Above- and belowground biomass and primary productivity of a Larix gmelinii stand near Tura, central Siberia

We assessed above- and belowground biomass and net primary production (NPP) of a mature Larix gmelinii (Rupr.) Rupr. forest (240-280 years old) established on permafrost soils in central Siberia. Specifically, we investigated annual carbon budgets in roots in relation to root system development and availability of soil resources. Total stand biomass estimated by allometry was about 39 Mg per ha. Root biomass (17 Mg per ha) comprised about 43% of total biomass. Coarse root (>/= 5 mm in diameter) biomass was about twice that of fine roots (< 5 mm). The aboveground biomass/root biomass ratio (T/R) of the larch stand was about unity, which is much less than that of other boreal and subalpine conifer forests. The proportion of fine roots in total root biomass (35%) was relatively high compared with other cold-climate evergreen conifer forests. Total NPP, defined as the sum of annual biomass increment of woody parts and needle biomass, was estimated to be 1.8 Mg per ha per year. Allocation of total NPP to needle production was 56%. The proportion of total NPP in belowground production (27%) was less than for evergreen taiga forests. However, belowground NPP was probably under-estimated because root mortality was excluded. We conclude that L. gmelinii trees invested annual carbon gains largely into needle production or roots, or both, at the expense of growth of aboveground woody parts. This carbon allocation pattern, which resulted in the construction of exploitative root networks, appeared to be a positive growth response to the nutrient-poor permafrost soil of central Siberia.     

Contrasting net primary productivity and carbon distribution between neighboring stands of Quercus robur and Pinus sylvestris

Standing biomass, net primary production (NPP) and soil carbon (C) pools were studied in a 67-year-old pedunculate oak (Quercus robur L.) stand and a neighboring 74-year- old Scots pine (Pinus sylvestris L.) stand in the Belgian Campine region. Despite a 14% lower tree density and a lower tree height in the oak stand, standing biomass was slightly higher than in the pine stand (177 and 169 Mg ha(-1) in oaks and pines, respectively), indicating that individual oak trees contained more biomass than pine trees of similar diameter. Moreover, NPP in the oak stand was more than double that in the pine stand (17.7 and 8.1 Mg ha(-1) year(-1), respectively). Several observations indicated that soil organic matter accumulated at higher rates under pines than under oaks. We therefore hypothesized that the pines were exhibiting an age-related decline in productivity due to nutrient limitation. The poor decomposability of pine litter resulted in the observed accumulation of organic matter. The subsequent immobilization of nutrients in the organic matter, combined with the already nutrient-poor soil conditions, resulted in a decrease in total NPP over time, as well as in a substantial shift in the allocation of NPP toward fine roots. In the oak stand, litter is less recalcitrant to decay and soil acidity is less severe; hence, organic matter does not accumulate and nutrients are recycled. This probably explains why NPP was much higher in the oaks than in the pines and why only a small proportion of NPP was allocated to oak fine roots.     

Fine root biomass and production in Scots pine stands in relation to stand age

We determined fine root biomass and production of 15-, 35- and 100-year-old Scots pine (Pinus sylvestris L.) stands during three growing seasons. Fine roots were sampled by the soil core method. Mean (+/- SE) annual fine root biomass of Scots pine in the 15-, 35- and 100-year-old stands was 220 +/- 25, 357 +/- 21 and 259 +/- 26 g m(-2), respectively. Fine root biomass of the understory vegetation was 159 +/- 54 g m(-2), 244 +/- 30 and 408 +/- 81 g m(-2), and fine root necromass was 500 +/- 112, 1,047 +/- 452 and 1,895 +/- 607 g m(-2) in the sapling, pole stage and mature stands, respectively. Both understory and Scots pine fine root production increased with stand age. Mean annual Scots pine fine root production was 165 +/- 131, 775 +/- 339 and 860 +/- 348 g m(-2) year(-1) in the sapling, pole stage and mature stand, respectively. The respective mean annual production of all fine roots (Scots pine and understory) was 181 +/- 129, 1,039 +/- 497 and 1,360 +/- 869 g m(-2) year(-1). The Scots pine and understory fine root biomass, necromass and production varied in relation to stand age, although the variation was not statistically significant.     

Equations for estimating above- and belowground biomass of Norway spruce,Scots pine,birch spp. and European aspen in Latvia

This study aims to derive allometric functions to estimate the above- and belowground biomass components of the most important tree species in Latvia. The study material included a total of 81 Norway spruce (Picea abies [L.] Karst), 102 Scots pine (Pinus sylvestris L.), 105 birch spp. (mainly silver birch (Betula pendula Roth)) and 84 European aspen (Populus tremula L.) trees sampled in 124 forest stands. The suitability of three mathematical models for the prediction of total aboveground biomass, stem biomass, total live and dead branch biomass, belowground biomass and small root biomass was evaluated. Our analysis revealed that the use of the Intergovernmental Panel on Climate Change mean default values for the root-to-shoot ratio recommended for temperate and boreal ecological zones leads to the overestimation of root biomass of young trees, especially Scots pine and Norway spruce. Our findings indicate that biomass functions recommended for other Baltic Sea countries are not appropriate for the assessment of the biomass stock in Latvia’s forests because these lead to biased estimates. The biomass functions derived in our study are recommended for reporting the biomass stock in Latvia.     

Fine root biomass in relation to site and stand characteristics in Norway spruce and Scots pine stands

Variations in fine root biomass of trees and understory in 16 stands throughout Finland were examined and relationships to site and stand characteristics determined. Norway spruce fine root biomass varied between 184 and 370 g m(-2), and that of Scots pine ranged between 149 and 386 g m(-2). In northern Finland, understory roots and rhizomes (< 2 mm diameter) accounted for up to 50% of the stand total fine root biomass. Therefore, the fine root biomass of trees plus understory was larger in northern Finland in stands of both tree species, resulting in a negative relationship between fine root biomass and the temperature sum and a positive relationship between fine root biomass and the carbon:nitrogen ratio of the soil organic layer. The foliage:fine root ratio varied between 2.1 and 6.4 for Norway spruce and between 0.8 and 2.2 for Scots pine. The ratio decreased for both Norway spruce and Scots pine from south to north, as well as from fertile to more infertile site types. The foliage:fine root ratio of Norway spruce was related to basal area and stem surface area. The strong positive correlations of these three parameters with fine root nitrogen concentration implies that more fine roots are needed to maintain a certain amount of foliage when nutrient availability is low. No significant relationships were found between stand parameters and fine root biomass at the stand level, but the relationships considerably improved when both fine root biomass and stand parameters were calculated for the mean tree in the stand. When the northern and southern sites were analyzed separately, fine root biomass per tree of both species was significantly correlated with basal area and stem surface area per tree. Basal area, stem surface area and stand density can be estimated accurately and easily. Thus, our results may have value in predicting fine root biomass at the tree and stand level in boreal Norway spruce and Scots pine forests.     

Estimating net primary productivity of Chinese pine forests based on forest inventory data

Forest inventory data (FID) include forest resources informationat large spatial scale and long temporal scale. They are importantdata sources for estimating forest net primary productivity(NPP) and carbon budget at landscape and regional scales. Inthis study, more than 100 datasets of biomass, volume, NPP andstand age for Chinese pine forests (Pinus tabulaeformis) fromthe literature were synthesized to develop regression equationsbetween biomass and volume, and between NPP and biomass as wellas stand age. Using these regression equations and the fourthFID surveyed by the Forestry Ministry China from 1989 to 1993,NPP values of Chinese pine forests were estimated. The meanNPP of Chinese pine forests was 4.35 Mg ha^–1 yr^–1.NPP varied widely among provinces, ranging from 1.5 (Neimenggu)to 13.73 Mg ha^–1 yr^–1 (Guizhou). Total NPP of Chinesepine was 10.87 Tg yr^–1 (1 Tg = 10¹² g). NPP values ofChinese pine forests were not distributed evenly across differentprovinces in China. This study may be useful not only for estimatingforest carbon of other forest types but also for evaluatingterrestrial carbon balance at regional and global levels.     

Foliage, fine-root, woody-tissue and stand respiration in Pinus radiata in relation to nitrogen status

We measured respiration of 20-year-old Pinus radiata D. Don trees growing in control (C), irrigated (I), and irrigated + fertilized (IL) stands in the Biology of Forest Growth experimental plantation near Canberra, Australia. Respiration was measured on fully expanded foliage, live branches, boles, and fine and coarse roots to determine the relationship between CO(2) efflux, tissue temperature, and biomass or nitrogen (N) content of individual tissues. Efflux of CO(2) from foliage (dark respiration at night) and fine roots was linearly related to biomass and N content, but N was a better predictor of CO(2) efflux than biomass. Respiration (assumed to be maintenance) per unit N at 15 degrees C and a CO(2) concentration of 400 micro mol mol(-1) was 1.71 micro mol s(-1) mol(-1) N for foliage and 11.2 micro mol s(-1) mol(-1) N for fine roots. Efflux of CO(2) from stems, coarse roots and branches was linearly related to sapwood volume (stems) or total volume (branches + coarse roots) and growth, with rates for maintenance respiration at 15 degrees C ranging from 18 to 104 micro mol m(-3) s(-1). Among woody components, branches in the upper canopy and small diameter coarse roots had the highest respiration rates. Stem maintenance respiration per unit sapwood volume did not differ among treatments. Annual C flux was estimated by summing (1) dry matter production and respiration of aboveground components, (2) annual soil CO(2) efflux minus aboveground litterfall, and (3) the annual increment in coarse root biomass. Annual C flux was 24.4, 25.3 and 34.4 Mg ha(-1) year(-1) for the C, I and IL treatments, respectively. Total belowground C allocation, estimated as the sum of (2) and (3) above, was equal to the sum of root respiration and estimated root production in the IL treatment, whereas in the nutrient-limited C and I treatments, total belowground C allocation was greater than the sum of root respiration and estimated root production, suggesting higher fine root turnover or increased allocation to mycorrhizae and root exudation. Carbon use efficiency, the ratio of net primary production to assimilation, was similar among treatments for aboveground tissues (0.43-0.50). Therefore, the proportion of assimilation used for construction and maintenance respiration on an annual basis was also similar among treatments.     

Forest biomass and root–shoot allocation in northeast China

Temperate and boreal forests act as major sinks for atmospheric CO₂. To assess the magnitude and distribution of the sinks more precisely, an accurate estimation of forest biomass is required. However, the determinants of large-scale biomass pattern (especially root biomass) are still poorly understood for these forests in China. In this study, we used 515 field measurements of biomass across the northeast part of China, to examine factors affecting large-scale biomass pattern and root–shoot biomass allocation. Our results showed that, Picea & Abies forest and coniferous & broadleaf mixed forest had the highest mean biomass (178–202 Mg/ha), while Pinus sylvestris forest the lowest (78 Mg/ha). The root:shoot (R/S) biomass ratio ranged between 0.09 and 0.67 in northeast China, with an average of 0.27. Forest origin (primary/secondary/planted forest) explained 31–37% of variation in biomass (total, shoot and root), while climate explained only 8–15%, reflecting the strong effect of disturbance on forest biomass. Compared with shoot biomass, root biomass was less limited by precipitation as a result of biomass allocation change. R/S ratio was negatively related to water availability, shoot biomass, stand age, height and volume, suggesting significant effects of climate and ontogeny on biomass allocation. Root–shoot biomass relationships also differed significantly between natural and planted forests, and between broadleaf and coniferous forests. Shoot biomass, climate and forest origin were the most important predictors for root biomass, and together explained 83% of the variation. This model provided a better way for estimating root biomass than the R/S ratio method, which predicted root biomass with a R² of 0.71.     

Biomass and biomass change in lodgepole pine stands in Alberta

We describe methods and results for broad-scale estimation and mapping of forest biomass for the Canadian province of Alberta. Differences over successive decades provided an estimate of biomass change. Over 1500 permanent sample plots (PSP) were analyzed from across the range of lodgepole pine (Pinus contorta var. latifolia Engelm.), the major forest tree species of Alberta. The PSP network is densest in stands aged between 70 and 100 years and is well-represented by stands of all ages to 150 years of age. Stand biomass (Mg ha(-1)) was estimated for each PSP plot as the sum of the respective biomass components for each tree (live and standing dead). The biomass components for live trees were stem, bark, branches, foliage and roots. The components for standing dead trees excluded foliage. Equations from previous biomass studies were used for biomass component estimation. Biomass estimates of additional non-tree components were attempted, but without much success. Biomass of the soil organic layer was estimated once on 452 PSPs and a mean estimate of total dead fuels on the ground (28.4 Mg ha(-1)) was available only for the entire distribution of lodgepole pine. However, values of these two components were essentially constant over time and therefore did not alter the analysis or conclusions obtained by analyzing total tree biomass alone. We then used this spatial network of 1549 plots as the basis for mapping biomass across Alberta. Mapping methods were based on Australian National University SPLINe (ANUSPLIN) software, Hutchinson's thin-plate smoothing spline in four dimensions (latitude, longitude, elevation and biomass). Total tree biomass (mean = 172 Mg ha(-1)) was dominated by stem biomass (mean = 106 Mg ha(-1)), which was an order of magnitude greater than the mean estimates for the bark (11 Mg ha(-1)), branch (12 Mg ha(-1)) and foliage (12 Mg ha(-1)) components. A close relationship was found between total tree biomass and stand stem volume (R(2) = 0.992 with n = 3585; note that volume and biomass were calculated independently). We compared total tree biomass for two decades, the 1980s and the 1990s. After correcting for changes in harvest removals over time, the mean change in total biomass was positive (0.99 Mg ha(-1) year(-1)) and differed significantly from zero (n = 421; P < 0.001). Estimates ranged from -13.9 to 8.0 Mg ha(-1) year(-1). The heart of the lodgepole pine distribution (primarily the Foothills subregions) showed an increase in biomass, whereas isolated pockets of lodgepole pine in the boreal northern subregion indicated a decline in biomass.     

Fine root morphological adaptations in Scots pine, Norway spruce and silver birch along a latitudinal gradient in boreal forests

Variability in short root morphology of the three main tree species of Europe's boreal forest (Norway spruce (Picea abies L. Karst.), Scots pine (Pinus sylvestris L.) and silver birch (Betula pendula Roth)) was investigated in four stands along a latitudinal gradient from northern Finland to southern Estonia. Silver birch and Scots pine were present in three stands and Norway spruce was present in all stands. For three fertile Norway spruce stands, fine root biomass and number of root tips per stand area or unit basal area were assessed from north to south. Principal component analysis indicated that short root morphology was significantly affected by tree species and site, which together explained 34.7% of the total variability. The range of variation in mean specific root area (SRA) was 51-74, 60-70 and 84-124 m(2) kg(-1) for Norway spruce, Scots pine and silver birch, respectively, and the corresponding ranges for specific root length were 37-47, 40-48 and 87-97 m g(-1). The range of variation in root tissue density of Norway spruce, Scots pine and silver birch was 113-182, 127-158 and 81-156 kg m(-3), respectively. Sensitivity of short root morphology to site conditions decreased in the order: Norway spruce > silver birch > Scots pine. Short root SRA increased with site fertility in all species. In Norway spruce, fine root biomass and number of root tips per m(2) decreased from north to south. The differences in morphological parameters among sites were significant but smaller than the site differences in fine root biomass and number of root tips.     

Dependence of the aboveground respiration of hinoki cypress (Chamaecyparis obtusa) on tree size

Nighttime respiration was measured at monthly intervals over one year on the aboveground parts of five sample trees in an 8-year-old hinoki cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) stand, by an enclosed standing-tree method. The respiration rate rose rapidly from early spring to a maximum in June, and decreased abruptly in July and then gradually toward autumn and winter. The seasonal change in the respiration rate was synchronized with stem volume increment rather than with monthly mean air temperature. The respiration rate, r, of individual trees increased with increasing tree dimensions, such as stem volume, v(S), and stem girth at the base of the live crown, G(B). The dependence of respiration rate on tree size was successfully represented by a power function. The r - v(S) dependence was rather stronger than the r - G(B) (2) dependence, especially toward the end of the growing season (from July to September). The observed respiration rate was almost the same as the respiration rate corrected for the monthly mean air temperature. The annual respiration of individual trees was directly proportional to their phytomass or to its increment. Although the annual respiration of individual trees decreased proportionally to the square root of the leaf mass, it decreased abruptly in the range close to the smallest sample tree. Combining the monthly relationship between respiration rate and stem volume with the tree size distribution in the stand, the stand aboveground annual respiration was estimated to be 20.4 Mg CO(2) ha(-1) year(-1) (= 12.5 Mg dry mass ha(-1) year(-1)) for an aboveground biomass of 17.4 Mg ha(-1) with an annual increment of 6.51 Mg ha(-1) year(-1), i.e., the stand aboveground annual respiration amounted to the equivalent of 72% of the biomass or to almost twice the biomass increment.     

Maintenance and growth respiration of the aboveground parts of young field-grown hinoki cypress (Chamaecyparis obtusa)

Aboveground respiration of five 8-year-old trees of field-grown hinoki cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) was nondestructively measured at monthly intervals over 1 year with an enclosed standing tree method. The relationship between monthly specific respiration rate and monthly mean relative growth rate at the individual tree level was described by a linear equation. During the dormant season, respiration was used mainly for maintenance purposes, whereas during the growing season, more than 40% of the respiration was used for growth purposes, i.e., 60 to 70% in May. We conclude that annual maintenance and growth respiration of a tree are directly proportional to the aboveground phytomass and its annual increment, respectively. The maintenance coefficient was estimated to be 0.504 +/- 0.039 (SE) kg kg(-1) year(-1), indicating that the amount respired for maintaining already existing phytomass was equivalent to about half of the existing phytomass. The growth coefficient was estimated to be 0.772 +/- 0.043 (SE) kg kg(-1), indicating that the amount respired for constructing new phytomass was equivalent to about three-fourths of the annual phytomass increment. The annual stand maintenance and growth respiration were, respectively, 8.8 Mg ha(-1) year(-1) for an aboveground biomass of 17.4 Mg ha(-1) and 5.0 Mg ha(-1) year(-1) for an annual stand aboveground biomass increment of 6.5 Mg ha(-1) year(-1). About two-thirds of the total respiration was used to maintain already existing biomass, and about one-third was used to construct new biomass.     

Hydraulic conductance, light interception and needle nutrient concentration in Scots pine stands and their relations with net primary productivity

Aboveground xylem hydraulic conductance was determined in Scots pine (Pinus sylvestris L.) trees and stands from 7 to about 60 years of age. At the stand scale, leaf area index and net primary productivity (NPP, above- plus belowground) increased and reached a plateau at about 25-30 and 15-20 years, respectively; both parameters declined in mature stands. Stand hydraulic conductance followed a similar trend to NPP, with a maximum at about 15-20 years and a pronounced reduction in old stands. At the tree scale, annual biomass growth per unit of leaf area (growth efficiency) declined with tree age, whereas aboveground sapwood volume per unit leaf area, which is linearly related to maintenance respiration costs, steadily increased. Radiation interception per unit leaf area increased significantly with reduced leaf area index of mature stands, despite increased foliage clumping in the canopies of mature trees. Needle nutrient concentration did not change in the chronosequence. Tree hydraulic conductance per unit leaf area was strongly and positively correlated with growth efficiency. We discuss our findings in the context of growth reductions in mature and old trees, and suggest that increased hydraulic resistance and maintenance respiration costs may be the main causes of reduced carbon gain in mature and old trees.     

Biomass and carbon pools of disturbed riparian forests

Quantification of carbon pools as affected by forest age/development can facilitate riparian restoration and increase awareness of the potential for forests to sequester global carbon. Riparian forest biomass and carbon pools were quantified for four riparian forests representing different seral stages in the South Carolina Upper Coastal Plain. Three of the riparian forests were recovering from disturbance (thermal pollution), whereas the fourth represents a mature, relatively undisturbed riparian forest. Above and belowground carbon pools were determined from linear transects established perpendicular to the main stream channels and spanning the width of the riparian area. The objective of this study was to quantify the biomass and carbon pools in severely disturbed, early successional bottomland hardwood riparian forests and to compare these values to those of a less disturbed, mature riparian forest.

Aboveground biomass in all four riparian forests increased during the 2.5-year investigation period. The total carbon pool in these South Carolina Coastal Plain riparian forests increased with forest age/development due to greater tree and soil carbon pools. The mature riparian forest stored approximately four times more carbon than the younger stands. The importance of the herbaceous biomass layer and carbon pool declined relative to total aboveground biomass with increasing forest age. As stands grew older fine root biomass increased, but an inverse relationship existed between percentages of fine root biomass to total biomass. The root carbon pool increased with forest age/development due to a combination of greater fine root biomass and higher root percent carbon.

Aboveground net primary production (NPP) in young riparian forests rapidly approached and exceeded NPP of the more mature riparian forest. As a woody overstory became established (after 8–10 years) annual litterfall rate as a function of NPP was independent of forest age and litterfall amount in the young riparian forests was comparable to mature riparian forests. Biomass in the riparian forest floor and carbon pool declined with increasing riparian forest development. Woody debris in these riparian forests comprised a relatively small carbon pool. An understanding of bottomland hardwood riparian forest carbon pools at different stages of succession allows us to assess how time since disturbance influences these pools, leading to a better understanding of the recovery processes.     

Trends in bole biomass accumulation, net primary production and tree mortality in Pseudotsuga menziesii forests of contrasting age

Although it is generally accepted that the rate of accumulation of biomass declines as forests age, little is known about the relative contributions to this decline of changes in net primary production (NPP) and tree mortality. We used 10-15 years of observations of permanent plots in three small watersheds in and near the H.J. Andrews Experimental Forest, Oregon, to examine these issues. The three watersheds are of similar elevation and potential productivity and support young (29 years at last measurement), mature (approximately 100 years) and old (approximately 400 years) forest dominated by Pseudotsuga menziesii (Mirb.) Franco and Tsuga heterophylla (Raf.) Sarg. Accumulation of tree bole biomass was greatest in the young stand, reaching approximately 7 Mg ha-1 year-1 in the last measurement interval. Bole biomass accumulation was relatively constant (approximately 4-5 Mg ha-1 year-1) in the mature stand, and there was no net accumulation of bole biomass in the old-forest stand. The NPP of boles increased with time in the young stand, from approximately 3 to approximately 7 Mg ha-1 year-1, but was nearly constant in the mature and old-forest stands, at approximately 6 and 3-4 Mg ha-1 year-1, respectively. Mortality increased slowly in the young stand (from < 0.1 to 0.3 Mg ha-1 year-1), but fluctuated between 1-2 and 2-6 Mg ha-1 year-1 in the mature and old-forest stands, respectively. Thus, declining biomass accumulation with stand age reflects, in approximately equal amounts, both decreasing NPP and increasing mortality.     

Regional assessment of boreal forest productivity using an ecological process model and remote sensing parameter maps

An ecological process model (BIOME-BGC) was used to assess boreal forest regional net primary production (NPP) and response to short-term, year-to-year weather fluctuations based on spatially explicit, land cover and biomass maps derived by radar remote sensing, as well as soil, terrain and daily weather information. Simulations were conducted at a 30-m spatial resolution, over a 1205 km(2) portion of the BOREAS Southern Study Area of central Saskatchewan, Canada, over a 3-year period (1994-1996). Simulations of NPP for the study region were spatially and temporally complex, averaging 2.2 (+/- 0.6), 1.8 (+/- 0.5) and 1.7 (+/- 0.5) Mg C ha(-1) year(-1) for 1994, 1995 and 1996, respectively. Spatial variability of NPP was strongly controlled by the amount of aboveground biomass, particularly photosynthetic leaf area, whereas biophysical differences between broadleaf deciduous and evergreen coniferous vegetation were of secondary importance. Simulations of NPP were strongly sensitive to year-to-year variations in seasonal weather patterns, which influenced the timing of spring thaw and deciduous bud-burst. Reductions in annual NPP of approximately 17 and 22% for 1995 and 1996, respectively, were attributed to 3- and 5-week delays in spring thaw relative to 1994. Boreal forest stands with greater proportions of deciduous vegetation were more sensitive to the timing of spring thaw than evergreen coniferous stands. Similar relationships were found by comparing simulated snow depth records with 10-year records of aboveground NPP measurements obtained from biomass harvest plots within the BOREAS region. These results highlight the importance of sub-grid scale land cover complexity in controlling boreal forest regional productivity, the dynamic response of the biome to short-term interannual climate variations, and the potential implications of climate change and other large-scale disturbances.     

Photosynthetic utilization efficiency of absorbed photosynthetically active radiation by Scots pine and birch forest stands in the southern Taiga

Absorption and utilization of photosynthetically active radiation (PAR) were investigated in Scots pine (Pinus sylvestris L.) and birch (Betula pendula Roth.) stands that were 41 years old at the end of the experimental period. Canopy depth of the Scots pine stand was about half that of the birch stand (6.5 versus 11.0 m), but absorption of PAR was similar in the two stands. The Scots pine forest canopy, with a leaf area index of 8.9, absorbed 90% of the incoming PAR (APAR), whereas the birch forest canopy, with a leaf area index of 5.9, absorbed 92% of APAR. During maximum foliage development, the upper Scots pine canopy absorbed more PAR than the upper birch canopy (75 versus 66%). The upper, middle and lower layers of the Scots pine canopy contained 37, 48 and 15% of the total needle surface area, respectively. The corresponding distribution of foliage surface area in the three layers of the birch canopy was 50, 30 and 20%, respectively. Measurements of photosynthetic rate were combined with estimates of leaf area index and stand phytomass to determine rates of primary production on a sunny day, a cloudy day, and on an annual basis. The energy equivalents of short- and long-term carbon gain were used with determinations of APAR to calculate photosynthetic utilization efficiency. Throughout the growing season, photosynthetic utilization efficiency of APAR in the upper canopy layer of the Scots pine forest was almost twice that in the lower canopy layer. In the birch forest, photosynthetic utilization efficiency was greater in the lower canopy layer than in the upper canopy layer. In all cases, utilization efficiency was higher in the birch stand than in the Scots pine stand (52 versus 29 J kJ(-1)). Taking account of respiration of the non-photosynthetic parts of each stand (night respiration of needles or leaves; respiration of branches, trunk and roots), estimated utilization efficiency of APAR for net primary production was 11 J kJ(-1) for Scots pine and 19 J kJ(-1) for birch. Solar conversion ratios, expressed as whole-plant net primary productivity per unit of APAR for the growing season, were 0.81 g MJ(-1) for Scots pine and 0.93 g MJ(-1) for birch.     

Quantification and Understanding of Above and Belowground Biomass in Medium Saline Zone of the Sundarbans,Bangladesh: The Relationships with Forest Attributes

ABSTRACT

The relationship between stand attributes and biomass accumulation pattern in a mangrove forest has been intensively studied in this study. We assessed above (AGM) and belowground mass (BGM) and examined the relationship between forest attributes and aboveground mass in the Sundarbans, Bangladesh. The study was conducted with 18 plots having total area of 1.08 ha. The mean AGM and BGM of the study sites were 234.08 and 132.85 Mg ha^?1 respectively. H. fomes contributed the highest amount (82.9% of total AGM and 80.53% of total BGM) of above (193.56 Mg ha^?1) and belowground mass (107.09 Mg ha^?1) at the study site. Our study revealed structural attributes (tree diameter, height, and basal area) positively correlated with AGM. In contrast, species richness and species diversity negatively correlated with AGM. Our study indicated that lack of positive relationship between species diversity and AGM which may be attributable to high AGM of the dominant species (H. fomes) and may have a considerable consequence in AGM of the study area. Thus, maintaining large trees (DBH and height) rather than species diversity in the Sundarbans mangrove forest might be an effective approach for increasing aboveground mass.     

Quantitative relationships between fine roots and stand characteristics

Fine roots absorb nutrients and water for photosynthesizing leaves, which in return provide them with hydrocarbon products. Knowledge of the fine root biomass (FRB) at the individual tree level and its relationships with other components related to tree growth, especially leaves aboveground, is scarce. Therefore, we reviewed the FRB of major forest-forming species using a database of 518 forest stands compiled from the literature, including 21 tree species and 16 shrub species, in order to confirm the relationships between environmental or forest stand variables and FRB at the stand and tree levels, and we further determine the relationships between fine roots belowground and leaves aboveground. Correlations between FRB and site characteristics (latitude, elevation, age, density, and basal area) appeared to be species-specific. There were hardly any significant correlations between stand FRB and latitude, elevation, stand age and stand density. Tree FRB was better correlated with tree basal area than stand FRB with stand basal area. There was a significant linear relationship between tree FRB and tree basal area. In addition, individual FRB was significantly linearly related to leaf biomass for all analyzed species. When these species were grouped into coniferous and deciduous, or all species together, there were still significant linear relationships between tree FRB and tree basal area and leaf biomass. The ratios of FRB to leaf biomass varied between and among species and even among regions for the same species. For both Picea abies and Pinus sylvestris, the ratio of FRB to leaf biomass was negatively related to the ratio of annual actual evapotranspiration to annual potential evapotranspiration, which was an indicator of water availability.     

Aboveground biomass and nutrient allocation in an age-sequence of <Emphasis Type="Italic">Larix olgensis</Emphasis> plantations

Biomass and nutrient (N, P, K, Ca, Mg) stock in various aboveground tree components (stemwood, stembark, branches and leaves) were quantified in an age sequence of pure Larix olgensis planta- tions (20, 35, 53 and 69 years old) in Northeast China. The results show that the aboveground biomass allocation in various tree components was in the order of stemwood (62%-83%), branches (9%-21%), stembark (7%-11%) and leaves (1%-6%) for all stands. The proportion of stemwood biomass to total aboveground biomass increased whereas that of other tree components decreased consistently with stand age from 20 to 53 years old, but kept relatively constant with stand age from 53 and 69 years old. The nutrient allocation in various tree components generally followed the same pattern as the biomass allocation (i.e. stemwood > branches > stembark > leaves). The proportion of nutrient stock in leaves to total aboveground nutrient stock decreased consistently with increasing stand age, while that in stemwood increased with stand age from 20 to 53 years old but then decreased from 53 to 69 years old. The rate of nutrient removal for stands was estimated at different stand ages under different logging schemes, showing that the rate of nutrient removal would be unchanged when the rotation length was shortened to 20 years by the harvest of stem only, but greatly increased by the harvest of total aboveground biomass. The rate of nutrient removal would be a considerable reduction for all elements by debarking, especially for Ca.