Tree biomass allocation is governed by allometry but modulated by optimization

Variations in crown forms promote canopy space-use and productivity in mixed-species forests. However, we have a limited understanding on how this response is mediated by changes in within-tree biomass allocation. Here, we explored the role of changes in tree allometry, biomass allocation and architecture in shaping diversity-productivity relationships (DPRs) in the oldest tropical tree diversity experiment. We conducted whole-tree destructive biomass measurements and terrestrial laser scanning. Spatially explicit models were built at the tree level to investigate the effects of tree size and local neighbourhood conditions. Results were then upscaled to the stand level, and mixture effects were explored using a bootstrapping procedure. Biomass allocation and architecture substantially changed in mixtures, which resulted from both tree-size effects and neighbourhood-mediated plasticity. Shifts in biomass allocation among branch orders explained substantial shares of the observed overyielding. By contrast, root-to-shoot ratios, as well as the allometric relationships between tree basal area and aboveground biomass, were little affected by the local neighbourhood. Our results suggest that generic allometric equations can be used to estimate forest aboveground biomass overyielding from diameter inventory data. Overall, we demonstrate that shifts in tree biomass allocation are mediated by the local neighbourhood and promote DPRs in tropical forests.

Studying tree biomass dynamics and allocation is crucial to understanding the forest carbon cycle and the adaptation of trees to the environment. However, traditional biomass surveys are time-consuming and labor-intensive, so few studies have specifically examined biomass formation in terms of the increase in individual tree biomass, and the role that tree age and site conditions play in this process, especially tree roots, is unclear. We studied the tree ring characteristics of 87 sample trees (8–40 years old) from 29 Chinese fir plantations with different site conditions and measured the biomass of their stems, crowns, and roots. The biomass increment at various age stages during tree growth was determined via using tree ring analysis, and a generalized additive mixed model (GAMM) was used to analyze biomass formation and allocation, as well as the specific impact of site conditions on them. The results showed that the biomass increment of Chinese fir trees first increased and then decreased with age, and improving site conditions delayed the carbon maturation of the trees. The proportion of stem biomass increased with age, while the proportion of crown biomass decreased and the proportion of root biomass increased and then decreased. The effect of the site conditions on the tree biomass allocation showed a nonlinear trend. Tree ring analysis provides a feasible and effective method for assessing tree growth and biomass dynamics. Forest managers can use the findings of this study to scientifically optimize the management of increasing forest carbon sequestration.

Large-scale tree planting programs that store carbon provided by wood and non-wood products are being promoted to mitigate climate change. Assessing the biomass pool of plantations is thus an essential task in forest ecology. This study investigated biomass allocation and allometric equations for above- and belowground components along an age-sequence of Pinus tabuliformis plantations (8, 18, 32, and 46 years old) in northern Hebei Province, China. The biomass of each tree component (root, stem, branch, foliage) was quantified by destructive harvesting. Allometric equations and biomass conversion and expansion factors (BCEFs) were subsequently developed for each tree component. The mean above- and belowground biomass was 5.86, 20.05, 41.26, and 135.28 kg tree−1 and 1.73, 3.42, 11.39, and 27.30 kg tree−1 in the 8-, 18-, 32-, and 46-year-old stands, respectively. The proportion of stem biomass to total tree biomass increased from 28.7% for the 8-year-old stand to 55.8% for 46-year-old stand. In contrast, the contributions of foliage and branch decreased along the chronosequence. The root contribution to total tree biomass also showed a declining trend with stand age. Allometric models based on diameter at breast height showed a good fit (p < 0.001) and incorporating stand age as an additional variable improved the fit of allometric equations (higher R2 and lower ACI) for branch, aboveground, root, and total tree biomass. BCEFs decreased for all tree components as stand age increased. These findings indicate that changes in tree biomass allocation and allometry across stand development must be considered to improve estimates of plantation biomass and carbon stocks at regional and national scales.

Pinus taeda plantations have been facing declining productivity in South America, especially due to competition for natural resources such as light, water, and nutrients. Competition with spontaneous vegetation in the early years is one of the main constraints on growth and biomass allocation in trees. However, the best method and timing for weed control and its impact on the productivity of Pinus taeda plantations are unknown. This study aims to evaluate whether weed control increases the growth and above-ground biomass production of Pinus taeda plantations in southern Brazil. This study was conducted at two sites with five-year-old Pinus taeda plantations in southern Brazil, with each being submitted to different weed control methods. This study was conducted in randomized blocks, with nine treatments: (i) NC—no weed control, i.e., weeds always present; (ii) PC—physical weed control; (iii) CC–T—chemical weed control in the total area; (iv) CC–R—chemical weed control in rows (1.2 m wide); (v) C6m, (vi) C12m, (vii) C18m, and (viii) C24m—weed control up to 6, 12, 18, and 24 months after planting; and (ix) COC—company operational weed control. The following parameters were evaluated: the floristic composition and weed biomass, height, diameter, stem volume, needle biomass, branches, bark, and stemwood of Pinus taeda. Control of the weed competition, especially by physical means (PC), and chemical control over the entire area (CC–T) promoted significant gains in the growth and above–ground biomass production of Pinus taeda at five years of age, particularly at the Caçador site. The results reinforce the importance of using appropriate strategies for managing weed control to maximize productivity, especially before canopy closure. In addition, the strong correlation between growth variables and the total biomass and stemwood indicates the possibility of obtaining indirect estimates through dendrometric measurements. The results contribute to the improvement of silvicultural management in subtropical regions of southern Brazil.

Climate change, especially modified courses of temperature and precipitation, has a significant impact on forest functioning and productivity. Moreover, some alterations in tree biomass allocation (e.g. root to shoot ratio, foliage to wood parts) might be expected in these changing ecological conditions. Therefore, we attempted to model fir stand biomass (t ha−1) along the trans-Eurasian hydrothermal gradients using the data from 272 forest stands. The model outputs suggested that all biomass components, except for the crown mass, change in a common pattern, but in different ratios. Specifically, in the range of mean January temperature and precipitation of −30°C to +10°C and 300 to 900 mm, fir stand biomass increases with both increasing temperature and precipitation. Under an assumed increase of January temperature by 1°C, biomass of roots and of all components of the aboveground biomass of fir stands increased (under the assumption that the precipitation level did not change). Similarly, an assumed increase in precipitation by 100 mm resulted in the increased biomass of roots and of all aboveground components. We conclude that fir seems to be a perspective taxon from the point of its productive properties in the ongoing process of climate change.

A comparative discussion on advantages and disadvantages of natural stands and plantations, including their productivity and resistance, began from the moment of first forest plantings and continues to this day. In the context, progressive replacement of natural forests by plantations, the question of how that will change the carbon storage capacity of forest cover when replacing natural forests with planted ones in a changing climate becomes extremely relevant. This article presents the first attempt to answer this question at the transcontinental level on a special case for two-needles pine trees (subgenus Pinus L.). The research was carried out using the database compiled by the authors on the tree biomass allocation structure for major tree species of Eurasia, in particular, the 1880 and 1967 data of naturally regenerated and planted sample pine trees, respectively. Multi-factor regression models were calculated after combining the matrix of initial data on the structure of tree biomass with the mean temperature of January and mean annual precipitation; their adequacy indices allow us to consider them reproducible. It is found that the aboveground biomass of equal-sized and equal-aged natural and planted trees increases with the rise in the temperature in the month of January and annual precipitation. This pattern is only partially valid for the branches’ biomass. Iit has a specific character for the foliage one. The biomass of all components of planted trees is higher than that of natural trees, but the percentage excess varies among different components and depends on the level of January’s temperature, but does not depend at all on the level of annual precipitation. The uncertainties of estimations, as well as the nature of the obtained regularities, are discussed in the text.

Our research of common aspen (Populus tremula L.) focused on the forested mountainous area in central Slovakia. Forest stands (specifically 27 plots from 9 sites) with ages between 2 and 15 years were included in measurements and sampling. Whole tree biomass of aspen individuals was destructively sampled, separated into tree components (leaves, branches, stem, and roots), and then dried and weighed. Subsamples of fresh leaves from three crown parts (upper, middle, and lower) were scanned, dried, and weighed. Allometric biomass models with stem base diameter as an independent variable were derived for individual tree components. Basic foliage traits, i.e., leaf mass, leaf area, and specific leaf area, were modelled with regard to tree size and leaf position within the crown. Moreover, biomass stock of the woody parts and foliage as well as the leaf area index were modelled using mean stand diameter as an independent variable. Foliage traits changed with both tree size and crown part. Biomass models showed that foliage contribution to total tree biomass decreased with tree size. The total foliage area of a tree increased with tree size, reaching its maximum value of about 12 m2 for a tree with a diameter of 120 mm. Leaf area index increased with mean stand diameter, reaching a maximum value of 13.5 m2 m−2. Since no data for biomass allocation for common aspen had been available at either the tree or stand levels, our findings might serve for both theoretical (e.g., modelling of growth processes) and practical (forestry and agro-forestry stakeholders) purposes.

We studied the effects of stand age on allocation and equation fitting of aboveground and below-ground biomass in four Quercus acutissima stands (14, 31, 46, and 63 years old) in the Central Loess Plateau of China. The stem wood, stem bark, branch, foliage, and belowground biomass of each of the 20 destructive harvesting trees were quantified. The mean total biomass of each tree was 28.8, 106.8, 380.6, and 603.4 kg/tree in the 14-, 31-, 46-, and 63-year-old stands, respectively. Aboveground biomass accounted for 72.25%, 73.05%, 76.14%, and 80.37% of the total tree biomass in the 14-, 31-, 46-, and 63-year-old stands, respectively, and stem wood was the major component of tree biomass. The proportion of stem (with bark) biomass to total tree biomass increased with stand age while the proportions of branch, foliage, and belowground biomass to total tree biomass decreased with stand age. The ratio of belowground biomass to aboveground biomass decreased from 0.39 in the 14-year-old stand to 0.37, 0.31, and 0.24 in the 31-, 46-, and 63-year-old stands, respectively. Age-specific biomass equations in each stand were developed for stem wood, stem bark, aboveground, and total tree. The inclusion of tree height as a second variable improved the total tree biomass equation fitting for middle-aged (31-year-old and 46-year-old) stands but not young (14 years old) and mature (63 years old) stands. Moreover, biomass conversion and expansion factors (BCEFs) varied with stand age, showing a decreasing trend with increasing stand age. These results indicate that stand age alters the biomass allocation of Q. acutissima and results in age-specific allometric biomass equations and BCEFs. Therefore, to obtain accurate estimates of Q. acutissima forest biomass and carbon stocks, age-specific changes need to be considered.

Aboveground and belowground biomass allocation in native Prosopis caldenia Burkart secondaries woodlands in the semi-arid Argentinean pampas

Honey mesquite (Prosopis glandulosa Torr.) which grows on grasslands and rangelands in southwestern USA may have potential as a bioenergy feedstock because of existing standing biomass and regrowth potential. However, regrowth mesquite physiognomy is highly different from undisturbed mesquite physiognomy and little is known regarding growth rates and structural biomass allocation in regrowth mesquite. We compared canopy architecture, aboveground biomass and relative allocation of biomass components in regrowth (RG) trees of different known ages with undisturbed (UD) trees of similar canopy height to each RG age class. RG trees in most age classes (2–12 years old) had greater canopy area, leaf area, basal stem number, twig (<0.5 cm diameter) mass and small stem (0.5–3 cm diameter) mass than UD trees of the same height. Large stem (>3 cm diameter) mass was similar between RG and UD trees in all height classes. Ages of UD trees were determined after harvest and further comparisons were made between age, canopy structure and biomass in RG and UD trees. Relationships between age and total mass, age and height, and age and canopy area indicated a faster growth rate in RG than in UD trees. Large stem mass as a percentage of total tree mass accumulated more rapidly with age in RG than UD trees. Leaf area index and leaf : twig mass ratio were maintained near 1 in all RG and UD trees. Regrowth potential may be one of the most important features of mesquite in consideration as a bioenergy feedstock.

The biomass and carbon (C) storage of the main ecosystem components were examined in an age-sequence of six Japanese red pine forest stands in central Korea. The tree biomass was determined by the destructive method, and the C storage of the tree biomass, forest floor and mineral soil was estimated by analyzing the C concentration of each component. The above-ground and total tree biomass increased from 21.76 and 28.82 Mg ha−1 in the 17-year-old stand to 308.83 and 385.74 Mg ha−1 in the 73-year-old stand. The comparisons of above-ground tree and tree root biomass in replicate stands indicated that stand density has an effect on tree biomass partitioning for Japanese red pine, especially on the biomass allocation of above-ground tree and tree roots. The C concentrations were lowest in the tree roots while the highest concentrations were found in the foliage across the six Japanese red pine forest stands. The C storage in the forest floor and mineral soil were age-independent. The above-ground and total ecosystem C stocks increased from 19.40 and 43.49 Mg ha−1 in the 17-year-old stand to 162.72 and 247.39 Mg ha−1 in the 73-year-old stand. Although the total tree biomass C showed considerable accumulation with stand age, the relative contribution of the below-ground ecosystem to the total ecosystem C storage demonstrated large variation. The results of this study assist in understanding C storage and its change with stand development in Japanese red pine forests, which makes this species a large sink for atmospheric C at the regional scale.

Eucalyptus camaldulensis × globulus and E.camaldulensis × grandis hybrids have been developed to combine the salt–waterlogging tolerance and high-quality wood fibre of their respective parents. The aim is to develop trees that will grow in relatively dry and/or saline environments and provide commercial wood products. Previous studies indicate that the hybrids exhibit faster growth than either of their pure species parents, and that there are significant differences in growth rates between them. We undertook a comparative study of the partitioning of above-ground biomass (AGB) to examine biomass and chloride (Cl) allocation of trees growing on two saline-irrigated sites in south-eastern Australia. Eucalyptus camaldulensis × globulus had a higher proportion of AGB in leaves (20–29% cf. 15–16%), and lower proportion in live branches (3–10% cf. 6–14%) than E.camaldulensis × grandis. The concentration of Cl was highest in the stembark (4.2–9.6 g kg−1) and lowest in the stemwood (0.6–2.0 g kg−1), suggesting that trees can export Cl through bark shedding. Total Cl content was strongly related to volume under bark (R2 = 0.99), and differences in partitioning of Cl into tree components differed between the hybrids in the same way as AGB. Preferential partitioning of Cl to live branches rather than foliage in E. camaldulensis × grandis suggests that this hybrid may be compartmentalising Cl to reduce the risk of Cl toxicity in the leaves.

Effect of spacing and water availability on root:shoot ratio in Eucalyptus camaldulensis

In the conditions of Central Europe, silver birch (Betula pendula Roth) has so far not been an important species for commercial purposes but might be relevant from ecological point of view. For instance, prompt succession by birch and other pioneer tree species at large-scale post-disturbance areas is necessary to compensate for previous carbon losses by natural disasters. Therefore, our attention was focused on 14-year-old birch trees growing at the wind-thrown area in the High Tatra Mts. (northern Slovakia). We sampled aboveground biomass of 20 silver birch trees representing four classes of crown competition: 0 – competition free crowns, 1 – crowns under mild competition, 2 – crowns under moderate competition, 3 – crowns under severe competition. We studied biomass allocated to stems, branches and foliage, and basic properties of stems and foliage. The crown-competition free birches were nearly 13 m high, and their aboveground tree biomass was 150 kg. The biomass of birches under severe competition was five times lower. Crown competition modified biomass ratios of foliage to branch as well as of branch to stem. Our results showed that birches under severe competition stress invest more in height than in diameter. At the same time, crown competition modified foliage weight and specific leaf area (SLA), which was clear mostly in the upper part of the crowns. However, foliage area was influenced by crown competition only to a negligible extent. Our main finding is that foliage position (upper, middle or lower third of crown) affected foliage properties more than intraspecific crown competition. Finally, we pointed out that silver birch is a rather productive species that is not ecologically demanding. Therefore, it might be a prospective tree species under the ongoing climate change and the present period of intensification in renewable resources utilisation.

The significant role of tropical forest ecosystems in the global carbon budget has increased the need for accurate estimates of tropical forest biomass. The lack of large-scale biomass allometric equations hampers the understanding of the spatial distribution of tree biomass and carbon stocks and their influencing factors in West Africa. This study aimed to develop allometric equations to estimate aboveground biomass of African oak (Afzelia africana Sm.) in Burkina Faso and to analyze factors affecting the variability of tree biomass and carbon storage. Sixty individual trees were destructively sampled in four protected areas along two climatic zones. In each climatic zone, log–log models were tested and fitted to each aboveground biomass component and to the total aboveground biomass. Carbon content in tree aboveground components was evaluated using the ash method. All validated equations showed good fit and performance with high explained variance. Allometric equations differed between the Sudano-sahelian zone and the Sudanian zone, except for leaf biomass equations. Both biomass allocation and carbon content varied significantly between tree components but not between climatic zones. Carbon content in tree components followed the patterns of biomass allocation with branches accounting for the highest proportion. In the two climatic zones, carbon contents were 50.18–52.62% for leaves, 54.78–54.94% for stems and 54.96–55.99% for branches. Dry biomass ranged from 509.05 to 765.56 kg tree−1 at site level and from 620.21 to 624.48 kg tree−1 along climatic zones. Carbon content varied from 53.90% in the Sudano-sahelian zone to 54.39% in the Sudanian zone. This study indicated that climate does not influence aboveground biomass production and carbon sequestration of Afzelia africana along the Sudano-sahelian and the Sudanian climatic zones of Burkina Faso. Future studies on climate–growth relationships should contribute to better understanding climate effects on biomass production and carbon storage.