Effects of nutrition and soil water availability on water use in a Norway spruce stand

We investigated effects of nutrition and soil water availability on sap flux density, transpiration per unit leaf area (EL), and canopy stomatal conductance (GS) of Norway spruce (Picea abies L. (Karst.)) in northern Sweden during the 1996 growing season. Our objectives were to determine (1) if artificially imposed drought (65% rain diversion) reduces soil water sufficiently to cause physiological limitations to whole-tree and plot-scale water transport, and (2) whether increased capacity for water transport resulting from fertilization-induced increases in leaf (> 3-fold) and sapwood areas (> 2.3-fold) deplete soil water sufficiently to cause a negative feedback on GS and EL. We monitored soil water content (theta) and soil water potential (PsiS) in control (C), drought (D), fertilized (F) and irrigated + fertilized (IL) treatment plots, along with site meteorological conditions. Ten trees per plot were monitored for sap flow. Although there were significant treatment differences in mean daily EL (C > D > F; P < 0.01) and GS (C > D > F; P < 0.05), variation in absolute magnitudes was small. Therefore, transpiration differences on a unit ground area basis (EC) were nearly proportional to leaf area differences. Precipitation was well distributed throughout the study period and so PsiS remained high, except during short dry periods in Plot F when it declined rapidly. Thus, although soil water was not limiting to GS, EL or EC when precipitation was uniformly distributed throughout the growing season, we cannot conclude that water availability would not limit GS in fertilized stands if the seasonal distribution of precipitation were altered.     

Long-term growth and yield effects of respacing natural regeneration of Sitka spruce in Britain

Stands of Sitka spruce (Picea sitchensis (Bong.) Carr.) originating from natural regeneration can be extremely dense with high mortality, poor growth form and low volume production. Although costly, respacing (pre-commercial thinning) can reduce resource competition for the remaining trees, altering the final crop produced. Three experiments were established to examine the effect of respacing 1–2 m tall trees to different target densities. The impact on stand growth and yield was measured 11–17 years after respacing, and the longer-term impact was modelled to year 50. Unrespaced control treatments self-thinned at a similar and constant rate at two sites. At the third, extremely high initial density resulted in a higher rate of self-thinning and lower density after 11 years. Wider respacing treatments resulted in larger mean tree diameter, but there was no significant effect of respacing on stand volume 11–17 years after respacing; greater diameter growth did not compensate for low tree number. Results indicate a volume penalty associated with delaying respacing until trees were 4 m tall, but this treatment was unreplicated. Modelled stand volume in year 50 was higher for the 2.6 m × 2.6 m respacing treatment than for the 1.8 m × 1.8 m, 2.1 m × 2.1 m or 3.3 m × 3.3 m treatments. However, open-grown conditions may reduce timber quality compared to closer spacing treatments. The results are presented using a stand density management diagram for Sitka spruce growing in Canada and support recent suggestions that British stands have a shallower size–density relationship than Canadian stands.     

Changes in soil,seepage water and needle chemistry between 1984 and 2004 after liming an N-saturated Norway spruce stand at the Höglwald,Germany

In 1984, a liming experiment with a surface application of 4 t ha⁻¹ of dolomitic limestone was started at the acidic N-saturated Norway spruce forest “Höglwald” in southern Germany and monitored until 2004. The decay of surface humus due to the accelerated mineralisation accounted for 18.5 ± 2.7 t ha⁻¹ C or 50% of the initial pool and 721.6 ± 115.0 kg ha⁻¹ N or 46% for N. Due to some translocation of organic material to the mineral soil the values to 40 cm depth are slightly lower (13.5 ± 4.4 t ha⁻¹ C or 15% of the initial pool and 631.6 ± 192.8 kg ha⁻¹ N or 13% for N). In the control plot NO₃⁻ concentrations at 40 cm depth were above the European level of drinking water (0.8 mmolc l⁻¹ or 50 mg NO₃⁻ l⁻¹) for nearly the whole investigation period. Liming increased NO₃⁻ concentrations in seepage water for approximately 15 years, and accelerated leaching losses by 396.2 NO₃⁻–N kg ha⁻¹ from 1984 to 2003. The increase in pH of the soil matrix was more or less restricted to the humus layer and the upper 5 cm of the mineral soil during the whole time span, while the base cations Ca and Mg reached deeper horizons with seepage water. From 1984 to 2003, an amount that nearly equalled the applied Mg, was leached out of the main rooting zone, while most of the applied Ca was retained. The time series of the elemental concentrations in needles showed minor changes. Ca concentrations in needles increased with liming, while Mg remained nearly unchanged, and P decreased in older needles.     

Response of artificial acid irrigation, liming, and N-fertilisation on elemental concentrations in needles, litter fluxes, volume increment, and crown transparency of a N saturated Norway spruce stand

The response of N-fertilisation, irrigation, acid irrigation, and liming on concentrations of elements in needles, fluxes of elements in litter, volume increment, and crown density on a full stocked, healthy, and vigorously growing mature spruce stand was investigated. The plots of the Höglwald site in Southern Bavaria exhibited a high volume increment with regularly more than 20 m³ ha⁻¹ per year, despite the high age of the stand (77 years at the beginning of the experiment in 1984). Neither a distinct growth reduction of the stand due to acid irrigation, or N oversaturation, nor an enhancement of growth due to N-fertilisation, irrigation, or liming of the stand could be detected. For the years 1984 and 1985 a marked decrease in crown density was detected for all plots. This was followed by a stagnation for 2 years. Afterwards the crown density improved until the end of the investigation for all plots. Neither acid irrigation, nor liming altered the amount of litter fall. Ca fluxes in litter, and concentrations in needles were enhanced on most of the limed plots five to six years after liming. Acid irrigation reduced Ca content in litter, but other elements were not or only slightly influenced. The nutritional status of all plots as shown by the concentrations of elements in needles indicates that for most of the years a sufficient to high supply of N, P, Ca, and Mg for all investigated plots, whereas K concentration in needles exhibited wide year to year variations. Most of the elemental concentrations in needles and fluxes in litter were not influenced by any of the treatments. Also, N-fertilisation did not enhance the N concentrations in needles significantly.     

Trade-offs between seedling growth,thinning and stand stability in Sitka spruce stands: a modelling analysis

The forest industry is increasingly adopting alternative silvicultural systems, involving regeneration beneath an existing forest canopy, rather than clear-felling and replanting. To apply these silvicultural systems in windy regions such as Britain and Ireland, it is essential that the interactions between thinning intensity, stand stability and seedling growth are properly understood. Here, we present a modelling analysis of the three key relationships between: (i) stand density and the proportion of incident radiation transmitted through a forest canopy as a stand is thinned; (ii) transmitted radiation and seedling growth, and (iii) stand density and stand stability. These relationships were examined using separate models of radiative transfer (MAESTRO), seedling growth, and stand stability/wind risk (ForestGALES). Output from the three models was synthesised to calculate whether a given stand thinned to a pre-defined stability limit would allow sufficient light to penetrate the canopy for seedling growth. A minimum transmittance of 20% was identified as a requirement for seedling growth, which corresponds to removing 45% of stand basal area. A thinning of this intensity left some stands susceptible to unacceptable wind damage, especially in old or previously thinned stands on soils where rooting is impeded. The results emphasised the fact that rooting conditions, thinning history and age of intervention are major constraints on the silvicultural options. In general, older stands are not suitable for conversion to continuous cover forestry (CCF) systems, and the transformation process should begin at pole stage, when heavy thinning does not leave the stand unstable. The analysis approach used here illustrates the potential for combining models to address complex forest management issues.     

Bending and shear properties of compressed Sitka spruce

We examined the bending and shear properties of compressed wood using small and clear specimens of Sitka spruce (Picea sitchensis Carr.). For measuring the bending properties, three-point bending tests were conducted under the span/depth ratio of 14, which is standardized in the American Society for Testing and Materials [ASTM D143-94 (2005a)] and Japanese Industrial Standards [JIS Z2101-94 (1994)]. In the bending test, the load, deflection at the midspan, and strain at the bottom of the midspan were simultaneously measured, and Young’s modulus and bending strength were obtained by elementary beam theory. For obtaining the shear modulus and shear strength, asymmetric four-point bending tests were conducted using the specimens with rectangular and side-grooved cross sections, respectively, and the influence of the compression ratio on the shear properties was examined. The results are summarized as follows: (1) Young’s modulus increased with increasing compression ratio when it was determined by the load–strain relation. Nevertheless, this tendency was rather obscured when Young’s modulus was determined by the load–deflection relation. Hence, it is preferable that Young’s modulus is measured from the load–strain relation. (2) The shear modulus in the longitudinal–tangential plane was maximum at the compression ratio of 50%, whereas that in the longitudinal–radial plane was minimum at the compression ratio of 50%. (3) The influence of the compression on the bending and shear strength ratio was not significant.     

The water vapour sorption properties of Sitka spruce determined using a dynamic vapour sorption apparatus

The water vapour sorption properties of Sitka spruce (Picea abies) have been investigated over a range of temperatures (14.2–43.8°C) using a dynamic vapour sorption apparatus. The sorption kinetics behaviour was evaluated using the parallel exponential kinetics model which has been found to give very accurate fits to the data in studies of foodstuffs or plant fibres, but has not been previously applied to sorption studies with wood. Both the adsorption and desorption kinetics curve can be deconvoluted into a fast and slow exponential process. Under conditions of adsorption, the fast process appears to be associated with the formation of monolayer water (determined using the Hailwood Horrobin model) up to a relative humidity of 20%. Under desorption, there is no clear differentiation between fast and slow processes. The area bounded by the sorption hysteresis loop reduced as the temperature at which the isotherm was measured increased, due to movement of the desorption curve only, with the adsorption curve remaining the same at all temperatures. This behaviour is consistent with sorption processes taking place on nanoporous glassy solids below the glass transition temperature. The heat of wetting was determined from the temperature dependence of the desorption isotherms by using the Clausius–Clapeyron equation, yielding results that are consistent with literature values. However, doubts are raised in this paper as to the applicability of using the Clausius–Clapeyron equation for analyses of this type.     

Vibrational properties of Sitka spruce heat-treated in nitrogen gas

Sitka spruce (Picea sitchensis Carr.) wood was heated for 0.5–16.Oh at temperatures of 120°–200°C in nitrogen gas or air. The values for Young's modulus, shear modulus, and loss tangent were measured by free-free flexural vibration tests. X-ray diffractometry was carried out to estimate the crystallinity index and crystallite width. The results obtained are as follows: (1) Density decreased at higher temperatures and longer heating times. The specific Young's modulus, specific shear modulus, crystallinity index, and crystallite width increased during the initial stage and were constant after this stage at 120°C and 160°C, whereas they increased during the initial stage and decreased later when the temperature was high. Loss tangent in the longitudinal direction increased under all conditions, whereas that in the radial direction increased at 120°C and decreased at 160°C and 200°C. (2) From the relation between Young's modulus and moisture content, it can be safely said that Young's modulus is increased by the crystallization and the decrement in equilibrium moisture content, and that crystallization (rather than degradation) is predominant at the initial stage of the heat treatment, whereas the latter is predominant as the heating time increases. (3) It is implied that the specific Young's modulus, specific shear modulus, crystallinity index, and crystallite width decreased more in air than in nitrogen gas because of oxidation in air.This study was presented in part at the 43th Annual Meeting of Japan Wood Research Society at Morioka, August 1993, the 44th Annual Meeting of Japan Wood Research Society at Nara, April 1994, and the 45th Annual Meeting of Japan Wood Research Society at Tokyo, April 1995     

Long-term effects of stump harvesting on soil properties and tree growth in Scots pine and Norway spruce stands

Abstract

Effects of stump harvesting on the properties of surface soil and on the density, structure and growth of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) stands were estimated in a field trial in western Finland. The experiment was established in 1977 and measured in 2010. Stems and logging residues were harvested after clear-cutting, and stumps were lifted and removed from half of the experimental area. Sixteen plots were planted with pine seedlings and 16 with spruce. The main effects of stump harvesting were improved survival of planted trees and an increase in natural regeneration. No clearly negative effects were noted in the stand development. Stump harvesting had no or minimal effects on the properties of the organic layer and those of the 0- to 10-cm mineral-soil layer. Soil properties did not differ between tree species. Pine production was higher on plots with stump removal compared to plots without soil treatment.     

Characterization of glutathione S-transferases in needles of Norway spruce trees from a forest decline stand

Glutathione S-transferases (GST) detoxify many electrophilic xenobiotics, including several volatile organic compounds and pesticides. The GST activity for the conjugation of several xenobiotic substances was isolated from needles of Norway spruce (Picea abies L. Karst.) trees from a forest decline stand in the northern alps. Trees that exhibited different degrees of damage were selected from several stands in an altitude profile. The GST activity toward 1-chloro-2,4-dinitrobenzene (CDNB) in crude protein extracts of needles showed a seasonal pattern with highest activity during summer. The GST activity exhibited a strong dependence on the altitude of the stand showing highest activities in trees growing in the valley and lowest activities in trees growing in the summit regions of the mountain. When cytosolic GST from needles of healthy and damaged trees was purified, trees of healthy appearance exhibited three distinct GST isozymes with activities for the conjugation of CDNB and 1,2-dichloro-4-nitrobenzene (DCNB), whereas severely defoliated trees exhibited four GSTs with additional activity for the conjugation of ethacrynic acid. The main GST isozymes catalyzing the conjugation of CDNB differed in molecular weight, isoelectric point and catalytic properties between damaged and healthy trees.     

Interception of precipitation by pole-stage Sitka spruce and lodgepole pine and mature Sitka spruce at Kielder Forest, Northumberland

Interception loss was measured indirectly for 3 years in adjacentunthinned 25-year-old stands of Sitka spruce and lodgepole pineand for 2 years in a 63-year-old selectively thinned stand ofSitka spruce. For each stand measurements were made of grossprecipitation, throughfall and stemflow. Interception loss wasderived by subtracting the sum of throughfall and stemflow (netprecipitation) from gross precipitation. For the years 1977–78, 1978–79 and 1979–80,interception losses in the pole-stage Sitka spruce were 32,28 and 27 per cent of gross precipitation respectively whilethose in the lodgepole pine were 33, 29 and 26 per cent. For1979–80 and 1980–81 interception losses in the matureSitka spruce were 44 and 53 per cent of gross precipitation.The average interception loss was 29 per cent for both pole-stagecrops and 49 per cent for mature Sitka spruce. The proportions of net precipitation reaching the ground asstemflow and throughfall were 0.18 and 0.82 respectively forthe pole-stage Sitka spruce, 0.14 and 0.86 for the pole-stagelodepole pine and 0.02 and 0.98 for the mature Sitka spruce.     

The effect of soil water-table depth on root-plate development and stability of Sitka spruce

Stability was tested on 46-year-old Sitka spruce trees (over20 m tall) growing on gleyed soils. Trees with a range of rootdepths on peaty gley and surface-water gley soils were selectedby studying their water regime. Trees were pulled over horizontallywith a winch and the vertical displacement of the soil-rootplate was measured as the load was applied. The resistive turningmoment at the point of soil failure and the maximum resistiveturning moment were measured. Thin plates, which developed overa shallow water table, had a greater surface area than thickplates that developed over deeper water tables. However, theyalso had a smaller soil mass and tended to shear in soil materialof lower shear strength and were particularly flexible. Thickerplates sheared in stronger soil and were more rigid. Soil-rootplate rigidity was shown to be a major factor affecting thecontribution of soil resistance to overturning. Regression analysisrevealed that the more rigid the plate the greater the appliedturning moment at soil failure. Trees that develop rigid soil-rootplates through adaptive growth of their structural roots willtherefore also have improved stability. The results indicatethat intensive drainage of peaty gleys will increase rootingdepth and the resistance to overturning.     

Dynamic soil processes on heathland due to changes in vegetation to oak and Sitka spruce

Soil and soil water chemistry under three different vegetation-types on a former heath area were investigated in Jutland, Denmark. An unmanaged heathland left undisturbed since the start of this century is now invaded by the oaks. Together with a small Sitka plantation (age 60 years), soil development due to vegetation changes was investigated. The typical heathland podzol soil had changed within decades under the oak trees towards an acid brown soil. The soil under the Sitka spruce had changed towards a more strongly podzolized soil compared to the former heath soil. Present processes occurring within the soil were investigated by analysis of soil solution chemistry. The leaching of inorganic ions and soluble organic matter had increased manifold under Sitka spruce. Nitrification increased markedly with soil depth probably due to effects of higher pH. The increase of nitrate with soil depth was followed by a similar increase in dissolved aluminium. The equivalence of the dissolved organic carbon varies under oak and heath between 6 and 10 m eqv g⁻¹ TOC and under Sitka the equivalence of the dissolved carbon was between 10 and 15 m eqv g⁻¹. No nitrate and only traces of ammonium were found in soil water from the upper 30 cm of the soil under heath. Soil nitrogen was almost exclusively (more than 90%) connected to the carbon cycle. Under Sitka spruce generally less than 20% of soil water nitrogen was coupled to the carbon cycle, i.e. most of the nitrogen in the soil water leaving the rooting zone under Sitka spruce was nitrate. In the oak forest, nitrate only appeared as traces in the soil water in the winter period and in the summer period in relatively higher amounts. Generally, between 80% and 90% of the nitrogen was coupled to the carbon cycle in the winter and in the summer only 50%.     

Biophysical controls on rhizospheric and heterotrophic components of soil respiration in a boreal black spruce stand

We conducted a root-exclusion experiment in a 125-year-old boreal black spruce (Picea mariana (Mill.) BSP) stand in 2004 to quantify the physical and biological controls on temporal dynamics of the rhizospheric (R(r)) and heterotrophic (R(h)) components of soil respiration (R(s)). Annual R(r), R(h) and estimated moss respiration were 285, 269 and 57 g C m(-2) year(-1), respectively, which accounted for 47, 44 and 9% of R(s) (611 g C m(-2) year(-1)), respectively. A gradual transition from R(h)-dominated (winter, spring and fall) to R(r)-dominated (summer) respiration was observed during the year. Soil thawing in spring and the subsequent increase in soil water content (theta) induced a small and sustained increase in R(h) but had no effect on R(r). During the remainder of the growing season, no effect of theta was observed on either component of R(s). Both components increased exponentially with soil temperature (T(s)) during the growing season, but R(r) showed greater temperature sensitivity than R(h) (Q(10) of 4.0 and 3.0, respectively). Temperature-normalized variations in R(r) were highly correlated with eddy covariance estimates of gross ecosystem photosynthesis, and the correlation was greatest when R(r) was lagged by 24 days. Within diurnal cycles, variations in T(s) were highly coupled to variations in R(h) but were significantly decoupled from R(r). The patterns observed at both time scales strongly suggest that the flow of photosynthates to the rhizosphere is a key driver of belowground respiration processes but that photosynthate supply may control these processes in several ways.     

Effects of repeated urea doses on soil chemistry and nutrient pools in a Norway spruce stand

Pools of macro-nutrients in soil and vegetation were studied in an old fertilization experiment with a large previous input of N. Different doses of N, in the form of urea, had been added four times during a 20-year period. In total, between 480 and 2400 kg N ha⁻¹ had been given. The experiment was established in a relatively productive Norway spruce stand and the expectation was that the large N input would cause an accelerated leaching of N, especially nitrate, accompanied by soil acidification and losses of several nutrients. The aim was to test for possible residual effects. Thirteen years after the last N addition, samples from the aboveground part of trees, field layer, S-layer, humus layer and mineral soil (0–10 cm) were analyzed for concentrations of most major nutrients. Nutrient pools were calculated. In the humus layer, the concentration of N increased and the C/N-ratio decreased with increasing N dose. The calculated recovery of added N in soil including ground vegetation was complete for the lowest N dose, while it was 25–50% for higher doses. The amount of N retained was unaffected by the N dose. The amount of extractable P in the upper part of the mineral soil was negatively correlated with N dose, as was also the concentration of total P in the S-layer. Neither soil pH, nor concentrations or amounts of Ca, Mg and K were affected by the previous fertilization. The calculated total soil-plant pool was only influenced by N dose in the case of P, which was 20% lower at the highest N dose compared with unfertilized conditions. Despite the large extra N input, the nutritional changes in plants and soil of the actual study site seemed surprisingly small.