Recovery of Aboveground Plant Biomass and Productivity After Fire in Mesic and Dry Black Spruce Forests of Interior Alaska

Plant biomass accumulation and productivity are important determinants of ecosystem carbon (C) balance during post-fire succession. In boreal black spruce (Picea mariana) forests near Delta Junction, Alaska, we quantified aboveground plant biomass and net primary productivity (ANPP) for 4 years after a 1999 wildfire in a well-drained (dry) site, and also across a dry and a moderately well-drained (mesic) chronosequence of sites that varied in time since fire (2 to ∼116 years). Four years after fire, total biomass at the 1999 burn site had increased exponentially to 160 ± 21 g m⁻² (mean ± 1SE) and vascular ANPP had recovered to 138 ± 32 g m⁻² y⁻¹, which was not different than that of a nearby unburned stand (160 ± 48 g m⁻² y⁻¹) that had similar pre-fire stand structure and understory composition. Production in the young site was dominated by re-sprouting graminoids, whereas production in the unburned site was dominated by black spruce. On the dry and mesic chronosequences, total biomass pools, including overstory and understory vascular and non-vascular plants, and lichens, increased logarithmically (dry) or linearly (mesic) with increasing site age, reaching a maximum of 2469 ± 180 (dry) and 4008 ± 233 g m⁻² (mesic) in mature stands. Biomass differences were primarily due to higher tree density in the mesic sites because mass per tree was similar between sites. ANPP of vascular and non-vascular plants increased linearly over time in the mesic chronosequence to 335 ± 68 g m⁻² y⁻¹ in the mature site, but in the dry chronosequence it peaked at 410 ± 43 g m⁻² y⁻¹ in a 15-year-old stand dominated by deciduous trees and shrubs. Key factors regulating biomass accumulation and production in these ecosystems appear to be the abundance and composition of re-sprouting species early in succession, the abundance of deciduous trees and shrubs in intermediate aged stands, and the density of black spruce across all stand ages. A better understanding of the controls over these factors will help predict how changes in climate and fire regime will affect the carbon balance of Interior Alaska. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Density effects and self-thinning in even-aged pure stands ofEucalyptus camaldulensis Dehn

The competition density effect and changes of mean total tree weight (w) and stand density (ρ) during course of self-thinning were examined in even-aged pure stands ofEucalyptus camaldulensis Dehn. which were planted in the tropical monsoon region. The level of competition was controlled by changing the initial stand density from 625 trees ha⁻¹ to 40,000 trees ha⁻¹. Hozumi's model was used to describe thew-ρ trajectory with aging of each stand and thew-ρ relation between stands of different densities at each time. The higher density produced trees of smaller mean tree sizes. The higher the density, the sooner self-thinning began. The growth curve ofE. camaldulensis followed the logistic growth curve where both maximum size and intrinsic growth rate change with time. Mean intrinsic growth rate was maximized at initiation of growth after lag time and then gradually decreased as time progressed. Hozumi's model was considered to be the best model with wide applicability for describing and comparing the growth characteristics during the course of self-thinning among different species, especially in tropical forest plantations, in which many diverse species were used for reforestation.     

Structure, above-ground biomass and dynamics of mangrove ecosystems: new data from French Guiana

The article presents new results on the structure and the above-ground biomass of the various population types of mangroves in French Guiana. Nine mangrove stands were studied, each composed of three to ten adjoining plots with areas that varied depending on the density of the populations. Structural parameters and indices were calculated. Individuals representative of the three groups of taxa present were felled:Avicennia germinans (L) Stearn, Rhizophora spp., and Laguncularia racemosa (L) Gaertn. The trunks, branches and leaves were sorted and weighed separately. The biomass was obtained by determining the allometric relationships, the general equation selected being of the type y = a _o x ^a1, where the diameter (x) is the predictive variable. The total above-ground biomass varied from 31 t ha⁻¹ for the pioneer stages to 315 t ha⁻¹ for mature coastal mangroves, but with large variations depending on the structural characteristics at each site. The results place the Guianese mangroves among those with high biomass, although lower than those reported for Asia. Based on the relationships between structural parameters and standing biomass, in particular with the use of the “self-thinning rule”, population dynamics models are proposed. Received: 16 August 1996 / Accepted 17 January 1998     

Self-thinning lines of organs and aboveground parts based on allometric relationships in overcrowded Pinus densiflora stands

The allometric relationships of mean tree height [`(H)]( μ [`(w)]_x^q ) \bar{H}( \propto \bar{w}_{\rm {x}}^{\theta } ) and of organ mass density [`(d)]<sub>x</sub> ( μ [`(w)]_x^d ) \bar{d}_{\rm {x}} ( \propto \bar{w}_{\rm {x}}^{\delta } ) to mean organ mass [`(w)]_x \bar{w}_{\rm {x}} were studied in self-thinning Pinus densiflora Sieb. et Zucc. stands. Tree height increased significantly with increasing mean mass of organs and aboveground parts on log–log coordinates. The value of the allometric constant θ ranged from 0.2878 to 0.3349. On the other hand, the constant value δ was not significantly different from zero, except for leaves. The value of the allometric constant δ ranged from −0.2926 to 0.0120. According to Weller’s allometric model, the slope of the self-thinning line was calculated from the allometric constants θ and δ. The thinning slope was estimated to be −1.51 in stem, −1.39 in branches, −1.00 in leaf and −1.41 in aboveground parts, respectively. Mass density was high in stem, medium in branches and low in leaves. Mean leaf mass density decreased significantly with decreasing stand density on log–log coordinates, which could be interpreted as indicating the importance of the constant final leaf biomass in overcrowded P. densiflora stands. The self-thinning exponents of branch, stem and aboveground parts were not significantly different from 3/2, which indicated that the 3/2 power law of self-thinning holds for stem mass, branch mass and aboveground mass in overcrowded P. densiflora stands.     

Linking stand-level self-thinning allometry to the tree-level leaf biomass allometry

Long-term experimental plots of Norway spruce and European beech are investigated for a link between stand-level self-thinning and tree-level leaf biomass allometry. Self-thinning refers to the finding of Reineke (1933), who postulated for unthinned forest stands that with β = −1.605; i.e. an increase of mean (quadratic) diameter d _q by 1% results in a decrease of tree number N by 1.605%. On the individual tree level, leaf biomass (w _L) can be related allometrically to the tree diameter d: w _L = ad ^α. If we assume that (a) the stands have reached the ceiling leaf area, (b) the specific leaf area (leaf area/leaf weight) is constant, and (c) differences resulting from the use of mean quadratic diameter or individual tree diameter are negligible, then the decrease in the stands’ leaf biomass due to the trees lost in self-thinning must be compensated by an equivalent increase in the remaining trees’ leaf biomass. This means, the absolute slope of the individual trees’ leaf biomass allometry α and the self-thinning allometry β would be equal and just have the opposite sign: α = −β. The analysis of the two long-term plots reveals that α is stronger than β, both for spruce (β = −1.744, α = 1.840) and especially for beech (β = −1.791, α = 2.181). The cause is traced back to a changing average specific leaf area during stand development [assumption (b) is wrong]. The results do not only bridge a gap between tree and stand allometry, but also emphasize an important effect for the understanding and modelling of the resource allocations in trees and forests.     

Fire Severity and Long-term Ecosystem Biomass Dynamics in Coniferous Boreal Forests of Eastern Canada

The objective of this study was to characterize the effects of soil burn severity and initial tree composition on long-term forest floor dynamics and ecosystem biomass partitioning within the Picea mariana [Mill.] BSP-feathermoss bioclimatic domain of northwestern Quebec. Changes in forest floor organic matter and ecosystem biomass partitioning were evaluated along a 2,355-year chronosequence of extant stands. Dendroecological and paleoecological methods were used to determine the time since the last fire, the soil burn severity of the last fire (high vs. low severity), and the post-fire tree composition of each stand (P. mariana vs. Pinus banksiana Lamb). In this paper, soil burn severity refers to the thickness of the organic matter layer accumulated above the mineral soil that was not burned by the last fire. In stands originating from high severity fires, the post-fire dominance by Pinus banksiana or P. mariana had little effect on the change in forest floor thickness and tree biomass. In contrast, stands established after low severity fires accumulated during the first century after fire 73% thicker forest floors and produced 50% less tree biomass than stands established after high severity fires. Standing tree biomass increased until approximately 100 years after high severity fires, and then decreased at a logarithmic rate in the millennial absence of fire. Forest floor thickness also showed a rapid initial accumulation rate, and continued to increase in the millennial absence of fire at a much slower rate. However, because forest floor density increased through time, the overall rate of increase in forest floor biomass (58 g m⁻² y⁻¹) remained constant for numerous centuries after fire (700 years). Although young stands (< 200 years) have more than 60% of ecosystem biomass locked-up in living biomass, older stands (> 200 years) sequester the majority (> 80%) of it in their forest floor. The results from this study illustrate that, under similar edaphic conditions, a single gradient related to time since disturbance is insufficient to account for the full spectrum of ecosystem biomass dynamics occurring in eastern boreal forests and highlights the importance of considering soil burn severity. Although fire severity induces diverging ecosystem biomass dynamics in the short term, the extended absence of fire brings about a convergence in terms of ecosystem biomass accumulation and partitioning.     

The relationship between fire history and an exotic fungal disease in a deciduous forest

Exotic diseases have fundamentally altered the structure and function of forest ecosystems. Controlling exotic diseases across large expanses of forest has proven difficult, but fire may reduce the levels of diseases that are sensitive to environmental conditions. We examined Cornus florida populations in burned and unburned Quercus–Carya stands to determine if burning prior to anthracnose infection has reduced the impacts of an exotic fungal disease, dogwood anthracnose, caused by Discula destructiva. We hypothesized that fire has altered stand structure and created open conditions less conducive to dogwood anthracnose. We compared C. florida density, C. florida health, and species composition and density among four sampling categories: unburned stands, and stands that had burned once, twice, and 3 times over a 20-year period (late 1960s to late 1980s). Double burn stands contained the greatest density of C. florida stems (770 stems ha⁻¹) followed by triple burn stands (233 stems ha⁻¹), single burn stands (225 stems ha⁻¹) and unburned stands (70 stems ha⁻¹; P < 0.01). We observed less crown dieback in small C. florida trees (<5 cm diameter at breast height) in burned stands than in unburned stands (P < 0.05). Indicator species analysis showed that burning favored species historically associated with Quercus–Carya forests and excluded species associated with secondary succession following nearly a century of fire suppression. Our results suggest that fire may mitigate the decline of C. florida populations under attack by an exotic pathogen by altering forest structure and composition. Further, our results suggest that the burns we sampled have had an overall restorative effect on forest communities and were within the fire return interval of the historic fire regime. Consequently, prescribed fire may offer a management tool to reduce the impacts of fungal disease in forest ecosystems that developed under historic burning regimes.     

Density effects on the growth of self-thinning Eucalyptus urophylla stands

Density effects on the growth of self-thinning Eucalyptus urophylla stands were examined for 7 years. Tree height and stem diameter at breast height were measured during the experimental period. Stems, branches, leaves, bark and roots of 45 E. urophylla trees were sampled in three different density stands in order to establish their biomass equations. Change trends of the biological time τ and density ρ were described used corresponding equations. The stem weight ratio increased and leaf weight ratio decreased, whereas those of branch, bark and root were relatively steady from 2 years after the planting. The competition-density (C-D) effect equation of mean organ weight w _o was derived by combining the allometric power relationship between mean tree weight w and w _o with the C-D effect equation of self-thinning stands. The equations of the C-D effect for w and ρ and for w _o and ρ were used to describe the C-D effects in tree and organs during course of self-thinning, respectively, and showed a good fit to the data. Leaf biomass of different density stands reached a more or less constant level with time elapse. High density produced the greatest biomass and stem biomass, so that it is the best choice in silvicultural practice.     

Stand and self-thinning dynamics in natural <Emphasis Type="Italic">Abies</Emphasis> stands in northern Hokkaido,Japan

Stand dynamics and self-thinning were analyzed in relation to the dynamics of above-ground biomass in natural Abies sachalinensis stands growing on sand dunes in northern Hokkaido, Japan. This was done in order to examine wave-type regeneration in the stands. Fifty-two plots were established in almost pure Abies stands that ranged from saplings to the mature and collapsing growth stages. Above-ground biomass and tree height reached asymptotic levels prior to the collapsing phase, unlike wave-regeneration Abies stands in central Japan and North America. Stand density was high in the young growth stages, but the self-thinning rate, that is, the density decrease per biomass growth in the study stands was greater than in wave-regeneration stands in central Japan, as indicated by a large self-thinning exponent (–1.26 by reduced major axis regression). The range of tree height distribution was very narrow, and the stands vertical structure was typically single-layered. The slenderness ratio of trees was large, except in young stands. In mature and collapsing stands, advanced seedling density increased markedly. These stand and tree characteristics were considered to be correlated with the wave-type regeneration in the study stands, and it is assumed that prevailing winds affect tree mortality.     

Ectomycorrhizal root growth in scots pine stands in response to manipulation of litter and humus layers

 The effect on ectomycorrhizal root growth in a nitrogen-enriched planted stand of Scots pine (Pinus sylvestris L.) on podzolic sandy soil to manipulation of litter and humus layers (removal, doubling and control treatments) was examined, and compared to ectomycorrhizal root growth in an untreated naturally established Scots pine stand on nutrient-poor non-podzolic sandy soil. Half a year after manipulation of litter and humus layers in the planted stand, ingrowth-cores to a depth of 60 cm were installed in both stands. Scots pine roots were sampled four times during two growing seasons. Ectomycorrhizal roots were found at all sampled soil depths to 60 cm in all plots. Root growth and ectomycorrhizal development were greater in the naturally established stand than in all plots in the planted stand. Numbers of ectomycorrhizal root tips in the litter and humus removal plots were generally higher than in the control plots in the planted stand until May 1992. Doubling litter and humus did not significantly affect root length or the numbers of ectomycorrhizal root tips. The N _dissolved , NH₄ ⁺ and NO₃ ^– concentrations and the organic matter content in the upper 5 cm of the mineral soil in the planted stand on podzolic sandy soil were generally higher and the pH significantly lower than in the naturally established stand on non-podzolic sandy soil. Root growth and ectomycorrhizal development in the secondary stand may have been negatively affected by the chemical composition of the podzolic sandy soil. Accepted: 19 March 1997     

Density effect, self-thinning and size distribution in Pinus densiflora Sieb. et Zucc. stands

The competition-density (C-D) effect for given times and self-thinning over time in even-aged, natural, pure stands of Pinus densiflora Sieb. et Zucc. were analyzed with the reciprocal equation of the C-D effect in self-thinning stands, and the equation describing the time-trajectory of mean stem volume and stand density. The C-D effect and self-thinning were consistently well explained by the two equations. Differences in mean stem volume and in stand density among the stands tended to merge with increasing stand age. The self-thinning line with a slope of approximately –3/2 was reached by the higher density stand prior to the medium and lower density stands. The skewness of tree height distribution showed positive values, which means that the distribution is more or less L-shaped, and in addition the skewness decreased with increasing mean tree height, which indicates that smaller trees died as the stands grew. This trend is consistent with the asymmetric (one-sided) competition hypothesis that self-thinning is driven by competition for light. The tree height distribution was analyzed using the Weibull distribution. The location parameter h _min of the Weibull distribution increased with increasing stand age, and the scale parameter a tended to increase slightly with increasing stand age. The range of the shape parameter b of the Weibull distribution corresponded to that of the skewness.     

Natural 15N abundance in two nitrogen saturated forest ecosystems

Natural ¹⁵N abundance values were measured in needles, twigs, wood, soil, bulk precipitation, throughfall and soil water in a Douglas fir (Pseudotsuga menziesii (Mirb.) and a Scots pine (Pinus sylvestris L.) stand receiving high loads of nitrogen in throughfall (>50 kg N ha⁻¹ year⁻¹). In the Douglas fir stand δ¹⁵N values of the vegetation ranged between −5.7 and −4.2‰ with little variation between different compartments. The vegetation of the Scots pine stand was less depleted in ¹⁵N and varied from −3.3 to −1.2‰δ¹⁵N. At both sites δ¹⁵N values increased with soil depth, from −5.7‰ and −1.2‰ in the organic layer to +4.1‰ and +4.7‰ at 70 cm soil depth in the Douglas fir and Scots pine stand, respectively. The δ¹⁵N values of inorganic nitrogen in bulk precipitation showed a seasonal variation with a mean in NH₄ ⁺-N of −0.6‰ at the Douglas fir stand and +10.8‰ at the Scots pine stand. In soil water below the organic layer NH₄ ⁺-N was enriched and NO₃ ⁻-N depleted in ¹⁵N, which was interpreted as being caused by isotope fractionation accompanying high nitrification rates in the organic layers. Mean δ¹⁵N values of NH₄ ⁺ and NO₃ ⁻ were very similar in the drainage water at 90 cm soil depth at both sites (−7.1 to −3.8‰). A dynamic N cycling model was used to test the sensitivity of the natural abundance values for the amount of N deposition, the ¹⁵N ratio of atmospheric N deposited and for the intrinsic isotope discrimination factors associated with N transformation processes. Simulated δ¹⁵N values for the N saturated ecosystems appeared particularly sensitive to the ¹⁵N ratio of atmospheric N inputs and discrimination factors during nitrification and mineralization. The N-saturated coniferous forest ecosystems studied were not characterized by elevated natural ¹⁵N abundance values. The results indicated that the natural ¹⁵N abundance values can only be used as indicators for the stage of nitrogen saturation of an ecosystem if the δ¹⁵N values of the deposited N and isotope fractionation factors are taken into consideration. Combining dynamic isotope models and natural ¹⁵N abundance values seems a promising technique for interpreting natural ¹⁵N abundance values found in these forest ecosystems. Received: 5 May 1996 / Accepted: 10 April 1997     

Comparison of Nitrogen Fixation for North- and South-facing <Emphasis Type="Italic">Robinia pseudoacacia</Emphasis> Stands in Central Korea

The nitrogenase activity, root nodule biomass, and rates of nitrogen (N) fixation were measured in 25-year-old pure north- and south-facing Robinia pseudoacacia stands in an urban forest of Seoul (Kkachisan Mountain) in central Korea. The nitrogenase activity was estimated using an acetylene reduction (AR) assay, which showed an increasing trend during the early growing season, with sustained high rates from June through to September with a decrease thereafter. July had the highest nitrogenase activity rate (micromoles C₂H₄ per gram dry nodule per hour), averaging 95.8 and 115.1 for the north- and south-facing stands, respectively. The maximum root nodule biomass (kilograms per hectare) was 45.7 and 9.1 for the north- and south-facing stands in July, respectively. The AR rate appeared to be strongly correlated to the soil temperature (r ² = 0.68, P < 0.001) and soil pH (r ² = 0.59, P < 0.001) while root nodule biomass was correlated to the soil temperature (r ² = 0.36, P < 0.01) and water content (r ² = 0.35, P < 0.05). The soil temperature showed clear differences between seasons, while there was a significant difference in soil pH, organic matter, total N concentrations, and available phosphorus between the north- and south-facing stands. The N₂ fixation rates during the growing season varied from 0.1 to 37.5 kg N ha⁻¹ month⁻¹ depending on the sampling location and time. The annual N₂ fixation rate (kg N per hectare per year) was 112.3 and 23.2 for the north- and south-facing stands, respectively. The differences in N₂ fixation rate between the two stands were due mainly to the differences in total nodule biomass.     

Rainfall partitioning and related hydrochemical fluxes in a diverse and in a mono specific (Phenakospermum guyannense) secondary vegetation stand in eastern Amazonia

Rainfall partitioning into throughfall and stemflow was studied in a diverse and in a mono specific stand of secondary vegetation in Eastern Amazonia. The nutrient concentrations in the water were analysed in order to quantify the related hydrochemical fluxes. Secondary vegetation forms the fallow in the local shifting cultivation system and is usually dominated by shrubs and trees. Phenakospermum guyannense (Strelitziaceae), a banana-like herb, is one of the predominant non-woody species. The study was conducted during an 18-month period in a 2.5-year-old relatively species-rich stand and a 10-year-old stand dominated by P. guyannense. In a year with 1956 mm of rainfall 65% (1281 mm) of this quantity reached the soil as throughfall in the diverse stand and 38% (743 mm) in the mono specific stand. Stemflow was estimated to be 23% and 41% respectively. P. guyannense and Banara guianensis (Flacourtiaceae), a tree species, were causing these high funnelling effects. In the young diverse stand B. guianensis had a stemflow of more than 200 l year⁻¹ and P. guyannense had a median flux of 77 l year⁻¹ per pseudostem. In the older stand the taller plants of P.␣guyannense collected 644 l year⁻¹ per pseudostem on the median. The reason for these high values could be the banana-like growth form of P. guyannense and the crown morphology of B. guianensis, which has inclined branches. The low proportion of throughfall and the high stemflow values differ from all previous studies in Amazonian primary forests. The proximity to the Atlantic Ocean strongly influenced the nutrient fluxes via rainfall at our study site. This becomes obvious from the high Na and Cl fluxes with rainfall (19.7 kg Na ha⁻¹ year⁻¹, 37.2 kg Cl ha⁻¹ year⁻¹) which were approximately equal to the Na and Cl fluxes with the sum of throughfall and stemflow for both stands. K fluxes in throughfall and stemflow in both stands were higher than in rainfall by a factor of 8. The high K enrichment during the crown passage is assumed to be caused by a␣high K concentration in the leaf tissue resulting in enhanced leaching from the leaves. In months with low␣rainfall the concentrations of Ca, Mg, S and Cl in throughfall of the diverse stand were significantly higher than in months with high rainfall. This was mainly due to vegetation burns in the dry period, which resulted in ash deposition on the canopy and subsequent wash-off and solution of ash particles. Received: 11 May 1997 / Accepted: 2 November 1997     

Leaf biomass changes with stand development in hinoki cypress (<Emphasis Type="Italic">Chamaecyparis obtusa</Emphasis> [Sieb. et Zucc.] Endl.)

We monitored a permanent plot of 3-year-old Chamaecyparis obtusa seedlings for 11 years after planting. As the stem cross-sectional area at the crown base can be regarded as a good predictor of leaf mass according to the pipe model theory, we measured this parameter to determine temporal trends in leaf biomass. The mean values showed asymptotic growth, maintaining a near-constant level after a stand age of 9 years. Peak values were found at 9 years, followed by a slight decrease because of a continuous reduction in stand density. This temporal trend suggests that the leaf biomass per unit land area attains a peak at an age of 9 years. As the stand density changes with stand age, the relationship between stand stem cross-sectional area at the crown base and stand density showed an optimum curve in which the optimum density was around 9200 ha⁻¹. We propose hypothetical trends in primary productivity and biomass density with stand age, based on the results of measurements of stem cross-sectional area at the crown base and stand density under the assumption of the 3/2 power law of self-thinning.     

Biomass Expansion Factors of Natural Japanese Red Pine (<Emphasis Type="Italic">Pinus densiflora</Emphasis>) Forests in Korea

Biomass expansion factors, which convert the timber volume (or dry weight) to biomass, are used to estimate the forest biomass and account for the carbon budget at the national or regional level. This study estimated the biomass conversion and expansion factors (BCEF), root to shoot ratio (R), biomass expansion factors (BEF) of natural Japanese Red Pine (Pinus densiflora Sieb. et Zucc.) forests based on direct field measurements and publications in Korea. This study attempted to fit the non-linear relationships between the biomass expansion factors (BCEF and BEF) and main stand factors [stand age, tree height, and diameter at breast height (DBH)]. The relationship between BEF and each main stand factor was expressed as a simple logarithmical equation. The BCEF was also expressed as a logarithmical equation of the tree height, DBH, and stand volume, whereas there was no significant relationship between BCEF and stand age. The mean value for BCEF, BEF, and R was 0.5821 Mg m⁻³ (n = 22, SD = 0.1196), 1.4465 (n = 22, SD = 0.2905), and 0.2220 (n = 17, SD = 0.0687), respectively. The values of the biomass expansion factors in this study may indicate much representativeness to estimate forest biomass in natural Japanese Red Pine forests of Korea than the default values given by the IPCC (2003, 2006).     

Biomass and net production in a bambooPhyllostachys bambusoides stand

Biomass and net production were measured in aPhyllostachys bambusoides stand in Kyoto Prefecture, central Japan, which had carried out gregarious flowering in 1969 and has been recovering vegetatively. The culm density fluctuated around an average value of 12 040 ha⁻¹ during the research period (1985–91). Annual recruirment and mortality rates of culms were 1340 and 1133 ha⁻¹, respectively. The mean diameter at breast height increased from 7.28 cm in 1985 to 8.68 cm in 1991, and the biomass of culms increased from 71.3 to 111.6t ha⁻¹ over the same time period. Branch and leaf biomasses were almost constant, 10.0 and 9.4t ha⁻¹ on average, respectively. The leaf area index of the stand was 11.6 ha ha⁻¹, which is one of the largest values found in Japanese forests. The belowground biomass of 32.6t ha⁻¹ for rhizomes and 14.8t ha⁻¹ for fine roots resulted in the smaller ratio of aboveground parts to the root system (2.38) than those determined for forest stands. The amount of litterfall, excluding culms and large branches, was large (9.13t ha⁻¹ year⁻¹), corresponding to those measured in equatorial stands. The aboveground net production was 24.6t ha⁻¹ year⁻¹, larger than the average value reported for forest stands under similar weather conditions.     

Litterfall and fine root biomass contribution to nutrient dynamics in second- and old-growth Douglas-fir ecosystems

Litterfall and fine root production were measured for three years as part of a carbon balance study of three forest stands in the Pacific Northwest of the United States. A young second-growth Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] stand, a second-growth Douglas-fir with red alder (Alnus rubra Bong.) stand, and an old-growth (∼550 years) Douglas-fir stand were monitored for inputs of carbon and nitrogen into the soil from litterfall and fine root production, as well as changes in soil C and N. Fine root production and soil nutrient changes were measured through the use of soil ingrowth bags containing homogenized soil from the respective stands. Litterfall biomass was greatest in the Douglas-fir-alder stand (527 g m⁻² yr⁻¹) that annually returned nearly three times the amount of N as the other stands. Mean residence time for forest floor material was also shortest at this site averaging 4.6 years and 5.5 years for C an N, respectively. Fine root production in the upper 20 cm ranged from 584 g m⁻² in the N rich Douglas-fir-alder stand to 836 g m⁻² in the old-growth stand. Fine root production (down to one meter) was always greater than litterfall with a below:above ratio ranging from 3.73 for the young Douglas-fir stand to 1.62 for the Douglas-fir-alder stand. The below:above N ratios for all three stands closely approximate those for biomass. Soil changes in both C and N differed by site, but the soil C changes in the old-growth stand mirrored those obtained in an ongoing CO₂ flux study. Results from the soil ingrowth bags strongly suggest that this method provides a simple, but sufficient device for measuring potential fine root biomass production as well as soil chemical changes.     

The difference between above- and below-ground self-thinning lines in forest communities

Quantifying the self-thinning process in various plant communities has been a long-standing issue in both theoretical and empirical studies. Most studies on plant self-thinning have centered only on aboveground parts, and rarely on belowground parts. There is still a general lack of comparison between above- and belowground self-thinning processes, especially for forest communities. The fundamental mechanistic difference and the functional association between above- and belowground competition indicate that the self-thinning process of belowground parts may be different from that of aboveground parts. We investigated the self-thinning lines for above-ground (M _A), below-ground (M _B), and total biomass (M _T), respectively, across forest communities in China. The results showed that neither the classical self-thinning rule (−3/2 exponent) nor the universal scaling rule (−4/3 exponent) can apply to all the self-thinning relationships across these forest communities and that the self-thinning lines for belowground biomass were flatter and lower than those for aboveground biomass across most of these forest communities.     

Density effects on organs in self-thinning <Emphasis Type="Italic">Pinus densiflora</Emphasis> Sieb. et Zucc. stands

Although much research on the density effect in nonself-thinning populations has been conducted, there has been very little research on density effects in self-thinning populations. Furthermore, the density effect of plant organs in self-thinning populations is little reported. The present study analyzed the yield–density (Y–D) effects on organs, such as stem, branch and leaf, together with that on stands of self-thinning Pinus densiflora Sieb. et Zucc.. The stand yield- and organ Y–D effects were well described by reciprocal and parabolic equations, respectively, throughout the experiment. The value of coefficient B in the reciprocal equation decreased monotonically with increasing stand age and became significantly closer to zero at the end of experiment (33-year-old stand), indicating that the constant final stand yield was established regardless of the density realized. The value of the relative growth coefficient h in the allometric equation between mean organ weight and mean aboveground weight was significantly smaller than 1.0 for stem, indicating that stem yield increases monotonically with increasing realized density. The h-value was significantly larger than 1.0 for branch throughout the experiment, and for leaf except at 33 years old, indicating that optimum densities exist. The h-value for leaf was not significantly different from 1.0 at 33 years old, indicating that the leaf yield reached a constant level regardless of realized density. The constant final leaf yield was established at almost the same growth stage as the establishment of constant final stand yield.