Modelling biomass and leaf area index in a sub-arctic Scandinavian mountain area

Estimates of biomass and leaf area index (LAI) are important variables in ecological and climate models. However, very little is known about the biomass and LAI of the vegetation in the Scandinavian mountain area. In this study, extensive field data consisting of diameter at breast height for 13?000 trees and height for 550 trees were collected. Furthermore, biomass and leaf area (LA) measurements for 46 mountain birch trees [Betula pubescens ssp. czerepanovii (Orlowa) Hämet-Ahti] and biomass and LA measurements for shrubs (e.g. Salix spp., Betula nana) at 36 sample plots were carried out. Multiplicative linear models for trees were fitted to tree biomass and LA measurements using basal area at breast height, height, crown diameter and diameter at stump height as explanatory variables. Additive linear models were fitted to shrub biomass and LAI measurements using coverage of shrubs, topographic variables and soil type as explanatory variables. The functions were then used to predict the biomass and LAI for trees and shrubs for the entire test area, which covers an area of 84 km² and is located at latitude 68° N. The mean total biomass estimates were 27?493 kg ha^?1 for the forest and 7650 kg ha^?1 for snow-protected heath and meadow vegetation. The LAIs were 2.06 and 0.52, respectively. For monitoring biomass and LAI in the Scandinavian mountain area, the functions could also be applied to data from traditional field-based inventories and the estimates might further be improved by combining the estimates from the test area with auxiliary information such as remote sensing images.     

The chemical composition of needle and leaf litter from Scots pine, Norway spruce and white birch in Scandinavian forests

Needle litter from 14 stands of Scots pine (Pinus silvestris,L.), 13 stands of Norway spruce (Picea abies (L.) Karst.) andleaf litter from three stands of white birch (Betula pubescensEhrh.) were analysed for chemical composition. The concentrationsof the elements N, P, K, Ca, Mg and Mn as well as solid organiccomponents (lignin, cellulose and hemicelluloses) and solubleswere determined. When the average chemical compositions werecompared the Scots pine needle litter was clearly the most nutrient-poorlitter type. Of the solid organic-chemical components the ligninfraction dominated in the spruce and birch litter whereas thecellulose dominated in the pine needle litter. When Norway spruce and Scots pine were growing in adjacent standson soils with the same bedrock origin the spruce litter hadsignificantly higher concentrations of nutrients (N, P, K, Ca,Mg, Mn) than the pine needle litter. At sites where Norway spruceand white birch were growing in adjacent stands, the birch leaflitter had generally higher concentrations of nutrients. However, significant or nearly significant differences were onlyobtained for Mg (P = 0.002), K (P = 0.056) and N (P = 0.087),probably due to the few replicates of stands compared. Concerningorganic chemical components, the spruce needle litter had significantlyhigher concentrations of lignin and mannan than all the otherlitters and lower levels of ethanol-soluble substances, celluloseand galactan than the pine needle litter. Further, it had lowerconcentrations of water solubles, rhamnan and xylan than thebirch litter. No relationships were established between the nutrient statusof the conifer litters and the site index H100 (the dominantheight of the trees at a reference age of 100 years) of thestands. Concentrations of solid carbohydrates in the litterswere, however, positively correlated with site index (P <0.001). Further, the concentration of nitrogen in the pine needlelitter was negatively correlated with the latitude of the sites(P < 0.01). The influence of litter chemistry on the decompositionof litter and nutrient cycling of forests is discussed.     

Formulating allometric equations for estimating biomass and carbon stock in small diameter trees

Biomass and carbon sequestration rate of a young (four year old) mixed plantation of Dalbergia sissoo Roxb., Acacia catechu Willd., and Albizia lebbeck Benth. growing in Terai region (a level area of superabundant water) of central Himalaya was estimated. The plantation is seed sown in the rainy season of year 2004 and spread over an area of 44 ha. Allometric equations for both above and below ground components were developed for three tree species. The density of trees in the plantation was 1322 trees ha⁻¹ The diameters of trees were below 10 cm. Five diameter classes were defined for D. sissoo and A. catechu and 3 for A. lebbeck. 5 trees were harvested in each diameter class. Individual tree allometry was exercised for developing the allometric equations relating tree component (low and above ground) biomass to d.b.h. Post analysis equations were highly significant (P > 0.001) for each component of all species. In the plantation Holoptelia integrifolia Roxb. (Family Ulmaceae) has been reduced to shrub form because of frost. Only the aboveground biomass of H. integrifolia and other shrubs were estimated by destructive harvesting method. Herbaceous forest floor biomass and leaf litter fall were also estimated. The total forest vegetation biomass was 10.86 Mg ha⁻¹ in 2008 which increased to 19.49 Mg ha⁻¹ in 2009. The forest is sequestering carbon at the rate of 4.32 Mg ha⁻¹ yr⁻¹.     

Temporal variation and efficiency of leaf area index in young mountain Norway spruce stand

Temporal variation of leaf area index (LAI) in two young Norway spruce stands with different densities was monitored during eight consecutive growing seasons (1998–2005). We focused on: (1) LAI dynamics and above-ground mass production of both spruce stands and their comparison, (2) leaf area duration (LADU), crop production index (CPI) and leaf area efficiency (LAE) evaluation, and (3) thinning impact on the above-mentioned parameters. Also, we tried to deduce the most effective LAI value for the Norway spruce forest investigated. The LAI values of both spruce stands showed a typical seasonal course. To describe the LAI dynamics of the stand, we recommend taking LAI measurements within short time intervals at the time of budding and needle expansion growth (i.e., in early spring) and close to the LAI peak, when the twig growth has been completed. The reason was that after reaching the seasonal maximum, no significant differences between subsequently obtained values were found in the following 2 months. Therefore, we recommend this period for the estimation of seasonally representative LAI values, enabling the comparison of various spruce stands. The maximum hemi-surface LAI value reached 12.4. Based on our results, the most effective LAI values for maximum above-ground biomass production were within the range of 10–11. We found an LAI over these values to be less effective for additional production of above-ground biomass. In forest practice, thinning intensity is mostly described by percentage of stocking reduction. We want to show that not only thinning intensity, but also the type of thinning is important information. The type of thinning significantly affected the stand above-ground biomass increment, canopy openness, stand LAI and LAI efficiency. The stimulating effect of high-type thinning was observed; the LAE as well as the CPI increased. Low-type thinning had no such effects on LAE increments compared to the high-type thinning with similar intensity.

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Interception of precipitation by pole-stage Sitka spruce and lodgepole pine and mature Sitka spruce at Kielder Forest, Northumberland

Interception loss was measured indirectly for 3 years in adjacentunthinned 25-year-old stands of Sitka spruce and lodgepole pineand for 2 years in a 63-year-old selectively thinned stand ofSitka spruce. For each stand measurements were made of grossprecipitation, throughfall and stemflow. Interception loss wasderived by subtracting the sum of throughfall and stemflow (netprecipitation) from gross precipitation. For the years 1977–78, 1978–79 and 1979–80,interception losses in the pole-stage Sitka spruce were 32,28 and 27 per cent of gross precipitation respectively whilethose in the lodgepole pine were 33, 29 and 26 per cent. For1979–80 and 1980–81 interception losses in the matureSitka spruce were 44 and 53 per cent of gross precipitation.The average interception loss was 29 per cent for both pole-stagecrops and 49 per cent for mature Sitka spruce. The proportions of net precipitation reaching the ground asstemflow and throughfall were 0.18 and 0.82 respectively forthe pole-stage Sitka spruce, 0.14 and 0.86 for the pole-stagelodepole pine and 0.02 and 0.98 for the mature Sitka spruce.     

Validation of an efficient visual method for estimating leaf area index in clonal Eucalyptus plantations

Leaf area index (LAI) is a key ecophysiological parameter in forest stands because it characterises the interface between atmospheric processes and plant physiology. Several indirect methods for estimating LAI have been developed. However, these methods have limitations that can affect the estimates. This study aimed to evaluate the accuracy and applicability of a visual method for estimating LAI in clonal Eucalyptus grandis × E. urophylla plantations and to compare it with hemispherical photography, ceptometer and LAI-2000^® estimates. Destructive sampling for direct determination of the actual LAI was performed in 22 plots at two geographical locations in Brazil. Actual LAI values were then used to develop a field guide with photographic images representing an LAI range of 1.0–5.0 m² m^?2 (leaf area/ground area). The visual LAI estimation guide was evaluated with 17 observers in the field. The average difference between actual LAI and visual LAI estimation was 12% and the absolute difference between the two methods was less than or equal to 0.5 m² m^?2 in 77% of plots. Pearson’s correlation coefficients were high between actual LAI and hemispherical photographs (0.8), visual estimation (0.93) and LAI-2000^® (0.99) and low for the ceptometer (0.18). However, absolute values differed among methods, with the average difference between the actual and estimated LAI of [12]% for visual estimation, 28% for the LAI-2000^®, 37% for the ceptometer and ?43% for hemispherical photographs. The LAI-2000^® and ceptometer overestimated LAI in all plots, whereas hemispherical photographs underestimated the values in all measurements, showing that these methods need calibration to be used. No differences were observed between actual LAI and visual estimates across stand ages of 2–8 years and LAI of 1.5–5.3 m² m^?2 (P > 0.05). The results show that visual estimation of LAI in Eucalyptus stands is a practical method that is unaffected by atmospheric characteristics and can be used on an operational scale.     

区域尺度杉木生物量通用相对生长方程整合分析

对已发表的我国南方不同地点的杉木林生物量数据和方程进行整合分析(Meta-analysis),建立适合大尺度的杉木生物量估算通用相对生长方程。结果表明:通用相对生长方程拟合杉木树干、地上部分和全株生物量数据程度最高(相关系数R20.92),根生物量的拟合较好(R20.87),叶和枝的拟合较差(R20.78)。仅用胸径(D)作为自变量与引入胸径、树高因子(D2、H)相比,拟合的相对生长方程的相关系数相差不大。估算的各组份生物量叠加后,与地上部分和全株树木生物量方程估算值的相容性较好,其相关系数在0.99以上。方程检验结果与发表的全株生物量方程比较表明,整合分析数据拟合的通用相对生长方程能满足估算大尺度杉木地上部分和全株生物量的精度要求,可以用来估算区域杉木林的生物量和碳贮量。     

Species-specific biomass allometric models and expansion factors for indigenous and planted forests of the Mozambique highlands

Secondary Miombo woodlands and forest plantations occupy increasing areas in Mozambique, the former due to anthropogenic activities. Plantations, mainly species of Eucalyptus and Pinus, are being established on sites previously covered by secondary Miombo woodlands. This affects the evolution, cycle and spatiotemporal patterns of carbon(C) storage and stocks in forest ecosystems. The estimation of C storage, which is indispensable for formulating climate change policies on sequestrating CO_2, requires tools such as biomass models and biomass conversion and expansion factors(BCEF). In Mozambique, these tools are needed for both indigenous forests and plantations. The objective of this study is to fit species-specific allometric biomass models and BCEF for exotic and indigenous tree species. To incorporate efficient inter-species variability, biomass equations were fitted using nonlinear mixed-effects models. All tree component biomass models had good predictability; however, better predictive accuracy and ability was observed for the 2-predictors biomass model with tree height as a second predictor. The majority of the variability in BCEF was explained by the variation in tree species. Miombo species had larger crown biomass per unit of stem diameter and stored larger amounts of biomass per stem volume. However, due to relatively rapid growth, larger stem diameters, heights, and stand density, the plantations stored more biomass per tree and per unit area.     

Optical and litter collection methods for measuring leaf area index in an old-growth temperate forest in northeastern China

Hemispherical photographs combined with litter collection were applied to determine seasonal dynamics of leaf area index (LAI) between the period of maximum leaf area and the leafless period from an old-growth temperate forest in the Xiaoxing’an Mountains, northeastern China. Our objective is to explore the change in the relationship between “true” LAI and effective LAI (calculated only from hemispherical photography) and to find the best LAI estimation models. Effective LAI in November is corrected for contribution of woody material and clumping at shoot and beyond shoot levels, to give minimum “true” LAI. The “true” LAI in each period is estimated as a sum of the minimum “true” LAI and litter collection LAI in each period. Power function regression calibration models were then carried out between “true” LAI and effective LAI in each period and the entire litter-fall period. Then, significance tests were applied to detect the differences among different models. The results showed that the average “true” LAI ranged from 2.74 ± 0.54 on November 1 to 6.64 ± 1.34 on July 1. For the entire season, average effective LAI was 53.16 % lower than the average “true” LAI. After significance tests, calibration models were classified into two types: (1) maximum LAI period and the period of maximum leaf fall; (2) the period during which leaves began falling and all deciduous leaves had fallen. Based on our experience, we believe that the classified models can produce reliable and accurate LA1 values for the needle and broad-leaved mixed forest stands under the non-destructive condition.     

Comparison of four methods for estimating leaf area index based on terrestrial three-dimensional laser scanning

The leaf area index (LAI) of 16 sample plots was estimated based on terrestrial three-dimensional laser scanning. The point-cloud data of stand canopy were first scaled and projected onto a hemisphere according to Lambert azimuthal equal-area projection or stereographic projection, and the resulting hemispherical point-cloud images were used to extract the canopy porosity coefficients. Then, single-angle inversion and Miller formula inversion methods were used, respectively, to calculate the effective leaf area indices with canopy porosity coefficients. Results showed that the effective LAIs estimated by single-angle inversion method with Lambert projection and stereographic projection were within the range of 2.14~5.36 and 1.83~4.67, respectively. The effective LAIs obtained by Miller formula inversion method with Lambert projection and stereographic projection were within the range of 1.84~4.67 and 1.68~4.34, respectively. As a comparison, the LAI measured with a fish-eye camera ranged from 1.55 to 3.87. The LAI values estimated with four different calculation methods were linearly correlated with those measured by a fish-eye camera. The highest coefficient of determination (R²) 90.28% was obtained by the Miller formula inversion method combined with stereographic projection, and Duncan’s new multiple range test also further showed that this method had a relatively higher precision compared to other three methods.     

Managing forests infested by spruce beetles in south-central Alaska: Effects on nitrogen availability,understory biomass,and spruce regeneration

In Alaska, an outbreak of spruce beetles (Dendroctonus rufipennis) recently infested over one million hectares of spruce (Picea spp.) forest. As a result, land management agencies have applied different treatments to infested forests to minimize fire hazard and economic loss and facilitate forest regeneration. In this study we investigated the effects of high-intensity burning, whole-tree harvest, whole-tree harvest with nitrogen (N) fertilization, and conventional harvest of beetle-killed stands 4 years after treatment, as well as clear-cut salvage harvest 6 years after treatment. We measured available soil ammonium and nitrate and estimated N loss from leaching using in situ cation and anion resin exchange capsules. We also assessed spruce regeneration and responses of understory plant species. Availability and losses of N did not differ among any of the management treatments. Even a substantial application of N fertilizer had no effect on N availability. Spruce regeneration significantly increased after high-intensity prescribed burning, with the number of seedlings averaging 8.9 m⁻² in burn plots, as compared to 0.1 m⁻² in plots that did not receive treatment. Biomass of the pervasive grass bluejoint (Calamagrostis canadensis) was significantly reduced by burning, with burn plots having 9.5% of the C. canadensis biomass of plots that did not receive treatment. N fertilization doubled C. canadensis biomass, suggesting that N fertilization without accompanying measures to control C. canadensis is the least viable method for promoting rapid spruce regeneration.     

An allometric equation for estimating stem biomass of Acacia auriculiformis in the north-eastern region of Bangladesh

Tree biomass plays an important role in sustainable management and in estimating forest carbon stocks. The objective of this study was to select the best model for measuring stem biomass of Acacia auriculiformis in the study area. Data from five hillocks and 120 individual trees from each hillock were used in this study. Twelve different forms of linear, power and exponential equations were compared in this study to select the best model. Two models (VI and XI) were selected based on R ², adjusted R ², the Akaike information criterion, F-statistics and the five assumptions of linear regression. Model VI was discarded based on the Durbin-Watson value of autocorrelation of the residuals, then the ARIMA (2, 0, 1) model was used to remove the autocorrelation from the model and the final bias-corrected model XI was derived. The model was validated with a test data set having the same range of DBH and stem height of the training data set on the basis of linear regression, Morisita's similarity index, and t-test for mean difference between predicted and expected biomass. A comparison between the best logarithmic and non-linear allometric model shows that the non-linear model produces systematic biases and overestimates stem biomass for larger trees. The overall results showed that the bias-corrected logarithmic model XI can be used efficiently for estimating stem biomass of A. auriculiformis in the northeastern region of Bangladesh.     

Stocks and decay dynamics of above- and belowground coarse woody debris in managed Sitka spruce forests in Ireland

Coarse woody debris (CWD) has become recognised as an important component of the carbon (C) pool in forest ecosystems. In Ireland, managed Sitka spruce (Picea sitchensis (Bong) Carr.) forests account for 52.3% of the total forest estate. To determine the stock and decay dynamics of above and belowground CWD, field surveys using fixed area sample plots, were conducted in six even-aged Sitka spruce stands, representing the young, intermediate and mature stages of a typical commercial rotation. The volume, mass, density loss and C:N ratio of all CWD types (logs, stumps, and coarse roots) were determined using a five-decay class (DC) system. The decay rates and half life of CWD was also determined. To estimate CWD coarse root mass; roots associated with stumps classified in different decay classes were excavated. The coarse roots were categorised into small (2-10 mm), medium (10-50 mm) and large (>50 mm) diameter classes.CWD C-mass ranged from 6.98 to 18.62 Mg ha⁻¹ and was highest in an intermediate forest (D35), while the aboveground volume varied from 6.31 to 42.27 m³ ha⁻¹. Coarse roots accounted for 21% to 85% of the total CWD C-pool in the surveyed stands. The total CWD C-mass was poorly correlated with the number of thinning events (R² = 0.29), when data from D35 was excluded. The density loss was significant in logs (45%), stumps (58%), and small- (38%), medium- (50%) and large roots (38%) as decay progress from DC 0 to 4. There was a 46%, 41%, 51%, 72% and 57% decline in C:N ratio of logs, stumps, small-, medium- and large roots, respectively, as decay progressed from DC 0 to 4. The density decay rates were 0.059, 0.048 and 0.036 kg m⁻³ year⁻¹ for logs, stumps and coarse roots, respectively. The size classification of roots did not significantly affect their decay rate. The half life (50% decomposition) of CWD was estimated has 12-, 14- and 19 years for logs, stumps and roots of Sitka spruce. Regression curves showed a strong correlation between the density and C:N ratio (R² = 0.69, 0.74 and 0.93 for logs, stumps and coarse roots, respectively). The long term storage of C and its slow rate of decomposition make CWD a vital structural and functional component of the CWD C-pool and a major controller of forest ecosystem C-retention.     

Estimating carbon stock in secondary forests: Decisions and uncertainties associated with allometric biomass models

Secondary forests are a major terrestrial carbon sink and reliable estimates of their carbon stocks are pivotal for understanding the global carbon balance and initiatives to mitigate CO₂ emissions through forest management and reforestation. A common method to quantify carbon stocks in forests is the use of allometric regression models to convert forest inventory data to estimates of aboveground biomass (AGB). The use of allometric models implies decisions on the selection of extant models or the development of a local model, the predictor variables included in the selected model, and the number of trees and species for destructive biomass measurements. We assess uncertainties associated with these decisions using data from 94 secondary forest plots in central Panama and 244 harvested trees belonging to 26 locally abundant species. AGB estimates from species-specific models were used to assess relative errors of estimates from multispecies models. To reduce uncertainty in the estimation of plot AGB, including wood specific gravity (WSG) in the model was more important than the number of trees used for model fitting. However, decreasing the number of trees increased uncertainty of landscape-level AGB estimates substantially, while including WSG had limited effects on the accuracy of the landscape-level estimates. Predictions of stand and landscape AGB varied strongly among models, making model choice an important source of uncertainty. Local models provided more accurate AGB estimates than foreign models, but high variability in carbon stocks across the landscape implies that developing local models is only justified when landscape sampling is sufficiently intensive.     

Estimating leaf area index of mature temperate forests using regressions on site and vegetation data

Canopy gap fraction and leaf area index (LAI) were measured using hemispherical photography in 91 mature forests across Switzerland, including coniferous, broadleaved and mixed stands. The gap fraction and LAI derived from five photographs per site could be reproduced with a high coefficient of determination (R² > 0.7) by regression against simple stand parameters obtained from vegetation surveys: coverages of the tree, shrub and herb layers, and tree height. The method appeared to be robust across the different types of forests. Applied to 981 sites across Switzerland, the regression model produced LAI values ranging from 1.4 to 6.7. These predictions were compared with site variables not included in the regression. LAI appeared limited by the altitude, with maximal values decreasing by one third from 400 to 2000 m above see level. Water availability was also clearly a limitation at sites with a negative water balance, i.e. where the yearly potential evapotranspiration exceeded the precipitation. High or low values of a humidity index based on the ground vegetation also corresponded to a limitation of the LAI, with shorter trees at dry sites and more open canopies at wet sites. Compared to optical measurements (including hemispherical photography), our regression method is fast and inexpensive. Such an approach appears very promising for obtaining reliable estimates of LAI for many sites with low costs. These estimates can then be fed into process models at the stand level.     

Integrating regional climate change into allometric equations for estimating tree aboveground biomass of Masson pine in China

Key message

A climate-sensitive aboveground biomass (AGB) equation, in combination with nonlinear mixed-effects modeling and dummy variable approach, was developed to examine how climate change may affect the allometric relationships between tree diameter and biomass. We showed that such changes in allometry need to be taken into account for estimating tree AGB in Masson pine.

Context

As a native species and being widely distributed in subtropical China, Masson pine (Pinus massoniana Lamb.) forests play a pivotal role in maintaining forest ecosystem functions and mitigation of carbon concentration increase at the atmosphere. Traditional biomass allometric equations do not account for a potential effect of climate on the diameter–biomass relationships. The amplitude of such an effect remains poorly documented.

Aims

We presented a novel method for detecting the long-term (2041–2080) effects of climate change on the diameter–biomass relationships and the potential consequences for long-term changes of biomass accumulation for Masson pine.

Methods

Our approach was based on a climate-sensitive AGB model developed using a combined nonlinear mixed-effects model and dummy variable approach. Various climate-related variables were evaluated for their contributions to model improvement. Heteroscedasticity was accounted for by three residual variance functions: exponential function, power function, and constant plus function.

Results

The results showed that diameter at breast height, together with the long-term average of growing season temperature, total growing season precipitation, mean temperature of wettest quarter, and precipitation of wettest quarter, had significant effects on values of AGB. Excessive rain during the growing season and high mean temperature in the wettest quarter reduced the AGB, while a warm growing season and abundant precipitation in the wettest quarter increased the AGB.

Conclusion

Climate change significantly affected the allometric scale of biomass equation. The new climate-sensitive allometric model developed in this study may improve biomass predictions compared with the traditional model without climate effects. Our findings suggested that the AGB of Masson pine trees with the same diameter at breast height under three climate scenarios including representative concentration pathway (RCP) 2.6, RCP 4.5, and RCP 8.5 in the future period 2041–2080 would increase by 24.8 ± 32.7% (mean ± standard deviation), 27.0 ± 33.4%, and 27.7 ± 33.8% compared with the constant climate (1950–2000), respectively. As a consequence, we may expect a significant regional variability and uncertainty in biomass estimates under climate change.

     

Comparison of allometric equations and biomass expansion factors for six evergreen broad-leaved trees in subtropical forests in southern Korea

This study was conducted to compare the allometric equations and biomass expansion factors (BEFs) of six dominant evergreen broad-leaved trees (Camellia japonica L, Castanopsis sieboldii Hatus, Quercus acuta Thunb, Q. glauca Thunb, Machilus thunbergii S. et Z., and Neolitsea sericea Koidz) in subtropical forests. A total of 86 trees were destructively sampled to quantify the aboveground biomass of each tree component (i.e., leaves, branches, and stem). Species-specific or generalized allometric equations and species-dependent BEFs were developed for each tree component of the six broad-leaved forest trees. Species-specific allometric equations were significant (P < 0.05), with the diameter at breast height (DBH) accounting for 68–99% of the variation, whereas generalized allometric equations explained 64–96% of the variation. The values of stem density ranged broadly from 0.49 g cm^?3 for C. sieboldii to 0.79 g cm^?3 for Q. glauca, with a mean value of 0.68 g cm^?3. The BEFs were significantly (P < 0.05) lower for C. sieboldii (1.25) than for M. thunbergii (2.02). Stem density and aboveground BEFs had a significant negative relationship with tree ages. The results indicate that species-specific allometric equations and species-dependent BEFs are applicable for obtaining accurate biomass estimates of subtropical evergreen broad-leaved forests.     

Internal cycling of nitrogen, potassium and magnesium in young Sitka spruce

Potassium (K) and magnesium (Mg) are essential macro-nutrients, but little is known about how they are cycled within plants. Stable isotope studies have shown that the internal cycling of nitrogen (N) is independent of current nutrient supply in temperate tree species. This is ecologically significant because it allows trees to produce rapid shoot growth in spring independent of current soil N uptake. We used stable isotopes to quantify N, K and Mg in new shoots of Sitka spruce (Picea sitchensis (Bong.) Carr.) seedlings and to compare the relative contributions from current uptake and internal cycling. Two-year-old Sitka spruce seedlings were labeled with (15)N, (41)K and (26)Mg in an abundant or a limited supply for one growing season. The trees were repotted in the subsequent dormant season to prevent further root uptake of enriched isotopes and provided with an abundant or a limited supply of unlabeled nutrients until they were harvested in early summer of the following year. The supply was switched for half the trees in the second year to create four nutrient regimes. Enrichment of (15)N, (41)K and (26)Mg in current-year growth was attributed to internally cycled N, K and Mg uptake from the previous year. The internal cycling of N, K and Mg in new growth was significantly affected by the first-year nutrient treatments. The second-year nutrient supply affected the growth rates of the trees, but had no effect on the amounts of N, K or Mg contributed from internal cycling. Thus, internal cycling of K and Mg in Sitka spruce are, like that of N, independent of current nutrient supply.     

Spatial and temporal variations in leaf area index, specific leaf area and leaf nitrogen of two co-occurring savanna tree species

Foliage growth, mass- and area-based leaf nitrogen concentrations (Nm and N a) and specific leaf area (SLA) were surveyed during a complete vegetation cycle for two co-occurring savanna tree species: Crossopteryx febrifuga (Afzel. ex G. Don) Benth. and Cussonia arborea A. Rich. The study was conducted in the natural reserve of Lamto, Ivory Coast, on isolated and clumped trees. Leaf flush occurred before the beginning of the rainy season. Maximum leaf area index (LAI), computed on a projected canopy basis for individual trees, was similar (mean of about 4) for both species. Seasonal courses of the ratio of actual to maximum LAI were similar for individuals of the same species, but differed between species. For C. febrifuga, clumped trees reached their maximum LAI before isolated trees. The LAI of C. arborea trees did not differ between clumped and isolated individuals, but maximum LAI was reached about 2 months later than for C. febrifuga. Leaf fall was associated with decreasing soil water content for C. arborea. For C. febrifuga, leaf fall started before the end of the rainy period and was independent of changes in soil water content. These features lead to a partial niche separation in time for light resource acquisition between the two species. Although Nm, N a and SLA decreased with time, SLA and N a decreased later in the vegetation cycle for C. arborea than for C. febrifuga. For both species, N a decreased and SLA increased with decreasing leaf irradiance within the canopy, although effects of light on leaf characteristics did not differ between isolated and clumped trees. Given relationships between N a and photosynthetic capacities previously reported for these species, our results show that C. arborea exhibits higher photosynthetic capacity than C. febrifuga during most of the vegetation cycle and at all irradiances.