O3] alone, but increased in seedlings in the O(3) + LP treatment. Stomatal conductance of current-year needles decreased with increasing N availability, but increased in response to elevated [O3]. However, the O3-induced increase in stomatal conductance was less in the LN and LP treatments than in the control treatment. In chloroplasts of current-year needles, increased N availability decreased mean starch grain area, but increased the number of plastoglobuli. We conclude that Norway spruce seedlings are relatively tolerant to slightly elevated [O3], and that nitrogen and phosphorus imbalances do not greatly affect the influence of O3 on this species when the exposure lasts for two growing seasons or less.     

Chemical composition of earlywood and latewood in Norway spruce heartwood,sapwood and transition zone wood

This study focused on the distribution of wood components along a cross section of a spruce stem. Thin samples of earlywood and latewood were analysed by special micro-scale analytical techniques. Heartwood contained significantly more lignin and less cellulose than sapwood. The total content of hemicelluloses was the same along the radial direction, but the distribution of sugar units differed. The amounts of arabinoglucuronoxylan and pectins were larger in the heartwood. The transition zone between heartwood and sapwood had a specific composition, with less lignin and lipophilic extractives than heartwood and sapwood. For earlywood and latewood, significant differences were found in the distribution of sugar units in hemicelluloses. Latewood contained clearly more galactoglucomannan than earlywood, and conversely less pectins. The lipophilic extractives were also less concentrated in the latewood.Abbreviations EW or E earlywood - LW or L latewood - HW heartwood - SW sapwood - TZ transition zone wood - A.R. annual ring - AcBr Acetyl bromide - Ara arabinose - Xyl xylose - Gal galactose - Glc glucose - Man mannose - Rha rhamnose - GlcA glucuronic acid - MGlcA 4-O-methyl-glucuronic acid - GalA galacturonic acid - o.d. oven dry