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.

Tree biomass allocation differs by mycorrhizal association

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.

1 Herbivore damage can select for tolerance in plant populations where genetic variation for tolerance exists. The causes underlying variation in tolerance are not fully resolved. We assessed the importance of two potential mechanisms for tolerance by examining its relationship with leaf photosynthetic rate and relative biomass allocation across plant organs. 2 We monitored responses of 12 aspen (Populus tremuloides) genotypes, grown in a common garden under two levels of nutrient availability, to defoliation in two successive seasons. Tolerance of each genotype was calculated as the difference in growth between defoliated and undefoliated trees grown under the same nutrient conditions. 3 Although light-saturated leaf photosynthesis increased in response to nutrient addition and defoliation, it did not vary among genotypes and was not correlated with tolerance. 4 Tolerance was, however, correlated with patterns of biomass allocation. Under low-nutrient conditions it was positively correlated with the proportion of biomass in stems just prior to defoliation, while under high-nutrient conditions it was correlated with greater allocation to stems in response to damage. 5 Herbivores may select for specific patterns of biomass allocation in trees, and do so differently in different environments. The positive correlation between tolerance and relative allocation to stems, as opposed to roots, runs counter to reports from studies of herbaceous species and underscores the need for further exploration of mechanisms of tolerance in woody plants.

Allometry and partitioning of above- and belowground tree biomass in an age-sequence of white pine forests

Casuarina equisetifolia is commonly planted and used in the construction of coastal shelterbelt protection in Hainan Island. Thus, it is critical to accurately estimate the tree biomass of Casuarina equisetifolia L. for forest managers to evaluate the biomass stock in Hainan. The data for this work consisted of 72 trees, which were divided into three age groups: young forest, middle-aged forest, and mature forest. The proportion of biomass from the trunk significantly increased with age (P<0.05). However, the biomass of the branch and leaf decreased, and the biomass of the root did not change. To test whether the crown radius (CR) can improve biomass estimates of C. equisetifolia, we introduced CR into the biomass models. Here, six models were used to estimate the biomass of each component, including the trunk, the branch, the leaf, and the root. In each group, we selected one model among these six models for each component. The results showed that including the CR greatly improved the model performance and reduced the error, especially for the young and mature forests. In addition, to ensure biomass additivity, the selected equation for each component was fitted as a system of equations using seemingly unrelated regression (SUR). The SUR method not only gave efficient and accurate estimates but also achieved the logical additivity. The results in this study provide a robust estimation of tree biomass components and total biomass over three groups of C. equisetifolia.

Ponderosa pines (Pinus ponderosa Dougl. ex. Laws) 21 to 60 yr old were used to assess the relative importance of environmental stressors (O3, drought) versus an enhancer (N deposition) on foliar retention, components of aboveground growth, and whole tree biomass allocation. Sites were chosen across a well-described gradient in ozone exposure (40 to 80 ppb per h, 24 h basis, 6 month growing season) and nitrogen deposition (5 to 40 kg ha-1 yr-1) in the San Bernardino Mountains east of Los Angeles, California. A high level of chlorotic mottle indicated high O3 injury at sites closest to the pollution source, despite potential for the mitigating effects of N deposition. At the least polluted site, foliar biomass was evenly distributed across three of the five needle-age classes retained. At the most polluted site, 95% of the foliar biomass was found in the current year's growth. High N deposition and O3 exposure combined to shift biomass allocation in pine to that of a deciduous tree with one overwintering needle age class. Based on whole tree harvests, root biomass was lowest at sites with the highest pollution exposure, confirming previous chamber exposure and field studies. Aboveground growth responses in the high-pollution sites were opposite to those expected for O3 injury. Needle and lateral branch elongation growth, and measures of wood production increased with increasing proximity to the pollution source. An enhancement of these growth attributes suggested that N deposition dominated the ponderosa pine response despite high O3 exposure.

Inter-annual climatically driven growth variability of above-ground biomass compartments (for example, tree stems and foliage) controls the intensity of carbon sequestration into forest ecosystems. However, understanding the differences between the climatic response of stem and foliage at the landscape level is limited. In this study, we examined the climate-growth response of stem and leaf biomass and their relationship for Pinus sylvestris (PISY) and Picea abies (PCAB) in topographically complex landscapes. We used tree-ring width chronologies and time series of the normalized difference vegetation index (NDVI) derived from high-resolution Landsat scenes as proxies for stem and leaf biomass, respectively. We then compared growth variability and climate-growth relationships of both biomass proxies between topographical categories. Our results show that the responses of tree rings to climate differ significantly from those found in NDVI, with the stronger climatic signal observed in tree rings. Topography had distinct but species-specific effects: At moisture-limited PISY stands, stem biomass (tree rings) was strongly topographically driven, and leaf biomass (NDVI) was relatively insensitive to topographic variability. In landscapes close to the climatic optimum of PCAB, the relationship between stem and leaf biomass was weak, and their correlations with climate were often inverse, with no significant effects of topography. Different climatic signals from NDVI and tree rings suggest that the response of canopy and stem growth to climate change might be decoupled. Furthermore, our results hint toward different prioritizations of biomass allocation in trees under stressful conditions which might change allometric relationships between individual tree compartments in the long term.

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.

Applying allometric equations in combination with forest inventory data is an effective approach to use when qualifying forest biomass and carbon storage on a regional scale. The objectives of this study were to (1) develop general allometric tree component biomass equations and (2) investigate tree biomass allocation patterns for Pinus massoniana, a principal tree species native to southern China, by applying 197 samples across 20 site locations. The additive allometric equations utilized to compute stem, branch, needle, root, aboveground, and total tree biomass were developed by nonlinear seemingly unrelated regression. Results show that the relative proportion of stem biomass to tree biomass increased while the contribution of canopy biomass to tree biomass decreased as trees continued to grow through time. Total root biomass was a large biomass pool in itself, and its relative proportion to tree biomass exhibited a slight increase with tree growth. Although equations employing stem diameter at breast height (dbh) alone as a predictor could accurately predict stem, aboveground, root, and total tree biomass, they were poorly fitted to predict the canopy biomass component. The inclusion of the tree height (H) variable either slightly improved or did not in any way increase model fitness. Validation results demonstrate that these equations are suitable to estimate stem, aboveground, and total tree biomass across a broad range of P. massoniana stands on a regional scale.

Knowledge of tree biomass allocation is fundamental for estimating forest acclimation and carbon stock for global changes in the future. Optimal partitioning theory (OPT) and allometric partitioning theory (APT) are two major patterns of biomass allocation, and occurrences have been tested on taxonomical, ontogenetic, geographic and environmental scales, showing conflicting results and unclear ecophysiological mechanisms. Here, we examine the biomass allocation patterns of two young poplar (Populus) clones varying greatly in drought resistance under different soil water and nitrogen availabilities and the major physiological processes involved in biomass partitioning. We found that Biyu, a drought-sensitive hybrid poplar clone, had significant relations among biomass of leaf, stem and root, showing allometric partitioning. Xiaoye, a drought-tolerant poplar clone native to semi-arid areas, on the contrary, showed tightly regulated biomass allocation following optimal partitioning theory. Biyu had higher nitrate reductase activity in the fine roots, while Xiaoye had higher nitrate reductase activity in the leaves. Biochemical analyses and measurements of fluorescence and gas exchange showed that Xiaoye maintained more stable chloroplast membranes and photosystem electron flow, showing higher water use efficiency and a higher resistance to drought. A nitrogen addition could improve leaf photosynthesis and growth both in Biyu and Xiaoye seedlings under drought conditions. We concluded that the two poplar clones showed different biomass allocation patterns and suggest that the site of nitrate assimilation may play a role in biomass partitioning under varying water and nitrogen availabilities.

Understanding the determinants of tree biomass allocation patterns among organs is crucial for both predicting the rate and potential of forest carbon sinks and guiding future multifunctional forest management. However, it is still not clear how the site conditions (e.g., elevation) and stand structure (e.g., tree dominance, stand density) affect the biomass allocation of single trees in forests. This study was implemented in the Liupan Mountains of the Loess Plateau of Northwest China by collecting the related information of biomass data of 110 sample trees with different dominance and influencing factors within 23 sample plots of larch plantations set up along the elevation gradient. Based on these data, the response tendency and functions of biomass allocation of single trees to individual influencing factors of site conditions and forest structure were analyzed. Moreover, the results illustrated that the ratio between root biomass and aboveground biomass decreased significantly with rising stand age and tree density, but increased significantly with rising elevation, and there was no significant relationship with the dominance of individual trees. The results of this study revealed the importance of considering the influencing factors of site conditions and stand structure when developing dynamic models of tree biomass allocation. The results and research methods used in this study provide useful tools for quantifying the biomass allocation and carbon storage partitioning in the study area and other similar regions.

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.

Factorial combinations of soil compaction and organic matter removal were replicated at the Long Term Site Productivity study in the Croatan National Forest, near New Bern, North Carolina, USA. Ten years after planting, 18 preselected loblolly pine ( Pinus taeda L.) trees were destructively harvested to quantify treatment effects on total above- and below-ground tree biomass and to detect any changes in the absolute and relative allocation patterns. Stem volume at year 10 was not affected by compaction treatments, even though the ultisols on these sites continued to have higher bulk densities than noncompacted plots. However, even when site preparation treatments were undetectable aboveground, the treatments significantly altered absolute root growth and tree biomass allocation patterns. Soil compaction decreased taproot production and significantly increased the ratio of aboveground to belowground biomass. Decreased root production will decrease carbon and nutrient stores belowground, which may impact future site productivity.