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.     

Changes in the relationship between tree size and aboveground respiration in field-grown hinoki cypress (Chamaecyparis obtusa) trees over three years

Respiration measurements of aerial parts of 18-year-old hinoki cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) trees were made under field conditions over three years to study changing relationships with tree age between respiration and phytomass, phytomass increment, and leaf mass. The relationship between annual respiration (r(a)) and phytomass (w(T)) was approximated by a proportional function (r(a) = aw(T)), where the proportional constant (a) decreased year by year. The effect of time on the relationship between annual respiration and phytomass of each sample tree was fitted by a power function. Respiration of the tree suppressed by the canopy decreased year by year, but respiration of the other trees increased slightly with age. The relationship between annual respiration and leaf mass was also approximated by a generalized power function. Excluding the suppressed tree, the relationship between annual respiration (r(a)) and the annual increment of aboveground phytomass (Deltaw(T)) was described by a proportional function (r(a) = 2.27Deltaw(T)), where the proportional constant, 2.27, was independent of sample tree and year, indicating that about 2.3 times of the annual aboveground phytomass increment equivalent was respired annually. For any tree, the time constant relationships between annual respiration and leaf mass and phytomass increment for different-sized trees were similar to the corresponding time continuum relationships. In contrast, the time continuum relationship between annual respiration and phytomass differed from the time constant relationship, indicating that respiration of less active woody tissue contributed significantly to aboveground respiration. Based on the relationship between tree size and annual respiration, annual aboveground stand respiration was estimated to be 25.0, 26.9, and 25.8 Mg(dm) ha(-1) year(-1) for the three consecutive years, respectively, and the corresponding aboveground stand biomass was 60.0, 69.0, and 76.8 Mg(dm) ha(-1).     

Root respiration in Chamaecyparis obtusa trees

We examined the respiration rate of root segments, which had a constant length in relation to their diameter, from three small and two large 26-year-old Chamaecyparis obtusa (Siebold & Zucc.) Endl. trees. The dependence of respiration rate on segment diameter was described by a power function with an exponent of about 1.5, except for the smallest sample tree, for which the exponent was 1.74. Unlike stem segments, root segments of similar diameter showed similar rates of respiration regardless of the tree from which the root segments had been taken. On the basis of the power function, we propose a new equation to estimate the total root respiration rate of a tree. The relationship between root respiration rate per tree and root weight can be expressed by a power function with an exponent of 1.11. The ratio of the specific respiration rate of stems to that of roots was 0.7 for the three smaller trees, and 1.1 to 1.3 for the two larger trees. In November, the stand respiration rate of roots was estimated to be 0.36 kg CO(2) ha(-1) h(-1) for a root biomass (dry weight) of 28 Mg ha(-1).     

Spatial and Temporal Patterns in Stand Structure,Biomass, Growth,and Mortality in a Monospecific Nothofagus solandrivar. cliffortioides (Hook,f.) Poole Forest in New Zealand

Abstract

In 250 20 m X 20 m permanent plots in the Craigie-burn Range, Canterbury, New Zealand, 1970 stem density was 2,191/ha, basal area was 52.4 m²/ha, and stem biomass was 178.1 Mg/ha. Net production of stemwood (1974-1987) was 2.0 Mg/ha/yr; mortality was 3.5 Mg/ha/yr. By 1987 density had decreased by 30%, basal area by 12%, and stem biomass by 13%.

Stands with many short trees of small mean dbh were common at high elevation, whereas stands with fewer, taller trees with large mean dbh were common at low elevation. Stemwood production and mortality rate were higher in tall stands. Mortality was well distributed among plots, indicating small, frequent canopy openings; stand turnover calculations were 66 year (based on 2.2% annual biomass loss) to 83 year (based on 1.2% annual stemwood production). Larger canopy openings were also evident, but were more infrequent, so stand turnover times due to 'catastrophic' disturbances were in the range of 350-4000 yr. Consequently, the small, high-frequency disturbances blurred effects of larger disturbances on stand structure and also constrained the fluctuation in forest biomass.     

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.     

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.     

Above- and belowground biomass and net primary production in a 73-year-old Scots pine forest

We estimated above- and belowground biomass and net primary production (NPP) of a 73-year-old Scots pine (Pinus sylvestris L.) forest stand in the Belgian Campine region. Total biomass for the stand was 176 Mg ha(-1), of which 74.4% was found in stems. The root system contained 12.6% of total biomass, most of it in coarse roots (> 5 mm). Fine roots (< 5 mm) comprised only about 1.7% of total biomass, and more than 50% of fine root biomass was retrieved in the litter layer and the upper 15 cm of the mineral soil. The ratio of belowground biomass to aboveground biomass was 0.14, which is lower than that of other Scots pine forests and other coniferous forests. Between 1995 and 2001, mean annual NPP was 11.2 Mg ha(-1) year(-1), of which 68.7% was allocated to aboveground compartments. Stems, needles and cones made relatively high contributions to total NPP compared with branches. However, branch NPP was possibly underestimated because litterfall of big branches was neglected. The proportion of total NPP in belowground components was 31.3%. Coarse root NPP (2% of total) was low compared with its biomass. Fine root NPP was 3.3 Mg ha(-1) year(-1), representing about 29.5% of total NPP; however, the estimate of fine root NPP is much more uncertain than NPP of aboveground compartments. The ratio NPP/GPP (gross primary production) was 0.32, which was low compared with other coniferous forests.     

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.     

Changes in composition, structure and aboveground biomass over seventy-six years (1930-2006) in the Black Rock Forest, Hudson Highlands, southeastern New York State

We sought to quantify changes in tree species composition, forest structure and aboveground forest biomass (AGB) over 76 years (1930-2006) in the deciduous Black Rock Forest in southeastern New York, USA. We used data from periodic forest inventories, published floras and a set of eight long-term plots, along with species-specific allometric equations to estimate AGB and carbon content. Between the early 1930s and 2000, three species were extirpated from the forest (American elm (Ulmus americana L.), paper birch (Betula papyrifera Marsh.) and black spruce (Picea mariana (nigra) (Mill.) BSP)) and seven species invaded the forest (non-natives tree-of-heaven (Ailanthus altissima (Mill.) Swingle) and white poplar (Populus alba L.) and native, generally southerly distributed, southern catalpa (Catalpa bignonioides Walt.), cockspur hawthorn (Crataegus crus-galli L.), red mulberry (Morus rubra L.), eastern cottonwood (Populus deltoides Bartr.) and slippery elm (Ulmus rubra Muhl.)). Forest canopy was dominated by red oak and chestnut oak, but the understory tree community changed substantially from mixed oak-maple to red maple-black birch. Density decreased from an average of 1500 to 735 trees ha(-1), whereas basal area doubled from less than 15 m(2) ha(-1) to almost 30 m(2) ha(-1) by 2000. Forest-wide mean AGB from inventory data increased from about 71 Mg ha(-1) in 1930 to about 145 Mg ha(-1) in 1985, and mean AGB on the long-term plots increased from 75 Mg ha(-1) in 1936 to 218 Mg ha(-1) in 1998. Over 76 years, red oak (Quercus rubra L.) canopy trees stored carbon at about twice the rate of similar-sized canopy trees of other species. However, there has been a significant loss of live tree biomass as a result of canopy tree mortality since 1999. Important constraints on long-term biomass increment have included insect outbreaks and droughts.     

Carbon stocks and soil respiration rates during deforestation, grassland use and subsequent Norway spruce afforestation in the Southern Alps, Italy

Changes in carbon stocks during deforestation, reforestation and afforestation play an important role in the global carbon cycle. Cultivation of forest lands leads to substantial losses in both biomass and soil carbon, whereas forest regrowth is considered to be a significant carbon sink. We examined below- and aboveground carbon stocks along a chronosequence of Norway spruce (Picea abies (L.) Karst.) stands (0-62 years old) regenerating on abandoned meadows in the Southern Alps. A 130-year-old mixed coniferous Norway spruce-white fir (Abies alba Mill.) forest, managed by selection cutting, was used as an undisturbed control. Deforestation about 260 years ago led to carbon losses of 53 Mg C ha(-1) from the organic layer and 12 Mg C ha(-1) from the upper mineral horizons (Ah, E). During the next 200 years of grassland use, the new Ah horizon sequestered 29 Mg C ha(-1). After the abandonment of these meadows, carbon stocks in tree stems increased exponentially during natural forest succession, levelling off at about 190 Mg C ha(-1) in the 62-year-old Norway spruce and the 130-year-old Norway spruce-white fir stands. In contrast, carbon stocks in the organic soil layer increased linearly with stand age. During the first 62 years, carbon accumulated at a rate of 0.36 Mg C ha(-1) year(-1) in the organic soil layer. No clear trend with stand age was observed for the carbon stocks in the Ah horizon. Soil respiration rates were similar for all forest stands independently of organic layer thickness or carbon stocks, but the highest rates were observed in the cultivated meadow. Thus, increasing litter inputs by forest vegetation compared with the meadow, and constantly low decomposition rates of coniferous litter were probably responsible for continuous soil carbon sequestration during forest succession. Carbon accumulation in woody biomass seemed to slow down after 60 to 80 years, but continued in the organic soil layer. We conclude that, under present climatic conditions, forest soils act as more persistent carbon sinks than vegetation that will be harvested, releasing the carbon sequestered during tree growth.     

Net ecosystem productivity,net primary productivity and ecosystem carbon sequestration in a Pinus radiata plantation subject to soil water deficit

Tree carbon (C) uptake (net primary productivity excluding fine root turnover, NPP') in a New Zealand Pinus radiata D. Don plantation (42 degrees 52' S, 172 degrees 45' E) growing in a region subject to summer soil water deficit was investigated jointly with canopy assimilation (A(c)) and ecosystem-atmosphere C exchange rate (net ecosystem productivity, NEP). Net primary productivity was derived from biweekly stem diameter growth measurements using allometric relations, established after selective tree harvesting, and a litterfall model. Estimates of A(c) and NEP were used to drive a biochemically based and environmentally constrained model validated by seasonal eddy covariance measurements. Over three years with variable rainfall, NPP' varied between 8.8 and 10.6 Mg C ha(-1) year(-1), whereas A(c) and NEP were 16.9 to 18.4 Mg C ha(-1) year(-1) and 5.0-7.2 Mg C ha(-1) year(-1), respectively. At the end of the growing season, C was mostly allocated to wood, with nearly half (47%) to stems and 27% to coarse roots. On an annual basis, the ratio of NEP to stand stem volume growth rate was 0.24 +/- 0.02 Mg C m(-3). The conservative nature of this ratio suggests that annual NEP can be estimated from forest yield tables. On a biweekly basis, NPP' repeatedly lagged A(c), suggesting the occurrence of intermediate C storage. Seasonal NPP'/A(c) thus varied between nearly zero and one. On an annual basis, however, NPP'/A(c) was 0.54 +/- 0.03, indicating a conservative allocation of C to autotrophic respiration. In the water-limited environment, variation in C sequestration rate was largely accounted for by a parameter integrative for changes in soil water content. The combination of mensurational data with canopy and ecosystem C fluxes yielded an estimate of heterotrophic respiration (NPP' - NEP) approximately 30% of NPP' and approximately 50% of NEP. The estimation of fine-root turnover rate is discussed.     

Carbon budget for Scots pine trees: effects of size, competition and site fertility on growth allocation and production

Time series of carbon fluxes in individual Scots pine (Pinus sylvestris L.) trees were constructed based on biomass measurements and information about component-specific turnover and respiration rates. Foliage, branch, stem sapwood, heartwood and bark components of aboveground biomass were measured in 117 trees sampled from 17 stands varying in age, density and site fertility. A subsample of 32 trees was measured for belowground biomass excluding fine roots. Biomass of fine roots was estimated from the results of an earlier study. Statistical models were constructed to predict dry mass (DW) of components from tree height and basal area, and time derivatives of these models were used to estimate biomass increments from height growth and basal area growth. Biomass growth (G) was estimated by adding estimated biomass turnover rates to increments, and gross photosynthetic production (P) was estimated by adding estimated component respiration rates to growth. The method, which predicts the time course of G, P and biomass increment in individual trees as functions of height growth and basal area growth, was applied to eight example trees representing different dominance positions and site fertilities. Estimated G and P of the example trees varied with competition, site fertility and tree height, reaching maximum values of 22 and 43 kg(DW) year(-1), respectively. The site types did not show marked differences in productivity of trees of the same height, although height growth was greater on the fertile site. The G:P ratio decreased with tree height from 65 to 45%. Growth allocation to needles and branches increased with increasing dominance, whereas growth allocation to the stem decreased. Growth allocation to branches decreased and growth allocation to coarse roots increased with increasing tree size. Trees at the poor site allocated 49% more to fine roots than trees at the fertile site. The belowground parts accounted for 25 to 55% of annual G, increasing with tree size and decreasing with site fertility. Annual G and P per unit needle mass varied over the ranges 1.9-2.4 and 3.5-4.0 kg(DW) kg(-1), respectively. The relationship between P and needle mass in the example trees was linear and relatively independent of competition, site fertility and age.     

Fruit development, not GPP, drives seasonal variation in NPP in a tropical palm plantation

We monitored seasonal variations in net primary production (NPP), estimated by allometric equations from organ dimensions, gross primary production (GPP), estimated by the eddy covariance method, autotrophic respiration (R(a)), estimated by a model, and fruit production in a coconut (Cocos nucifera L.) plantation located in the sub-tropical South Pacific archipelago of Vanuatu. Net primary production of the vegetative compartments of the trees accumulated steadily throughout the year. Fruits accounted for 46% of tree NPP and showed large seasonal variations. On an annual basis, the sum of estimated NPP (16.1 Mg C ha(-1) year(-1)) and R(a) (24.0 Mg C ha(-1) year(-1)) for the ecosystem (coconut trees and herbaceous understory) closely matched GPP (39.0 Mg C ha(-1) year(-1)), suggesting adequate cross-validation of annual C budget methods. However, seasonal variations in NPP + R(a) were smaller than the seasonal variations in GPP, and maximum tree NPP occurred 6 months after the midsummer peak in GPP and solar radiation. We propose that this discrepancy reflects seasonal variation in the allocation of dry mass to carbon reserves and new plant tissue, thus affecting the allometric relationships used for estimating NPP.     

Component and whole-system respiration fluxes in northern deciduous forests

We measured component and whole-system respiration fluxes in northern hardwood (Acer saccharum Marsh., Tilia americana L., Fraxinus pennsylvanica Marsh.) and aspen (Populus tremuloides Michx.) forest stands in Price County, northern Wisconsin from 1999 through 2002. Measurements of soil, leaf and stem respiration, stem biomass, leaf area and biomass, and vertical profiles of leaf area were combined with biometric measurements to create site-specific respiration models and to estimate component and whole-system respiration fluxes. Hourly estimates of component respiration were based on site measurements of air, soil and stem temperature, leaf mass, sapwood volume and species composition. We also measured whole-system respiration from an above-canopy eddy flux tower. Measured soil respiration rates varied significantly among sites, but not consistently among dominant species (P < 0.05 and P > 0.1). Annual soil respiration ranged from 8.09 to 11.94 Mg C ha(-1) year(-1). Soil respiration varied linearly with temperature (P < 0.05), but not with soil water content (P > 0.1). Stem respiration rates per unit volume and per unit area differed significantly among species (P < 0.05). Stem respiration per unit volume of sapwood was highest in F. pennsylvanica (up to 300 micro mol m(3) s(-1)) and lowest in T. americana (22 micro mol m(3) s(-1)) when measured at peak summer temperatures (27 to 29 degrees C). In northern hardwood stands, south-side stem temperatures were higher and more variable than north-side temperatures during leaf-off periods, but were not different statistically during leaf-on periods. Cumulative annual stem respiration varied by year and species (P < 0.05) and averaged 1.59 Mg C ha(-1) year(-1). Leaf respiration rates varied significantly among species (P < 0.05). Respiration rates per unit leaf mass measured at 30 degrees C were highest for P. tremuloides (38.8 nmol g(-1) s(-1)), lowest for Ulmus rubra Muhlenb. (13.1 nmol g(-1) s(-1)) and intermediate and similar (30.2 nmol g(-1) s(-1)) for T. americana, F. pennsylvanica and Q. rubra. During the growing season, component respiration estimates were dominated by soil respiration, followed by leaf and then stem respiration. Summed component respiration averaged 11.86 Mg C ha(-1) year(-1). We found strong covariance between whole-ecosystem and summed component respiration measurements, but absolute rates and annual sums differed greatly.     

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.     

Thinning alters crown dynamics and biomass increment within aboveground tissues in young stands of Chamaecyparis obtusa

The stand density of a forest affects the vertical distribution of foliage. Understanding the dynamics of this response is important for the study of crown structure and function, carbon-budget estimation, and forest management. We investigated the effect of tree density on the vertical distribution of foliage, branch, and stem growth, and ratio of biomass increment in aboveground tissues; by monitoring all first-order branches of five trees each from thinned and unthinned control stands of 10-year-old Chamaecyparis obtusa for four consecutive years. In the control stand, the foliage crown shifted upward with height growth but the foliage quantity of the whole crown did not increase. In addition, the vertical distribution of leaf mass shifted from lower-crown skewed to upper-crown skewed. In the thinned stand in contrast, the foliage quantity of individual crowns increased two-fold within 4 years, while the vertical distribution of leaf mass remained lower-crown skewed. The two stands had similar production rates, numbers of first-order branches per unit of tree height, and total lengths of first-order branches. However, the mortality rate of first-order branches and self-pruning within a first-order branch were significantly higher in the control stand than in the thinned stand, which resulted in a higher ratio of biomass increment in branch. Thinning induced a higher ratio of biomass increment in foliage and lower in branch. The increased foliage quantity and variation in ratio of biomass increment after thinning stimulated stem growth of residual trees. These results provide information that will be useful when considering thinning regimes and stand management.     

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.     

Impacts of fructification on biomass production and correlated genetic effects in Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> [L.] Karst.)

For the period 2003–2006, fructification of Norway spruce (Picea abies [L.] Karst.) was recorded at the Kranzberg forest site in Southern Germany by employing a crane with access to the canopy of more than 266 trees. For each tree, stem diameter and growth parameters were assessed annually as well as biomass of cones and seeds, number of seeds per cone, and proportions of empty seeds for a total of 371 trees with cone crop. Genotypes at 19 enzyme coding gene loci of 110 trees were included in the study of correlations between morphological and genetic traits. Re-scaling the observed values for a virtual pure Norway spruce stand of 1 ha, cone biomass including winged seeds (oven-dried at 38°C) varied between 706.8 kg/ha in 2006 (average value per tree was 3.6 kg) and values close to zero in 2005. Corresponding values for vegetative biomass increment of the coning trees in 2006 were 9,273.0 kg/ha and 10.8 kg/tree. A significant higher biomass investment was determined for dominant trees in terms of absolute cone mass as well as in terms of cone mass relative to vegetative biomass and fructification frequency. No trade-off effects in decreased vegetative biomass growth were found in the fructification year, compared to trees that did not grow cones. Although the dominant trees invested proportionally considerable biomass in cones, they showed no significant reduction in vegetative biomass growth. In the following year no decrease in vegetative growth was detected. Based on logistic regressions and homogeneity tests, respectively, significant genetic effect became evident with respect to the gene loci AAP-B and AAT-C concerning fructification probability in the year with maximum generative biomass investment. These and closely related loci also have been found to be indicative for growth and viability, respectively, in other species.     

Growth and water relations of Eucalyptus marginata (jarrah) stands in response to thinning and fertilization

We studied the effects of five thinning treatments (T1 = 5.5, T2 = 11, T3 = 16.5, T4 = 22.5 and T5 = 28.5 m(2) ha(-1) basal area under bark) x two fertilizer treatments (F0 = unfertilized and F1 = fertilized with 400 kg ha(-1) N plus 229 kg ha(-1) P) on growth and water relations of pole-sized Eucalyptus marginata J. Donn ex Sm. trees growing in southwestern Australia. Thinning reduced leaf area index (LAI) from 2.1 in the T4 and T5 treatments to 0.8 in the T1F0 treatment. Fertilizer had no effect on LAI in the T2, T4 or T5 treatments, but increased LAI by 45 and 20% in the T1 and T3 treatments, respectively. Thinning plus fertilizing increased diameter growth most in the fastest growing trees, from 0.4 cm year(-1) for trees in the T5F0 and T5F1 treatments to 0.7 and 1.2 cm year(-1) for trees in the T1F0 and T1F1 treatments, respectively. In both fertilizer treatments, stand basal area and volume growth increased with increasing stand density up to 15 m(2) ha(-1), and thereafter declined with increasing stand density, such that the growth rate of trees in the T5 treatment was only half of that at a stand density of 15 m(2) ha(-1). In response to fertilizer, growth rates of the slowest and fastest-growing trees increased from 0.35 and 3.5 m(2) ha(-1) year(-1) (F0) to 0.56 and 5.4 m(3) ha(-1) year(-1) (F1), respectively. Stand growth efficiency (growth per unit LAI) increased in response to thinning, and fertilizer increased stand growth efficiency at all stand densities. Throughout the dry season, T5 trees had lower predawn shoot water potentials (Psi(pd)) (minimum of -1.5 MPa) than T1 or T2 trees (minimum of -0.7 MPa). Fertilizer decreased Psi(pd) in T5 trees (by -0.9 and -1.5 MPa, respectively, in F0 and F1), but not in T1 or T2 trees. Stand growth rate was closely related to cumulative midday water stress (CMWS) over the dry season, and volume growth rate declined sharply from 6 m(3) ha(-1) year(-1) at a CMWS of 130 MPa days, to zero at a CMWS of 220 MPa days. Application of fertilizer to thinned stands increased LAI, stand growth efficiency and stand growth. In unthinned stands, fertilizer increased stand growth efficiency and stand growth; however, it also increased tree water stress, which limited the fertilizer-induced increases in LAI and growth. We attribute the increase in tree and stand growth in response to application of fertilizer to increased photosynthetic rates, increased allocation to stem wood, and in thinned stands also to higher LAIs.     

Structure, allometry, and biomass of plantation Metasequoia glyptostroboides in Japan

We quantified structural features and the aboveground biomass of the deciduous conifer, Metasequoia glyptostroboides (Hu and Cheng) in six plantations in central Japan. In order to derive biomass estimates we dissected 14 M. glyptostroboides trees into three structural components (stem wood, branch wood and foliage) to develop allometric equations relating the mass of these components and of the whole tree to diameter at breast height (DBH). We found robust relationships at the branch and whole tree level that allow accurate prediction of component and whole tree biomass. Dominant tree height was similar within five older (>40 years) plantations (27–33 m) and shorter in a 20-year-old plantation (18 m). Average stem diameter varied from 12.8 cm in the youngest stand to greater than 35 cm in the oldest stand.

Metasequoia have relatively compact crowns distributed over the top 30% of the tree although the youngest stand had the deepest crown relative to tree height (up to 38%). At the individual tree level in older stands, 87% of the aboveground biomass was allocated to the stem, 9% to branch wood and 4% to foliage. We found little difference in the relative distribution of above ground biomass among the stands with the exception of lower foliage biomass in larger diameter trees. Total aboveground biomass of the older stands varied twofold, ranging from a maximum of 450 Mg ha⁻¹ in a 42-year-old stand to a minimum of 196 Mg ha⁻¹ in a 48-year-old stand. Total above ground biomass of the 20-year-old stand was 176 Mg ha⁻¹.