Phosphorus cycling and soil P fractions in Douglas-fir and red alder stands

Nitrogen-fixing species can dramatically increase soil acidity and organic matter content, and potentially alter biogeochemical P dynamics. We compared ecosystem P cycling in adjacent stands of N₂-fixing red alder (Alnus rubra Bong.) and non-fixing Douglas-fir (Pseudotsuga menziesii Franco) in order to determine whether P-cycling rates within stands were related to soil P forms as measured by sequential P fractionation. Above-ground annual P uptake was 61% greater in the red alder stand, although soil available P, as measured by Bray (NH₄F–HCl) extraction, was only 10% of that found in the Douglas-fir stand. Total ecosystem P in the alder stand was only 69% of that found in the Douglas-fir stand, and could indicate a pre-establishment difference between stands. However, the percentage of total soil P released by Bray or NaOH extraction was also lower in the alder stand, which suggests that differences in total P alone did not control the patterns observed in P fractions. Concentrations of inorganic P sorbed to Fe and Al minerals and contained in Fe minerals and apatite were greater under Douglas-fir, while organic P was slightly greater under red alder. While fluxes of P in litterfall, uptake and resorption were 94, 60 and 292% higher in the alder stand, soil extractable fractions meant to represent available P were lower under alder. Static measures of available P do not appear to adequately reflect P supply, and the development of techniques to assess P turnover is needed to better understand cycling and plant availability of P.     

A comparison of annual transpiration and productivity in monoculture and mixed-species Douglas-fir and red alder stands

Although much is known about drivers of productivity in Douglas-fir and red alder stands, less is known about how productivity may relate to stand transpiration and water use efficiency. We took advantage of a 15-year-old experiment involving Douglas-fir (Pseudotsuga menziesii) and red alder (Alnus rubra) in the western Cascade Range of western Oregon to test the following hypotheses: (a) more productive stands transpire more water, (b) the relationship between productivity and transpiration differs between species, and (c) the relationship between productivity and transpiration differs between sites varying in soil moisture and fertility. Furthermore, the experimental design included alder, a facultative nitrogen-fixing species, which could also affect fertility. Fixed area plots (20 × 20 m) were planted as monocultures of each species or in mixtures at a common density (1100 trees ha⁻¹) in a randomized-block design. Transpiration of Douglas-fir and red alder was measured using heat dissipation sensors installed in eight trees per plot and scaled to the plot level based on sapwood basal area for each species. Although up to 53% of the variability in tree transpiration was explained by basal area, irrespective of species or site conditions, the two stands with the highest biomass and sapwood basal area did not transpire the most. Instead of more productive stands transpiring more water, the greatest variability in both productivity and transpiration was determined by site conditions and to a lesser degree, species composition. For example, 70% of the variation in tree biomass increment (TBI) was determined by leaf area index, which was much higher at the site with higher fertility and soil moisture (p < 0.05). Despite marked phenological and physiological differences, Douglas-fir and red alder performed similarly. Only 19% of annual water use of Douglas-fir occurred between October and March when alder was leafless. Also, there was no evidence of a fertilization effect of the nitrogen-fixing red alder on the Douglas-fir: the nitrogen concentration and N-isotopic ratio of Douglas-fir needles did not differ whether trees were grown in monoculture or in mixtures with red alder. We conclude that lower soil fertility and contrasting microclimate at one site relative to the other suppressed NPP while maintaining higher transpiration, thus reducing water use efficiency.     

Ecosystem production in Douglas-fir plantations: Interaction of red alder and site fertility

Pure stands of Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) and mixed stands of Douglas-fir and naturally established red alder (Alnus rubra Bong.) were examined on two different sites for soil properties, tree growth and nutrition, and aboveground ecosystem biomass and net primary production. One site (Mt. Benson, Nanaimo, B.C.) was nitrogen (N)-deficient and had a low site index (expected Douglas-fir height of 24 m at 50 years). The other site (Skykomish, western Washington) was N-rich and had a site index of 45 m at 50 years. Soil N accretion on the red alder units was estimated at 65 (Mt. Benson) and 42 (Skykomish) kg ha^?1 year^?1 for 23 years to a soil depth of 50 cm. At the current stage of plantation development, presence of red alder at the infertile Mt. Benson site increased average Douglas-fir diameter but did not affect its basal area and basal area growth rate; including alder stem biomass increased total stand basal area and basal area growth 2.5 fold. Presence of red alder at the fertile Skykomish site decreased average diameter, basal area, and basal area growth of Douglas-fir; including alder biomass left total stand basal area and basal area growth unchanged. Douglas-fir foliar N concentrations on Mt. Benson increased from 0.93 without alder to 1.41% on the red alder unit but were 1.55% for both units at Skykomish. Although alder did not affect Douglas-fir aboveground biomass and net primary production on Mt. Benson, total ecosystem biomass doubled and production tripled when alder biomass was included. Conversely, at Skykomish, Douglas-fir biomass and production decreased, and total ecosystem values were essentially unchanged. Mixing red alder and Douglas-fir seems to have great potential for increasing Douglas-fir growth and ecosystem production on infertile, N-deficient sites but probably has limited value on fertile, N-rich sites.     

THE CHARACTERISTICS OF THE NUTRIENT CYCLING IN BIRCH STANDS WITH DIFFERENT AGE STAGES

SuccessionisthedynanucdeveloPInentofecosystemswhichcanbecharactetindbyenergyandndneralcyclings.Eachsuccessionstagehasitscharaceristicsofelementcycling.SomeschOlarshaveStatedbutbrieflawsofnutrienCyclinginsuccessionstagesI"3I.InthespaPer,thenutrientcyclingcharacterishcsofbirchsuccessionseriesinXiaokinganlingwerediscussed.SwrYAasAroarernoDSThestUdyareaislocatedinLiangshniNatUralReservesinthendddieofXiaox-ing'anlilg(l28"53,2o'rE,47olo'5O"N).Theclimat6belongstocontinentalmonsoonclimate,W…     

Annualized diameter and height growth equations for Pacific Northwest plantation-grown Douglas-fir, western hemlock, and red alder

Simulating the influence of intensive management and annual weather fluctuations on tree growth requires a shorter time step than currently employed by most regional growth models. High-quality data sets are available for several plantation species in the Pacific Northwest region of the United States, but the growth periods ranged from 2 to 12 years in length. Measurement periods of varying length complicate efforts to fit growth models because observed growth rates must be interpolated to a common length growth period or those growth periods longer or shorter than the desired model time step must be discarded. A variation of the iterative technique suggested by Cao [Cao, Q.V., 2000. Prediction of annual diameter growth and survival for individual trees from periodic measurements. Forest Sci. 46, 127–131] was applied to estimate annualized diameter and height growth equations for pure plantations of Douglas-fir, western hemlock, and red alder. Using this technique, fits were significantly improved for all three species by embedding a multi-level nonlinear mixed-effects framework (likelihood ratio test: p < 0.0001). The final models were consistent with expected biological behavior of diameter and height growth over tree, stand, and site variables. The random effects showed some correlation with key physiographic variables such as slope and aspect for Douglas-fir and red alder, but these relationships were not observed for western hemlock. Further, the random effects were more correlated with physiographic variables than actual climate or soils information. Long-term simulations (12–16 years) on an independent dataset using these annualized equations showed that the multi-level mixed effects models were more accurate and precise than those fitted without random effects as mean square error (MSE) was reduced by 13 and 21% for diameter and height growth prediction, respectively. The level of prediction error was also smaller than an existing similar growth model with a longer time step (ORGANON v8) as the annualized equations reduced MSE by 17 and 38% for diameter and height growth prediction, respectively. These models will prove to be quite useful for understanding the interaction of weather and silviculture in the Pacific Northwest and refining the precision of future growth model projections.     

Nutrition and bud removal affect biomass and nutrient allocation in Douglas-fir and western red cedar

Seedlings of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and western red cedar (Thuja plicata J. Donn ex D. Don) were grown at high (250 mg l(-1)) and low (20 mg l(-1)) nitrogen (N) supply for a year. Before the second growing season, half of the seedlings in each nutrient treatment were allocated to the other treatment. Half of the seedlings in each nutrient treatment then had all growing points removed. Biomass and N, phosphorus (P) and potassium (K) concentrations of old and new shoots and roots were measured three times in the second year to test the interaction of current-year and previous-year nutrient supply on biomass and nutrient allocation in these two species with different growth habits. Pruned seedlings served as controls. Unpruned seedlings of both species increased in height throughout the second growing season, except for Douglas-fir in the N250 --> N20 treatment. Repeated pruning did not prevent new shoot growth, but resulted in a 12 to 52% reduction in biomass of new shoots and new and old roots. Seedlings receiving a low N supply in the first growing season were more severely affected by pruning than seedings receiving a high N supply. Growth was reduced more by pruning in western red cedar than in Douglas-fir. Concentrations of N, P and K were higher in pruned seedlings than in unpruned seedlings. Although dry weights of all plant parts in all treatments increased throughout the second growing season, some retranslocation of N, P and K was observed from old shoots of both species in the N250 --> N20 and N20 --> N20 treatments after August. Quantities of N, P and K retranslocated were greatest in seedlings grown the previous year in the high-N treatment.     

Long-term effects of repeated urea fertilization in Douglas-fir stands on forest floor nitrogen pools and nitrogen mineralization

In six Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] stands in the Puget Sound Region in Western Washington/USA, forest floor C and N pools were quantified on control plots and on plots that had been fertilized repeatedly with urea 8–30 years ago (total amount of applied N 0.9–1.1 Mg ha⁻¹). Additionally, net N mineralization and nitrification rates were assessed in field and laboratory incubation experiments. Forest floor C/N ratios were decreased on the fertilized plots of all sites compared to the respective control plots. The decreases were particularly strong at sites with initial C/N ratios larger than 30. On sites with low productivity (site index at age 50: <33 m), N fertilization resulted in considerable increases in forest floor N pools. Net N mineralization and nitrification during 12-week field incubation was negligible for the unfertilized and fertilized plots of all except one site (Pack Forest), where the stand had been clear-cut 2 years ago. The increases in N mineralization rates during 12-week laboratory incubation induced by repeated N fertilization showed an inverse relationship to the time elapsed since the last fertilizer application, and were generally larger at sites with initial forest floor C/N ratios >30. For the investigated sites, fertilization effects on net N mineralization sustained for at least 11 years after the last fertilizer application. Nitrification correlated strongly with the forest floor pH; significant formation of NO₃⁻ was observed only for O layers with a pH (H₂O) higher than 4.5.     

Long-term effects of nitrogen fertilization on the productivity of subsequent stands of Douglas-fir in the Pacific Northwest

The carryover effects of N fertilization on five coastal Pacific Northwest Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) plantations were studied. “Carryover” is defined as the long-term impact of N fertilizer added to a previous stand on the growth of a subsequent stand. Average height and diameter at 1.3 m above-ground (DBH) of 7–9-year-old Douglas-fir trees and biomass and N-content of understory vegetation were assessed on paired control (untreated) and urea-N-fertilized plots that had received cumulative additions of 810–1120 kg N ha⁻¹ to a previous stand. Overall productivity was significantly greater in the fertilized stands compared to the controls. In 2006, the last growth measurement year, mean seedling height was 15% greater (p = 0.06) and mean DBH was 29% greater (p = 0.04) on previously fertilized plots compared to control plots. Understory vegetation biomass of fertilized plots was 73% greater (p = 0.005), and N-content was 97% greater (p = 0.004) compared to control plots. These results show that past N fertilization markedly increased seedling growth in these plantations as well as biomass and N-content of understory vegetation in a subsequent rotation. These findings suggest that N fertilization could potentially increase site productivity of young Douglas-fir stands found on low quality sites in the Pacific Northwest 15–22 years after application by a carryover effect. These plantations have not yet reached the age where marketable materials can be harvested from them, and the growth of trees should be monitored over a longer time period before potential impacts on older stands, if any, can be determined.     

Belowground carbon pools and processes in different age stands of Douglas-fir

Forest floor material and soil organic matter may act as both a source and a sink in global CO2 cycles. Thus, the ecosystem processes controlling these pools are central to understanding the transfers of carbon (C) between the atmosphere and terrestrial systems. To examine these ecosystem processes, the effect of stand age on temporal carbon source-sink relationships was examined in 20-year-old, 40-year-old and old-growth stands of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) in the Cascade Mountains of south-central Washington State. Belowground C and nitrogen (N) storage and soil respiration were measured. In addition, nylon mesh bags containing homogenized soils from each site were buried at the respective sites to quantify root ingrowth and potential C sequestration and loss. The sites supporting the 20- and 40-year-old stands had soil C stores reflecting the C contributions from logging residue, coarse woody debris and stumps left after harvest. Because the N-fixer red alder (Alnus rubra Bong.) comprised 33% of the 40-year-old stand, this site had significantly greater concentrations and pools of N in the forest floor than sites without red alder. This N-rich site had consistently lower soil CO2 efflux rates during the growing season than the sites supporting the 20-year-old and old-growth stands. Estimated annual soil C efflux was 1367, 883 and 1194 g m-2 for the sites supporting the 20-, 40- and old-growth stands, respectively. These values are higher than previously reported values. Root ingrowth was significantly less in the 40-year-old stand than in the 20-year-old stand, and both young stands showed markedly less fine root growth than the old-growth stand. At the sites supporting the young stands, C and N were lost from the soil bags, whereas there was an increase in C and N in the soil bags at the site supporting the old-growth stand. The fine root growth and soil respiration data support the hypothesis that belowground C allocation decreases with increasing fertility. Quantification of the source-sink relationship of soil C at the three stands based on litterfall, relative root ingrowth and soil respiration measurements was compromised because of significant CO2 flux from decaying organic matter in the young stands.     

Red alder leaf decomposition and nutrient release in alder and conifer riparian patches in western Washington,USA

Current management practices encourage conversion of red alder (Alnus rubra) riparian forests to conifers in the Pacific Northwest U.S. Patches of young naturally regenerated conifers are commonly present in alder dominated riparian areas and an understanding of the soil processes in these patches will be helpful in guiding future riparian management. Study objectives were to: (1) determine decomposition rates of red alder leaves in riparian alder and conifer patches, (2) relate decomposition rates to environmental factors and litter chemistry, and (3) determine nutrient release from decomposing alder leaves in these patches. Study sites were riparian areas adjacent to Brown and Le Bar creeks in the Skokomish River basin, Olympic National Forest, Washington. Red alder leaves were placed in litterbags in red alder and conifer riparian patches along each stream in November 2000 and collected after 1 and 3 years. There was rapid mass loss of alder leaves in the first year in both patch types, but decomposition was significantly faster (p < 0.05) in alder patches (43.2% mass remaining, k = 0.855 year⁻¹) than in conifer patches (48.4% mass remaining, k = 0.734 year⁻¹). There was little mass loss after the first year and no significant difference in decomposition rates. After 3 years mass remaining was 44.2% (k = 0.283 year⁻¹) and 47.8% (k = 0.48 year⁻¹) in alder and conifer patches, respectively. Decomposition rate differences were attributed more to the effects of the different litters in each patch and the influence on soil microbial and faunal communities than differences in soil temperature and moisture. The forest floor was deeper in conifer patches (3.7 cm) than alder (1.8 cm) patches. This was ascribed to slower decomposition rates in conifer patches, greater litterfall in conifer patches, and/or removal of alder surface litter by flooding. Alder patches were lower in elevation (0.8 m above bankfull width) than conifer patches (2.2 m). Forest floor and soil C and N concentrations and pHs were not significantly different in alder and conifer patches. Nutrient release from decomposing alder leaves was not significantly different in conifer and alder patches, although there was a trend for C, N, P, K, and Ca to be lost faster from leaves in alder patches than conifer patches in the first year. Red alder litter input to riparian conifer patches will initially decompose rapidly and provide nutrients, particularly N and P to conifers, as well as enhancing soil C since long-term decomposition rates are slow.     

Productivity and nutrient cycling of a Javanese homegarden

In a 0.13 ha homegarden in West Java, Indonesia, a 16-week study of production, nutrient in- and outputs as well as internal circulation was carried out in 1989. Total annual agricultural production was 11.4 t ha^–1, of which 6.8 t ha^–1 were timber and firewood. Two thirds of the production was sold, the rest consumed by the owners. The productivity is more than twice the productivity of ricefields in the area, but can be raised further. Nutrient inputs in rainfall, streamwater and via N-fixation were estimated to a total of: 33 kg N, 7 kg P, 115 kg K, 79 kg Ca and 55 kg Mg ha^–1 yr^–1. Outputs in streamwater, harvest sold and erosion were estimated at: 42 kg N, 11 kg P, 151 kg K, 278 kg Ca and 83 kg Mg. Litterfall and pruning returned 10.0 and 7.5 t dry matter, respectively, per hectare to the soil annually. Throughfall was on average 92% of incident rainfall. Total amounts of nutrients circulated internally in the homegarden each year were estimated at: 223 kg N, 38 kg P, 373 kg K, 135 kg Ca and 50 kg Mg per hectare. This corresponded to 22, 44, 50, 20 and 30% of nutrients stored in the plant biomass, respectively. The balance is negative for all elements, the magnitude depending on whether the part of harvest consumed by the family itself is regarded as internal circulation or as an output. Compared to available soil reserves the system is likely to be sustainable for many years ahead, but may be vulnerable because of the high fraction of plant nutrient storage cycled annually.     

Estimating understory vegetation response to multi-nutrient fertilization in Douglas-fir and ponderosa pine stands

Models were developed to predict understory vegetation response to multi-nutrient fertilization at six conifer-forested stands in the inland Northwest United States. Equations are presented to estimate how fertilization as well as other factors impacting understory production in the inland Northwest change total understory vegetation production and the production of three individual lifeforms (shrubs, forbs, and grasses and grass-likes). Overstory stand density was found to have the greatest impact on understory production, and regardless of factors such as fertilization or precipitation, large stand densities will limit understory production. At lower stand densities, multi-nutrient fertilization as well as greater amounts of precipitation will increase understory production. These factors were also found to be synergistic; thus, greater amounts of precipitation increase the effects of multi-nutrient fertilization on understory production. For sites of the same stand density, Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] was shown to have a greater negative impact on understory production than ponderosa pine (Pinus ponderosa Dougl. ex Laws.). The models predict that multi-nutrient fertilization of ponderosa pine stands will produce increases in understory production across a broader range of stand densities.     

Underplanting western hemlock in a red alder thinning: early survival,growth, and damage

Underplanting conifers beneath thinned hardwood stands could shorten conversion to a more valuable crop species. Thinning the hardwoods to final crop-tree spacing could increase growth and marketable volume while releasing the site resources necessary to support conifers planted in the understory. In western Oregon, an experimental thinning of a 14-year-old red alder (Alnus rubra Bong.) stand provided an opportunity to test this concept. Initial efforts were directed toward testing survival and growth of wilding seedlings of western hemlock (Tsuga heterophylla [Raf.] Sarg.) planted in thinned and unthinned alder stands. Survival averaged 78% and 52% after the first and fifth growing seasons. Fifth-year height growth of surviving seedlings averaged from 38 to 49 cm under various thinning regimes but only 9 cm in the unthinned. Wildlife browsing and the pinning of seedlings by falling debris reduced growth and survival.     

Branch mortality and potential litterfall from Douglas-fir trees in stands of varying density

Forest floor characteristics influence nutrient cycling and energy flow properties of forest ecosystems, and determine quality of habitat for many forest plants and animals. Differential crown recession and crown development among stands of differing density suggest that an opportunity may exist to control the input of fine woody litter into the system by manipulating stand density. The objective was to measure the rate of branch mortality among stands of differing density and to estimate the range in total per hectare necromass inputs. Although litter traps are reliable for estimating per hectare rates of litterfall, branch mortality dating on sectioned stems uniquely allows assessment of several other litterfall components: (1) individual tree contributions to total litterfall; (2) the amount of branch material released by mortality, regardless of whether the branches are shed to the forest floor; (3) the distribution of basal diameters characterizing the litterfall from a given tree and stand. Twenty-four trees were felled and sectioned on permanent plots that were part of a silvicultural study of stand density regimes in Douglas-fir (Pseudotsuga menziesii (Mirbel) Franco.). Whorl branches were dissected out of bole sections to determine the dates of mortality, and a branch biomass equation was applied to estimate potential rate of litterfall. Periodic annual rates were expressed in four ways: (1) number of branches per tree; (2) mass of branches per tree; (3) mass of branches per unit of crown projection area; (4) mass of branches per hectare. For the growth periods investigated, larger trees and trees growing on denser plots tended to release a greater necromass through branch mortality. Average branch basal diameter generally decreased with increasing stand density. Annual branch mortality ranged from 33 to 430 g m⁻² crown projection area for individual trees, and from 236 to 1035 kg ha⁻¹ for individual plots. These rates approached the low end of the range of previously published fine litterfall rates for Douglas-fir. Rates on these plots were relatively low owing to the temporary delay in crown recession imposed by artificial thinning. A conceptual model of branch litter dynamics is presented to depict consistencies with crown development among stands managed under different density regimes.     

Evidence for red:far red signaling and photomorphogenic growth response in Douglas-fir (Pseudotsuga menziesii) seedlings

In a greenhouse experiment, potted coastal Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings were grown in miniature "Nelder" (Nelder 1962) plots where growing space varied from 265 to 2555 cm(2) per plant. After thirty weeks, mean plant height, crown biomass and branch number increased significantly (P = 0.0141) with decreasing growing space (increasing plant density). Differences in height growth became apparent about six weeks after sowing. Furthermore, horizontally reflected radiation measured within the Nelder plots showed a decrease in red:far-red ratio (R:FR) from 1.2 at the lowest density to 0.71 at the highest. Plant height was strongly inversely correlated with estimated phytochrome photoequilibrium values (r(2) = 0.893). Field measurements made in a three-year-old variable density plantation also showed a decrease in R:FR with increasing planting density from 300 to 3,000 trees ha(-1). These results support the hypothesis that young Douglas-fir seedlings are able to detect, through the phytochrome system, the presence of nearby seedlings owing to the depletion of R relative to FR in the spectra reflected by the foliage of the adjacent plants. They then adjust their growth allometry in a way that reduces the possibility of being over topped by these future competitors.     

Evaluation through a simulation model of nutrient exports in fast-growing southern European pine stands in relation to thinning intensity and harvesting operations

The effects on nutrient exports of a range of thinning regimes for maritime pine and radiata pine plantations in northern Spain were simulated in this study. Growth models, tree biomass equations and nutrient concentration in tree fractions were used simultaneously to calculate the amounts of N, P, K, Ca and Mg removed and left in the logging residues for five thinning intensities, five site indexes and four harvesting scenarios for each species, considering the whole rotation. A more intense thinning regime decreases the total amount of nutrients exported and increases the proportion of nutrients returned to the soil before the clearfell, being a more progressive system of extracting nutrients from the ecosystem. A substantial amount of nutrients are located in the crown fractions and the bark, making desirable the harvesting of debarked logs. The results allow the calculation of fertilization needs to avoid the depletion of soil nutrient capital in a variety of silvicultural situations.     

Permanence of resilience and protection efficiency in mountain Norway spruce forest stands: A simulation study

An objective of mountain forest management is to increase the ability of forest stands to protect human activities against natural hazards such as rock-falls and snow avalanches in a sustainable way. The challenge is to find a compromise between efficient instantaneous protection, favoured by dense stands, and continuous renewal, minimizing time periods of low protection efficiency. We used a Norway spruce stand dynamics model to compare the respective advantages of individual tree and gap selection silviculture in this context. We simulated stand dynamics over 800 years with either individual tree or gap thinning every 20 years with several thinning intensities. At each time step, we evaluated stand resilience, protection efficiency against rock-falls, protection efficiency against avalanches, and structural complexity with four indicators based on stand structure. Every scenario produced short time periods with low stand resilience and protection efficiency. Such periods can be tolerated if they are sufficiently rare compared to the local disturbance regime. We characterized the permanence of resilience and protection of a forest stand as its ability to remain within boundary values of the different indicators, without going out of them during continuous time periods longer than fixed maximum durations. Permanence of resilience and permanence of protection decreased with thinning intensity. Efficient protection against rock-falls was obtained with gap thinning of intermediate intensity while protection against avalanches was obtained only for very low thinning intensities. For our ecological context, the best compromise between resilience and protection was obtained with three 10 m radius gaps per hectare every 20 years (9.5% of the area of a stand). This strategy led to uneven-aged stand structures with a high diversity of diameters classes. Our results suggest that small gap silviculture may be a good way to combine forest renewal and protection efficiency in mountain regions.     

Hybrid and clonal variability of nutrient content and nutrient use efficiency in Eucalyptus stands in Congo

For 20 years, there has been 42,000 ha estate of clonal Eucalyptus plantations around Pointe-Noire in Congo on sandy soils that have very low reserves of available nutrients. These plantations have been based on a natural hybrid (E. PF1). This hybrid is being replaced by E. urophylla × E. grandis (UG), a more productive hybrid developed by the breeding program of UR2PI. A study of biogeochemical cycles showed that nutrient removal by harvesting is the main nutrient output in the E. PF1 ecosystem. It is therefore important to quantify the nutrient content (NC) in both hybrids to compare corresponding nutrient removal values.

The work dealt with four UG clones and the most planted clone of E. PF1. Twelve trees per clone were sampled at the logging age (8 years) in a clonal test for UG clones and in a nearby stand for E. PF1. Tables were established to predict, from girth at breast height (C_1.30 m), the biomass and nutrient content of stemwood, bark, dead and living branches, leaves, and were applied to the inventory of the different stands to evaluate corresponding biomass, NC and nutrient use efficiency (NUE) on a per-hectare basis.

Total biomass differed between the two hybrids and among UG clones: 109 t ha⁻¹ for E. PF1 and 108–155 t ha⁻¹ for UG clones. In E. PF1 trees, total NC was globally lower for N, K, and Mg, but greater for P and Ca. In stemwood, nitrogen content was similar for both hybrids. By contrast, in UG clones, NC was much lower for P (−72%) and Ca (−40% to −55%). The same trends were observed for NUE: equivalent for both hybrids for N, but higher in UG clones for P (+72%) and Ca (+43% to +59%). A marked variability among clones was observed for K and Mg. UG clones allocated proportionally more nutrients in leaves than E. PF1.

These results show that clones should not be selected only on growth traits but also on NUE and on the concentration of nutrients in tree components removed by harvesting. It will be then possible to limit the cost of fertilising needed to maintain stand growth and soil fertility.     

Long-term dynamics of leaf and root decomposition and nitrogen release in a grey alder (Alnus incana (L.) Moench) and silver birch (Betula pendula Roth.) stands

The decomposition of the leaf litter, fine roots (d?<?2?mm) and coarser roots (2?≤?d?<?5?mm) of grey alder and silver birch, as well as of α-cellulose sheets using the litterbag method was studied in two experimental stands on Podzoluvisol soils in Southern Estonia. For both tree species, the coarser roots decomposed faster than the fine roots, (p?<?.05), tree species did not affect the decomposition rate of the roots (p?>?.5). The nitrogen (N) input to soil from aboveground litter was multiple times higher than the N flux from roots. The remaining relative ash-free mass of the leaves of grey alder and silver birch after three and a half years was similar. After 11 years the remaining relative ash-free mass of the fine and coarser roots of grey alder still accounted for around 10% of the initial value. For silver birch the remaining value was around 20% after 9 years. The litterbag method to underestimates in fertile soils the decomposition of organic matter and thus did not reflect the actual dynamics of decomposition.     

Biomass and nutrient allocation in Douglas-fir and amabilis fir seedlings: influence of growth rate and nutrition

Allocation of biomass and nutrients to shoots and roots was followed for three years in fast and slow growing populations of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), a fast growing pioneer species, and amabilis fir (Abies amabilis Dougl. ex J. Forbes), a slow growing shade-tolerant species. Seedlings were grown for three seasons in five nutrient treatments containing varying proportions of nitrogen and phosphorus (N:P). In both species, growth was greatest in the 250:20 N:P treatment followed by the 100:60 and 100:20 treatments. Vector analysis showed that, in both species, relative to the 100:20 treatment, seedlings in the 20:20 treatment were N deficient and seedlings in the 100:4 treatment were P deficient, i.e., where deficiency is defined to mean that an increase in nutrient supply increases nutrient content, nutrient concentration and plant dry weight. Seedlings in the 100:60 treatment had a higher P content than seedlings in the 100:20 treatment but the same dry weight, indicative of what Timmer and Armstrong (1987) termed luxury consumption. No nutrient retranslocation was observed in either species until the third growing season. In Douglas-fir, the greatest percentage of nutrients was exported from one-year-old shoots between May and July of the third growing season to support new growth. The total amount and percent of nutrients retranslocated was higher in Douglas-fir than in amabilis fir. Amabilis fir seedlings also exported N and P from older shoots, but this was later partially replenished. In both species, P retranslocation was greatest in treatments with a high N:P ratio. Nitrogen retranslocation was greatest in amabilis fir seedlings in treatments with a low N:P ratio, and greatest in Douglas-fir seedlings in the 250:20 and 100:60 treatments. Potassium retranslocation was correlated with seedling size. Douglas-fir retranslocated more of its shoot N reserves into new growth at the expense of older needles when soil fertility was high and sinks were strong. Otherwise, both species preferentially translocated the elements in short supply. Thus, retranslocation varied with the ecological characteristics of species, the relative availability of soil nutrients and sink strength.