Estimation of wetland vegetation height and leaf area index using airborne laser scanning data

Wetland vegetation is a core component of wetland ecosystems. Wetland vegetation structural parameters, such as height and leaf area index (LAI) are important variables required by earth-system and ecosystem models. Therefore, rapid, accurate, objective and quantitative estimations of wetland vegetation structural parameters are essential. The airborne laser scanning (also called LiDAR) is an active remote sensing technology and can provide accurate vertical vegetation structural parameters, but its accuracy is limited by short, dense vegetation canopies that are typical of wetland environments. The objective of this research is to explore the potential of estimating height and LAI for short wetland vegetation using airborne discrete-return LiDAR data.The accuracies of raw laser points and LiDAR-derived digital elevation models (DEM) data were assessed using field GPS measured ground elevations. The results demonstrated very high accuracy of 0.09 m in raw laser points and the root mean squared error (RMSE) of the LiDAR-derived DEM was 0.15 m.Vegetation canopy height was estimated from LiDAR data using a canopy height model (CHM) and regression analysis between field-measured vegetation heights and the standard deviation (σ) of detrended LiDAR heights. The results showed that the actual height of short wetland vegetation could not be accurately estimated using the raster CHM vegetation height. However, a strong relationship was observed between the σ and the field-measured height of short wetland vegetation and the highest correlation occurred (R² = 0.85, RMSE = 0.14 m) when sample radius was 1.50 m. The accuracy assessment of the best-constructed vegetation height prediction model was conducted using 25 samples that were not used in the regression analysis and the results indicated that the model was reliable and accurate (R² = 0.84, RMSE = 0.14 m).Wetland vegetation LAI was estimated using laser penetration index (LPI) and LiDAR-predicted vegetation height. The results showed that the vegetation height-based predictive model (R² = 0.79) was more accurate than the LPI-based model (the highest R² was 0.70). Moreover, the LAI predictive model based on vegetation height was validated using the leave-one-out cross-validation method and the results showed that the LAI predictive model had a good generalization capability. Overall, the results from this study indicate that LiDAR has a great potential to estimate plant height and LAI for short wetland vegetation.     

Environmental and stomatal control of transpiration, canopy conductance and decoupling coefficient in young lemon trees under shading net

Sap flow, as a measure of transpiration, was monitored in 2-year-old lemon trees growing in pots. Eight trees were used in the experiment, four of which were placed under a rectangular shading net, while the other four were maintained in the open air. Daily averages of canopy conductance and photosynthesis were not affected by shading; however, the daily transpiration was reduced in shaded plants, which displayed an increase in water use efficiency compared with exposed trees. The decoupling coefficient was higher in the shaded trees, indicating that the transpiration of lemon trees was efficiently controlled by stomata in exposed plants, while the transpiration rate was mainly influenced by radiation in the plants growing under the net. This influence was more pronounced in the afternoon, when the whole tree transpiration was largely dominated by equilibrium transpiration in the plants under netting, and the relationship between transpiration and radiation showed a steeper slope in shaded trees. The reduction in transpiration and the maintenance of photosynthesis in shaded plants with respect to exposed trees indicated that screen structures in semi-arid and arid environments could be considered as an intermediate solution for reducing plant water stress and increasing water use efficiency.     

Airborne LiDAR technique for estimating biomass components of maize: A case study in Zhangye City,Northwest China

Crop biomass is an important ecological indicator of growth, light use efficiency, and carbon stocks in agro-ecosystems. Light detection and ranging (LiDAR) or laser scanning has been widely used to estimate forest structural parameters and biomass. However, LiDAR is rarely used to estimate crop parameters because the short, dense canopies of crops limit the accuracy of the results. The objective of this study is to explore the potential of airborne LiDAR data in estimating biomass components of maize, namely aboveground biomass (AGB) and belowground biomass (BGB). Five biomass-related factors were measured during the entire growing season of maize. The field-measured canopy height and leaf area index (LAI) were identified as the factors that most directly affect biomass components through Pearson's correlation analysis and structural equation modeling (SEM). Field-based estimation models were proposed to estimate maize biomass components during the tasseling stage. Subsequently, the maize height and LAI over the entire study area were derived from LiDAR data and were used as input for the estimation models to map the spatial pattern of the biomass components. The results showed that the LiDAR-estimated biomass was comparable to the field-measured biomass, with root mean squared errors (RMSE) of 288.51 g/m² (AGB), and 75.81 g/m² (BGB). In conclusion, airborne LiDAR has great potential for estimating canopy height, LAI, and biomass components of maize during the peak growing season.     

Evaporation and canopy characteristics of coniferous forests and grasslands

Canopy-scale evaporation rate (E) and derived surface and aerodynamic conductances for the transfer of water vapour (g_s and g_a, respectively) are reviewed for coniferous forests and grasslands. Despite the extremes of canopy structure, the two vegetation types have similar maximum hourly evaporation rates (E _max) and maximum surface conductances (g_smax) (medians = 0.46 mm h^-1 and 22 mm s^-1). However, on a daily basis, median E _max of coniferous forest (4.0 mm d^-1) is significantly lower than that of grassland (4.6 mm d^-1). Additionally, a representative value of g_a for coniferous forest (200 mm s^-1) is an order of magnitude more than the corresponding value for grassland (25 mm s^-1). The proportional sensitivity of E, calculated by the Penman-Monteith equation, to changes in g_s is >0.7 for coniferous forest, but as low as 0.3 for grassland. The proportional sensitivity of E to changes in g_a is generally ±0.15 or less.Boundary-line relationships between g_s and light and air saturation deficit (D) vary considerably. Attainment of g_smax occurs at a much lower irradiance for coniferous forest than for grassland (15 versus about 45% of full sunlight). Relationships between g_s and D measured above the canopy appear to be fairly uniform for coniferous forest, but are variable for grassland. More uniform relationships may be found for surfaces with relatively small g_a, like grassland, by using D at the evaporating surface (D₀) as the independent variable rather than D at a reference point above the surface. An analytical expression is given for determining D₀ from measurable quantities. Evaporation rate also depends on the availability of water in the root zone.Below a critical value of soil water storage, the ratio of evaporation rate to the available energy tends to decrease sharply and linearly with decreasing soil water content. At the lowest value of soil water content, this ratio declines by up to a factor of 4 from the non-soil-water-limiting plateau. Knowledge about functional rooting depth of different plant species remains rather limited. Ignorance of this important variable makes it generally difficult to obtain accurate estimates of seasonal evaporation from terrestrial ecosystems.     

Environmental and physiological regulation of transpiration in tropical forest gap species: the influence of boundary layer and hydraulic properties

Environmental and physiological regulation of transpiration were examined in several gap-colonizing shrub and tree species during two consecutive dry seasons in a moist, lowland tropical forest on Barro Colorado Island, Panama. Whole plant transpiration, stomatal and total vapor phase (stomatal + boundary layer) conductance, plant water potential and environmental variables were measured concurrently. This allowed control of transpiration (E) to be partitioned quantitatively between stomatal (g _s) and boundary layer (g _b) conductance and permitted the impact of invividual environmental and physiological variables on stomatal behavior and E to be assessed. Wind speed in treefall gap sites was often below the 0.25 m s^–1 stalling speed of the anemometer used and was rarely above 0.5 m s^–1, resulting in uniformly low g _b (c. 200–300 mmol m^–2 s^–1) among all species studied regardless of leaf size. Stomatal conductance was typically equal to or somewhat greater than g _b. This strongly decoupled E from control by stomata, so that in Miconia argentea a 10% change in g _s when g _s was near its mean value was predicted to yield only a 2.5% change in E. Porometric estimates of E, obtained as the product of g _s and the leaf-bulk air vapor pressure difference (VPD) without taking g _b into account, were up to 300% higher than actual E determined from sap flow measurements. Porometry was thus inadequate as a means of assessing the physiological consequences of stomatal behavior in different gap colonizing species. Stomatal responses to humidity strongly limited the increase in E with increasing evaporative demand. Stomata of all species studied appeared to respond to increasing evaporative demand in the same manner when the leaf surface was selected as the reference point for determination of external vapor pressure and when simultaneous variation of light and leaf-air VPD was taken into account. This result suggests that contrasting stomatal responses to similar leaf-bulk air VPD may be governed as much by the external boundary layer as by intrinsic physiological differences among species. Both E and g _s initially increased sharply with increasing leaf area-specific total hydraulic conductance of the soil/root/leaf pathway (G _t), becoming asymptotic at higher values of G _t. For both E and g _s a unique relationship appeared to describe the response of all species to variations in G _t. The relatively weak correlation observed between g _s and midday leaf water potential suggested that stomatal adjustment to variations in water availability coordinated E with water transport efficiency rather than bulk leaf water status.     

Effect of sand shape on evaporation from soil surface

Summary Loss of water through evaporation for different types of soils was examined. Soils were prepared from a clay soil and multiangular or platy sand in different proportions. The evaporation curves are of the type z=a+bx+cx² but with different slopes. Both soils had the same percentage of clay and sand but platy-sand soils behaved as if they had a finer texture.     

Plant competition for light analyzed with a multispecies canopy model

Summary A multispecies canopy photosynthesis simulation model was used to examine the importance of canopy structure in influencing light interception and carbon gain in mixed and pure stands of wheat (Triticum aestivum L.) and wild oat (Avena fatua L.), a common weedy competitor of wheat. In the mixtures, the fraction of the simulated canopy photosynthesis contributed by wheat was found to decline during the growing season and this decline was closely related to reductions in the amount of leaf area in upper canopy layers. For both species in mixture and in monoculture, simulated photosynthesis was greatest in the middle or upper-middle canopy layers and sensitivity analyses revealed that canopy photosynthesis was most sensitive to changes in leaf area and leaf inclination in these layers. Changes in LAI and leaf inclination affected canopy carbon gain differently for mixtures and monocultures, but the responses were not the same for the two species. Results from simulations where the structural characteristics of the two species were substituted indicated that species differences in leaf inclination, sheath area and the fraction of leaf area alive were of minor consequence compared with the differences in total leaf area in influencing relative canopy carbon gain in mixtures. Competition for light in these species mixtures appears to be influenced most by differences in the positioning of leaf area in upper canopy layers which determines, to a great extent, the amount of light intercepted.     

Assessment of transpiration estimates for Picea abies trees during a growing season

Summary The tree-trunk heat balance method with internal heating and sensing of temperature was used to estimate sap-flow rate of spruce trees in a stand in southern Sweden. Sap-flow rate values were scaled up to stand transpiration and utilised for calculation of canopy conductance. The calculated values provided the basis for a function relating canopy conductance to vapour pressure deficit, which was implemented in the Penman-Monteith equation. The stand was mostly growing in non-limiting soil water conditions (irrigation regime applied during dry periods). The whole-season transpiration was assessed by two different approaches and then compared: the sap-flow rate measurements were scaled to stand transpiration and the adapted Penman-Monteith estimate. They gave similar results: the transpiration totals differed by 3% and the coefficient of determination of the linear regression was r² = 0.89. Similarly good was the assessment for a set of rainy days. The Penman-Monteith estimate adapted in this way proved to be reasonably precise and reliable in this forest stand and usable even in wet conditions. The seasonal transpiration of the spruce stand was 392 mm according to the adapted Penman-Monteith equation. Mean daily transpiration was 1.8 mm and daily maximum transpiration was 4.8–4.9 mm as estimated by sap-flow rate measurements.     

Oscillations in stomatal conductance of hybrid poplar leaves in the light and dark

Cycling of stomatal conductance in three hybrid poplar ( Populus sp.) cultivars was observed under a variety of conditions. Illumination of plants kept previously in the dark induced very large oscillations with a period of about 40 min and large oscillations with a shorter period (< 10 min) were superimposed on the longer cycles. During these oscillations, large changes in conductance could occur very rapidly (1.0 cm s⁻¹ in 3 min). Plants in constant light also displayed both long and short term cycles in conductance, but these were smaller in amplitude than those induced by sudden illumination. Stomatal oscillations were also observed in darkness and after darkening of previously illuminated plants. These oscillations had shorter (< 30 min) and less regular periods than those observed in the light. Such cycling in the dark is rare. Cycling of the two leaf surfaces was sometimes in synchrony in the light, and more so after a perturbation. Little synchrony between the two surfaces was observed in the dark. Stomatal movements of different leaves on a plant were usually relatively independent. Transient stomatal opening occurred following leaf excision in the light or dark, and often after sudden darkening of intact leaves. Also, stomata of intact leaves sometimes transiently closed following illumination.     

The applicability of empirical vegetation indices for determining leaf chlorophyll content over different leaf and canopy structures

Retrieving leaf chlorophyll content at a range of spatio-temporal scales is central to monitoring vegetation productivity, identifying physiological stress and managing biological resources. However, estimating leaf chlorophyll over broad spatial extents using ground-based traditional methods is time and resource heavy. Satellite-derived spectral vegetation indices (VIs) are commonly used to estimate leaf chlorophyll content, however they are often developed and tested on broadleaf species. Relatively little research has assessed VIs for different leaf structures, particularly needle leaves which represent a large component of boreal forest and significant global ecosystems. This study tested the performance of 47 published VIs for estimating foliar chlorophyll content from different leaf and canopy structures (broadleaf and needle). Coniferous and deciduous sites were selected in Ontario, Canada, representing different dominant vegetation species (Picea mariana and Acer saccharum) and a variety of canopy structures. Leaf reflectance data was collected using an ASD Fieldspec Pro spectroradiometer (400–2500 nm) for over 300 leaf samples. Canopy reflectance data was acquired from the medium resolution imaging spectrometer (MERIS). At the canopy level, with both leaf types combined, the DD-index showed the strongest relationship with leaf chlorophyll (R² = 0.78; RMSE = 3.56 μg/cm²), despite differences in leaf structure. For needleleaf trees alone the relationship with the top VI was weaker (D_[red], R² = 0.71; RMSE = 2.32 μg/cm²). A sensitivity study using simulated VIs from physically-modelled leaf (PROSPECT) and canopy (4-Scale) reflectance was performed in order to further investigate these results and assess the impacts of different background types and leaf area index on the VIs’ performance. At the leaf level, the MNDVI8 index showed a strong linearity to changing chlorophyll and negligible difference to leaf structure/type. At canopy level, the best performing VIs were relatively consistent where LAI ≥ 4, but responded strongly to differences in background at low canopy coverage (LAI = 2). This research provides comprehensive assessments for the use of spectral indices in retrieval of spatially-continuous leaf chlorophyll content at the leaf (MTCI: R² = 0.72; p < 0.001) and canopy (DD: R² = 0.78; p < 0.001) level for resource management over different spatial and temporal scales.     

基于多角度植被指数的马尾松林LAI反演方法

CHRIS/PROBA是目前具有最高空间分辨率（17 m×17 m）的星载多角度高光谱数据,该款数据在反演植被垂直结构参数,如树高、叶面积指数（leaf area index,LAI）等方面具有重要的应用前景。基于四尺度几何光学模型得到马尾松（Pinus massoniana Lamb.）冠层的归一化差分植被指数（normalized difference vegetation index,NDVI）各向异性分布规律,利用CHRIS红光特征波段和近红外特征波段构建一种新型多角度植被指数（normalized hotspot-dark-spot difference vegetation index,NHDVI）,并将其应用于CHRIS数据对马尾松林的LAI遥感估算上。结果显示：（1）相比归一化差分植被指数（NDVI）与土壤调节植被指数（soil adjusted vegetation index,SAVI）而言,NHDVI能很好地融合光谱信息与角度信息,与地面实测LAI的决定系数达到0.7278;（2）利用NHDVI-LAI统计回归模型方法来反演LAI值,将得到的LAI值与地面实测值进行相关性分析,结果拟合优度达到0.8272,均方根误差RMSE为0.1232。与传统植被指数相比,包含角度信息的多角度植被指数对LAI的反演在精度上有较大提升,同时比基于辐射传输模型的反演方法更简易、实用。     

小麦叶面积指数与冠层反射光谱的定量关系

在分析不同氮素水平下小麦叶面积指数（LAI）和冠层光谱反射率随生育期变化模式的基础上,确立了LAI与冠层光谱反射率及光谱参数的相关关系,提出了小麦LAI的敏感光谱参数及预测方程.结果表明,小麦LAI和近红外短波段（760～1 220 nm）反射率都随施氮量的增加呈上升趋势,可见光波段反射率则相反;从拔节期到成熟期,LAI和近红外短波段反射率均表现为先上升后下降的趋势,而可见光波段（460～710 nm）反射率随生育期的推进先降低后升高,以孕穗期反射率最低,近红外长波段区域（1 480～1 650 nm）反射率的变化与可见光部分相同.LAI与可见光波段反射率呈负相关,与近红外短波段反射率呈极显著正相关,其中以810 nm相关性最好.可以选择RVI(810,510)和DVI(810,560)作为反演小麦LAI的光谱参数.另外,在证明垂直植被指数PVI和转换型土壤调整指数TSAVI对LAI预测能力的同时,发现利用RVI(810,510)、DVI(810,560)和PVI 3个植被指数共同推算小麦LAI的准确度更高.     

A high resolution green area index for modelling the seasonal dynamics of CO<Subscript>2</Subscript> exchange in peatland vascular plant communities

We studied vegetation dynamics at peatlands, differing in their climate, land use management history and vegetation community in Ireland and Finland over a two-year period. Our aim was to develop a species-specific method to be used to (1) describe the seasonal dynamics of green (photosynthetic) area (GA) of the vegetation and (2) incorporate these changes into CO₂ exchange models. The extent of temporal and spatial variation between and within communities indicated the need for a two-step calculation approach for each community. Firstly, at biweekly to monthly intervals, GA of all species within a range of vascular plant communities was estimated by non-destructive field measurements. Gaussian or log-normal models were fitted to describe the seasonal dynamics of each species. Secondly, an estimate of community vascular green area (VGA) was obtained by summing the modelled daily GA of all species within the community. The highest values of VGA (2.1–6.0 m² m⁻²) occurred within the reed communities at the rewetted cutaway peatland in Ireland and the lowest at the ombrotrophic lawn communities in Finland (0.5–1.0 m² m⁻²). The relationship between light saturated gross photosynthesis (P _G) and VGA was either linear or hyperbolic depending on the degree of self-shading that occurred within each community. The addition of the VGA term into P _G models improved the explaining power of the model by 57.6, 24.5 and 23% within the Typha latifolia, Phalaris arundinacea and Eriophorum angustifolium/Carex rostrata communities, respectively. VGA proved useful in recording the seasonal development of a wide range of peatland vascular plant communities over geographically and climatically different regions.     

耦合叶肉导度的陆面过程模型最大叶肉导度参数的敏感性分析

陆面过程模型添加叶肉导度能有效改善模型模拟的CO₂施肥效应精度,但叶肉导度模拟受最大叶肉导度参数取值的影响,优化模型中最大叶肉导度参数是改进陆面过程模型叶肉导度和CO₂施肥效应模拟的重要途径。以EALCO（Ecological Assimilation of Land and Climate Observations）模型为例添加叶肉导度,通过人为改变最大叶肉导度值的取值,分析模型输出结果对最大叶肉导度的响应,揭示最大叶肉导度参数在模型中的敏感性,并与已有研究结果或观测数据比较,探讨耦合叶肉导度的陆面过程模型最大叶肉导度参数优化的途径。模拟试验以美国哈佛森林典型温带落叶阔叶林生态监测站（US-Ha1 site,Harvard Forest Environmental Monitoring site）数据为驱动。结果显示：（1）随最大叶肉导度增加,总初级生产力（GPP, Gross Primary Production）模拟精度增加,但最大叶肉导度取值大于1.0 mol m^-2 s^-1后模拟精度改善有限,最大叶肉导度小于1.0 mol m^-2 s^-1时GPP模拟精度对最大叶肉导度变化响应敏感;（2）证实了叶肉导度与气孔导度之间存在明显线性关系,最大叶肉导度取值的变化能明显影响这种线性关系的斜率。当最大叶肉导度取值从0.5 mol m^-2 s^-1增加到1.2 mol m^-2 s^-1时,气孔导度与叶肉导度的比值从0.75左右降至0.36,这个结果表明,通过明确某一植被功能型叶肉导度与气孔导度比值,可以间接确定模型最大叶肉导度的合理取值范围;（3）证实了陆面过程模型添加叶肉导度能改进CO₂施肥效应模拟精度,最大叶肉导度值能影响施肥效应模拟结果,当最大叶肉导度高于0.57 mol m^-2 s^-1后,随最大叶肉导度增加,模拟GPP随大气CO₂浓度增加的增长率呈下降趋势;（4）在月尺度上叶肉导度模拟对最大叶肉导度值的敏感性随不同生长季而不同,在生长盛期的7、8月份最大叶肉导度对叶肉导度模拟结果影响最大,其次是5、6、9月份等生长次盛期,其他月份的影响较小。     

Re-evaluation of the allometry of wet thermal conductance for birds

Wet thermal conductance is an important thermoregulatory parameter for birds and mammals. It is generally calculated as C(wet) (ml O2 g(-1) h(-1) degrees C(-1)) = VO2/(T(b)-T(a)), where VO2 is metabolic rate measured in ml O2 g(-1) h(-1), T(b) is body and T(a) is ambient temperature measured in degrees C. Minimum C(wet) is measured at T(a) at or below the lower critical temperature (T(lc)) of the thermoneutral zone, and is strongly influenced by time of day (rest or activity phase) and body mass [J. Aschoff, Comp. Biochem. Physiol. 69A (1981) 611]. Allometric analyses indicate differences in C(wet) for passerine and non-passerine birds, in their rest and active phases (Aschoff, 1981). The allometric slope for non-passerine rest-phase (-0.583) is lower than that for non-passerine active-phase (-0.484), and passerine rest-phase (-0.461) and active-phase (-0.463), although none of these slopes are significantly different. This different-sloped relationship for non-passerine rest-phase C(wet) extrapolates to lower-than-expected values at high body mass, and so this allometric relationship may be inappropriate for predictive purposes. Consequently, we have reanalysed Aschoff's (1981) data, as well as more recent compilations, to determine a more useful allometric relationship for C(wet) of non-passerine rest-phase birds. Re-analyses of minimum thermal conductance data from Drent and Stonehouse [Comp. Biochem. Physiol. 40A (1971) 689], Aschoff (1981) and Gavrilov and Dolnik [Acta XVIII Congressus Internationalis Ornithologici Moscow (1982) 421] indicate that the most appropriate regressions for predicting C(wet) (ml O2 g(-1) h(-1) degrees C(-1)) of birds from body mass (M; g) are the pooled regressions for non-passerine and passerine birds, in the active (alpha) and resting (rho) phases, using data tabulated by Aschoff (1981): alpha, C(wet)=0.994M(-0.509); rho, C(wet)=0.702M(-0.519). C(wet) is approximately 40% higher in the active phase than the rest phase. Regressions of various data sets for C(wet) of birds and mammals indicate a similar slope of approximately -0.5 for the allometric relationship, but significantly higher elevations for mammals compared to birds. The approximately 50% higher C(wet) for mammals than birds indicates a better physical insulation for birds than mammals of the same body mass. The general scaling of C(wet) with M(-0.5) indicates that (T(b)-T(lc)) should scale with M(0.22), if mass-specific metabolic rate scales with M(-0.28) [Reynolds and Lee, Am. Nat. 147 (1996) 735]. The observed scaling for (T(b)-T(lc)) of M(0.183) (calculated from Gavrilov and Dolnik, 1985) is consistent with this expectation.     

基于景观尺度过程模型的长白山地表径流量时空变化特征的模拟

运用已建立的景观尺度生态系统生产力过程模型EPPML对长白山自然保护区地表径流量的季节动态和空间分布特征进行了模拟,对其与环境因子间的相互关系进行了分析．模拟结果表明,长白山自然保护区地表径流量的季节进程表现出明显的三峰型,6月和9月出现低谷,8月达最大(2．58mm·d^-1),与土壤含水量的季节变化趋势十分相似．1995年地表径流量的模拟值平均为0．203m·年^-l,空间分布随海拔升高呈现明显增加的趋势,最高为高山流砾滩(0．619m·年^-1),最低为阔叶红松林(0．081m·年^-1)．年地表径流量与叶面积指数(LAI)呈极显著负相关指数关系(R^2＝0．857)．年地表径流量与气温和总辐射呈负相关关系(R^2分别为0．965和0．836),与降水量、相对湿度和风速呈正相关关系(R^2均大于0．950),还与土壤特性密切相关。     

Structural complexity and land‐surface energy exchange along a gradient from arctic tundra to boreal forest

Question: Current climate changes in the Alaskan Arctic, which are characterized by increases in temperature and length of growing season, could alter vegetation structure, especially through increases in shrub cover or the movement of treeline. These changes in vegetation structure have consequences for the climate system. What is the relationship between structural complexity and partitioning of surface energy along a gradient from tundra through shrub tundra to closed canopy forest? Location: Arctic tundra‐boreal forest transition in the Alaskan Arctic. Methods: Along this gradient of increasing canopy complexity, we measured key vegetation characteristics, including community composition, biomass, cover, height, leaf area index and stem area index. We relate these vegetation characteristics to albedo and the partitioning of net radiation into ground, latent, and sensible heating fluxes. Results: Canopy complexity increased along the sequence from tundra to forest due to the addition of new plant functional types. This led to non‐linear changes in biomass, cover, and height in the understory. The increased canopy complexity resulted in reduced ground heat fluxes, relatively conserved latent heat fluxes and increased sensible heat fluxes. The localized warming associated with increased sensible heating over more complex canopies may amplify regional warming, causing further vegetation change in the Alaskan Arctic.     

Leaf nitrogen distribution and whole canopy photosynthetic carbon gain in herbaceous stands

The amount of photosynthetically-active photon flux density incident upon a leaf and the nitrogen content of that leaf strongly affect the photosynthetic carbon gain of that leaf. Therefore, the canopy structure of a stand, affecting the light climate in the canopy, and the leaf nitrogen distribution pattern in the canopy, affect the carbon gain of the whole canopy. This review discusses the results of studies directed to this problem and obtained so far in open and in dense canopies of stands of herbaceous, monocotyledonous or dicotyledonous, plants in their growing or flowering stages. It is found that the leaf nitrogen distribution pattern in the canopy is vertically non-uniform, and in dense stands more strongly so than in open stands. The leaf nitrogen distribution pattern in most canopies closely approaches an optimal pattern in that it maximizes whole canopy potential carbon gain as calculated for the actual total leaf nitrogen content and leaf area index of the stand. The resulting increase in potential carbon gain as compared to a uniform leaf nitrogen distribution pattern is considerable and it is larger in dense stands than in open stands. For at least some dense stands simulation studies show that with the available total leaf nitrogen content, whole canopy carbon gains could still be considerable higher had a lower leaf area index been developed.     

Effects of mesophyll conductance on vegetation responses to elevated CO2 concentrations in a land surface model

Mesophyll conductance (g_m) is known to affect plant photosynthesis. However, g_m is rarely explicitly considered in land surface models (LSMs), with the consequence that its role in ecosystem and large‐scale carbon and water fluxes is poorly understood. In particular, the different magnitudes of g_m across plant functional types (PFTs) are expected to cause spatially divergent vegetation responses to elevated CO₂ concentrations. Here, an extensive literature compilation of g_m across major vegetation types is used to parameterize an empirical model of g_m in the LSM JSBACH and to adjust photosynthetic parameters based on simulated A_n ? C_i curves. We demonstrate that an explicit representation of g_m changes the response of photosynthesis to environmental factors, which cannot be entirely compensated by adjusting photosynthetic parameters. These altered responses lead to changes in the photosynthetic sensitivity to atmospheric CO₂ concentrations which depend both on the magnitude of g_m and the climatic conditions, particularly temperature. We then conducted simulations under ambient and elevated (ambient + 200 μmol/mol) CO₂ concentrations for contrasting ecosystems and for historical and anticipated future climate conditions (representative concentration pathways; RCPs) globally. The g_m‐explicit simulations using the RCP8.5 scenario resulted in significantly higher increases in gross primary productivity (GPP) in high latitudes (+10% to + 25%), intermediate increases in temperate regions (+5% to + 15%), and slightly lower to moderately higher responses in tropical regions (?2% to +5%), which summed up to moderate GPP increases globally. Similar patterns were found for transpiration, but with a lower magnitude. Our results suggest that the effect of an explicit representation of g_m is most important for simulated carbon and water fluxes in the boreal zone, where a cold climate coincides with evergreen vegetation.     

Light distribution and foliage structure in an oak canopy

 Leaf angle distribution and shoot bifurcation ratio were measured and related to photon flux density (PFD) distribution in an oak canopy. Leaf angle distribution deviated substantially from random and changed markedly throughout the canopy. The observed leaf angle distribution was described by an ellipsoidal function with the single parameter of the distribution, x, changing from 1.6 at the top of the canopy to 3.2 in the lowest branches. In vertically homogeneous canopies, the extinction coefficient for diffuse radiation is expected to decline with increasing leaf area index (LAI). However, in the canopy studied here, the leaf angle distribution changed with height such that the effective extinction coefficient remained constant. Both shoot bifurcation ratio and leaf number per shoot declined with decreasing PFD inside the canopy. Based on these observed relationships, a simple canopy growth model that assumes horizontal homogeneity of the canopy was constructed. Calculations showed that a steady state, when growth in the upper of the canopy is in equilibrium with decline of lower canopy, the total canopy LAI should equal to 4.3. This predicted value of equilibrium LAI is larger than that estimated from measurements of PFD transmission (LAI=3.3), but smaller than that directly determined by litter collection (LAI=6.2 in 1998). Possible reasons for these discrepancies are discussed. Received: 22 June 1998 / Accepted: 7 April 1999