The effects of annual fertilization and complete competition control on fascicle morphology of different aged loblolly pine stands

The goal of this study was to determine how increased nutrient availability affects foliage morphology of loblolly pine (Pinus taeda L.) without introducing the confounding influence of light availability. Morphology of fascicles from the terminal leader (radius, length, specific needle area, density, and needles per fascicle) and terminal leader traits (length, foliated length, total leaf area, and total fascicles) were measured for different aged (ranged from 5-year-old to 12-year-old) loblolly pine stands growing on the Piedmont and lower Coastal Plain of Georgia, USA. A factorial combination of annual fertilization and complete interspecific competition control was applied as stand level treatments. Competition control and stand age generally did not affect fascicle morphology. In contrast, annual fertilization significantly increased fascicle length (5%), needles per fascicle (4%), and total leaf area of the terminal (18%), while decreasing specific needle area (4%). Fertilization also increased terminal leader growth (5%) and total number of fascicles (6%). Therefore, loblolly pine foliage morphology does have plasticity to respond to increased nutrient availability. However, the magnitude of these morphological changes is small compared to changes in total canopy leaf area.     

Comparison of optical methods for estimating the opening of the canopy and leaf area in leafy forests

Based on inversion of gap fraction data (Poisson model of foliage distribution), three optical methods using the Demon, the Plant Canopy Analyzer LAI-2000 (PCA) and hemispherical photographs, have been compared to estimate canopy openness (CO) and leaf area index (LAI) in a mature, neutrophil, oak-beech-hornbeam forest on mull in eastern France. Mean CO over the whole hemisphere was similar for PCA (7.9%) and hemispherical photographs (8.0%). The needle method, a vertical point quadrat method, applied to the litter after leaf fall has served as a reference to LAI (4.7). The Demon provided the estimate (4.9) closest to the reference value. The PCA and hemispherical photographs underestimate mean LAI by 30% (3.3) and 19% (3.8), respectively, if used without correction. Based on fish-eye sensors, LAI estimates can be improved if 3 annuli (4.2) or 2 annuli (4.5) are used in place of 5 with the PCA, or by means of logarithmic averaging of gap fractions over azimuth at an appropriate angular resolution (180 degrees: 4.6, or 120 degrees: 5.2) with hemispherical photographs. Not taking into account azimuthal variation in gap fraction distribution generates a more important error than the error induced by light scattering near horizon. In order to improve LAI estimates, an original iterative procedure is presented, which allows the simultaneous calculation of LAI over a broad range of angular azimuthal resolutions.     

Calibration of LAI-2000 canopy analyser with leaf area index in a young eucalypt stand

Plant area index (PAI) measured with a LI-COR LAI-2000 plant canopy analyser (PCA) was calibrated with leaf area index (LAI) in a young stand of Eucalyptus grandis in the KwaZulu-Natal Midlands, South Africa. Destructive sampling and allometric equations were used to estimate LAI at 2 and 3 years after planting. Significant correlations (P<0.001) were found between LAI and PAI for each age with different equations being generated for the two ages (LAI=1.0594(PAI)−0.892 at 2 years of age, and LAI=1.0393(PAI) at 3 years of age). The equations differed from those reported in other eucalypt studies, as the PCA in this study over-predicted LAI at 2 years, and slightly under-predicted at 3 years, of age. It is argued that the stage of growth influenced this calibration, as the canopy and foliar structure may have been different in the young stands, affecting the basic assumptions for the PCA. A broad conversion from PCA derived PAI to LAI may not necessarily be valid for young, short rotation eucalypt plantations.     

Lodgepole pine forest age class dynamics and susceptibility to mountain pine beetle attack

Because mountain pine beetle attack mature pine stands, an understanding of forest age class dynamics is important to managing forests within the distribution of the beetle. The assumed theoretical negative exponential forest age distribution provides an estimate when ecosystem dynamics are in equilibrium. This study investigates the dynamics of forest age distribution for non-equilibrium ecosystem dynamics, which result primarily from large and irregular stand-replacement fire disturbances that alter the forest age distribution. A model experiment using the SEM-LAND model on a 1 million ha lodgepole pine forest landscape was conducted to estimate how the proportion of susceptible area could be influenced by different fire regimes. The results of the simulation suggest that the temporal dynamics of the area susceptible to mountain pine beetle attack are complex and depend on the fire history of the study area, if the area is experiencing large and irregular stand-replacement fires. The age range of the lodgepole pine forest stands susceptible to mountain pine beetle attack might significantly affect the estimate of the area susceptible to attack.     

千烟洲针叶林的比叶面积及叶面积指数

根据实测数据计算了湿地松(Pinus elliotii)、马尾松(P. massoniana) 和杉木(Cunninghamia lanceolata)不同年龄、不同类型叶片的生物量和比叶面积,并结合样地调查数据和相对生长方程计算了中国科学院千烟洲试验站20年生湿地松林、马尾松林、杉木林和针叶混交林的叶面积指数。根据拟合结果,选择如下方程计算3个树种的叶生物量:湿地松W=12.074 1D^{2.151 5}、马尾松W=6.972 7D^{2.197 3}和杉木W=5.261 9D^{2.302 7}。湿地松林的叶生物量(0.822 kg·m^-2)最大,其次为针叶混交林(0.679 kg·m^-2),马尾松林和杉木林相差不大(分别为林0.528和0.572 kg·m^-2)。不同树种、不同年龄、不同类型叶片的比叶面积比较发现,新叶的比叶面积大于老叶,三针一束叶的比叶面积略大于两针一束叶,马尾松的平均半比表面积(8.62 m²·kg^-1)大于湿地松(6.04 m²·kg^-1)和杉木(7.91 m²·kg^-1)。胸径与单木叶片半表面积之间的经验方程为:湿地松LA=0.073D^{2.151 5}、马尾松LA=0.060D^{2.197 3}和杉木LA=0.042D^{2.302 7}。据此计算湿地松林的叶面积指数为5.03,马尾松林和杉木林为4.31,针叶混交林为4.77,该结果比利用CI-110植被冠层数字图像仪测得的结果偏大。     

Crown structure and leaf area of the understorey species Prunus serotina

A detailed biometrical study of the exotic understorey invader Prunus serotina (Ehrh.) was carried out in a mixed coniferous forest stand in northern Belgium. Based on detailed destructive measurements of eight selected model trees, allometric relations of tree height, crown projected area, woody and leaf dry mass and leaf area on tree diameter at breast height (DBH) were derived. The scaling-up procedure from the tree to the stand level was done using the frequency distribution of DBH obtained at the selected experimental plot. The vertical and radial distributions of the tree foliage were estimated by the “cloud” technique. The vertical profile of leaf area showed a bimodal distribution pattern with maxima at heights of 4 and 6 m above the ground. The leaf area index (LAI) of the understorey Prunus serotina as estimated by the described up-scaling procedure (5.1) was significantly higher than the LAI (2.6) as measured by a plant canopy analyser and was also significantly higher than the LAI of the overstorey species Scots pine (1.5–3.0). The LAI of a neighbouring Rhododendron understorey reached only 1.25. This study emphasises the importance of an exotic understorey species in the total leaf area of mixed coniferous forests which might have important implications for the energy and mass exchanges of the entire forest.     

Leaf litterfall,leaf area index,and radiation transmittance in cypress wetlands and slash pine plantations in north-central Florida

The seasonal dynamics of leaf litterfall and leaf area index (LAI, all-sided basis), light penetration and the vertical distribution of surface area index, and the feasibility of estimating LAI from radiation transmittance were studied from April 1993 to March 1994 in the canopies of three cypress (Taxodium ascendens) wetlands and their surrounding slash pine (Pinus elliottii) uplands in Florida flatwoods. Annual leaf litterfall ranged from 324 to 359 g m^–2 in the wetlands, which was very close to the average for 11 sites throughout Florida of 340±26 g m^–2. The seasonal pattern of the normalized LAI obtained for the dominant tree species in the ecosystems could be used to construct the seasonal dynamics of LAI at the ecosystem scale. The vertical distribution of surface area index in the wetlands was significantly different from that in the surrounding pine uplands. The maximum LAI of cypress wetlands in this area was about 8 m² m^–2, which was higher than the maximum of slash pine plantations of 6 m² m^–2. Cypress leaves were strongly erectophile in space. Results showed that the LAI-2000 canopy analyzer could generally be used to estimate forest LAI, whether the forest canopy was closed or not, if an overall clumping index of 2.00 was applied. However, as LAI decreased, the relative error contained in the radiation-based LAI estimates increased. This indicated that foliage clumping at the stand scale was more important than that at the tree or branch scale.     

Variation of specific leaf area and upscaling to leaf area index in mature Scots pine

Reliable and objective estimations of specific leaf area (SLA) and leaf area index (LAI) are essential for accurate estimates of the canopy carbon gain of trees. The variation in SLA with needle age and position in the crown was investigated for a 73-year-old Scots pine (Pinus sylvestris L.) stand in the Belgian Campine region. Allometric equations describing the projected needle area of the entire crown were developed, and used to estimate stand needle area. SLA (cm² g⁻¹) as significantly influenced by the position in the crown and by needle age (current-year versus 1-year-old needles). SLA increased significantly from the top to the bottom of the crown, and was significantly higher near the interior of the crown as compared to the crown edge. SLA of current-year needles was significantly higher than that of 1-year-old needles. Allometric relationships of projected needle area with different tree characteristics showed that stem diameter at breast height (DBH), tree height and crown depth were reliable predictors of projected needle area at the tree level. The allometric relationships between DBH and projected needle area at the tree level were used to predict stand-level needle area and estimate LAI. The LAI was 1.06 (m² m⁻²) for current-year needles and 0.47 for 1-year-old needles, yielding a total stand LAI of 1.53.     

不列颠·哥伦比亚省北部小杆松及白云杉立地指数与生态立地质量的关系

为在ＰｒｉｎｃｅＲｕｐｅｒｔ林区的“亚北方”部分建立生态立地质量与森林生产力的联系,对从９３个小杆松林分和７７个白云杉林分获得的数据进行了分析．所研究的林分处于两个气候状况、８个土壤水分状况以及５个土壤养分状况．这些气候、土壤水分和养分状况被视为等级变量用于林地分类和回归分析．小杆松和白云杉的立地指数随土壤水分和养分状况变化而变化,但不依赖于气候变化．与土壤水分相关的变化格局对两个种来说很相似,但与土壤养分相关的变化格局则全然不同．在所建立的５类回归模型中,土壤小区模型对于两个种都显示出立地指数与土壤水分和养分状况具有很强的相互关系（Ｒ２＞０．８０,ＳＥＥ≤１．６ｍ）．可以认为土壤水分和养分的等级度量在大范围内可作为小杆松和白云杉立地指数的预测预报因子．     

Carbon and Nitrogen Dynamics of Boreal Jack Pine Stands With and Without a Green Alder Understory

We compared the species composition, structure and selected components of the carbon (C) and nitrogen (N) budgets of similar-aged, mature boreal jack pine (Pinus banksiana Lamb.) forests with and without green alder [Alnus crispa (Ait.) Pursh.] in two different boreal environments. The C and N content of the overstory biomass components (for example, stem, branch, and foliage), total vegetation, forest floor, and mineral soil were greater (P= 0.05 to P= 0.10) for jack pine with alder (JPA) stands than for jack pine without alder (JP) stands at both study areas. Jack pine foliage N isotopic discrimination (δ¹⁵N) and annual litterfall N content were significantly greater (P < 0.05) in the JPA than the JP stands at both study areas, suggesting that alder was fixing N and that N availability was greater in the JPA than the JP stands. The greater leaf area index (LAI) and overstory C accumulation in the JPA than the JP stands (P < 0.05) is likely because of the greater N availability in the JPA stands, but the effect of soil texture discontinuity on water availability in the JPA stands can not be dismissed. Percent ground cover by feathermoss varied among the jack pine communities and was positively correlated with overstory LAI (r ²= 0.83, P< 0.05). One index of N-use efficiency (NUE), defined as aboveground net primary productivity (ANPP) per litterfall N, was significantly greater (P < 0.05) for the JP than the JPA stands, but a second index of NUE, ANPP/N uptake, did not differ between the two jack pine communities. Jack pine trees growing without alder produced more organic matter per unit of N, but percent N retranslocation from senescing foliage and N mean residence time in the overstory did not differ between the JPA and the JP stands. A conceptual model is presented that illustrates the potential influence of alder on the species composition, structure, and function of boreal jack pine forests. Received 6 January 1998; accepted 15 April 1998.     

基于PROSAIL辐射传输模型的毛竹林叶面积指数遥感反演

采用PROSAIL辐射传输模型建立毛竹林叶面积指数(LAI) 冠层反射率查找表,并结合Landsat TM卫星遥感数据,实现了毛竹林LAI的定量反演.结果表明： PROSAIL模型各输入参数的敏感性由高到低依次为LAI>叶绿素含量(C_ab)>叶片结构参数(N)>平均叶倾角(ALA)>等效水厚度(C_w)>干物质含量(C_m),并以LAI、C_ab两个主要敏感因子用于构建毛竹林LAI 冠层反射率查找表;基于PROSAIL模型的毛竹林LAI遥感反演结果与实测LAI具有很好的一致性,二者相关系数为0.90,均方根误差和相关的均方根误差也较小,分别为0.58和13.0%,但也存在反演LAI平均值高于实际值的问题.     

Do Variations in Local Leaf Irradiance Explain Changes to Leaf Nitrogen within Row Maize Canopies?

Numerous studies have dealt with the relationship between leafnitrogen content and leaf irradiance. However, most of themrefer to dense stands presenting reduced horizontal heterogeneityof foliage distribution. Both gradients of leaf nitrogen andleaf irradiance related to canopy depth are significant undersuch conditions, and modelling radiative exchange using a turbid-mediumanalogy and dividing the canopy into vegetation layers is sufficient.Conversely, row crops such as maize are characterized by stronghorizontal heterogeneity of foliage distribution and the one-dimensional(1D) approach may be unsuitable. We thus modelled the three-dimensional(3D) geometry of maize canopies with varying densities and atdifferent developmental stages using plant digitizing underfield conditions. The nitrogen content per unit area of eachleaf part was obtained subsequently by nitrogen analysis. Wenext calculated radiative exchange using a 3D volume-based approachwithin the canopies in order to estimate local leaf irradianceon a daily integration scale. Vertical gradients in leaf nitrogencontent per unit area observed in dense stands during the vegetativephase corresponded largely to those reported in the literature.We also identified significant gradients in nitrogen contentalong the leaves, which had not before been clearly demonstrated.Our study shows that local light climate during plant developmentplays a major role in leaf nitrogen distribution and remobilization.Moreover, brutal plant thinning involves rapid changes in leafnitrogen partitioning. It is concluded that taking account ofthe 3D heterogeneity of nitrogen and irradiance distributionmay have implications for modelling crop photosynthesis andproduction. Copyright 1999 Annals of Botany Company 3D plant architecture, horizontal gradients in leaf nitrogen, leaf irradiance, leaf nitrogen content per unit area, maize, nitrogen partitioning, nitrogen remobilization, virtual plant, Zea mays L.     

Estimation of deciduous forest leaf area index using direct and indirect methods

This study evaluated one semi-direct and three indirect methods for estimating leaf area index (LAI) by comparing these estimates with direct estimates derived from litter collection. The semi-direct method uses a thin metallic needle to count a number of contacts across fresh litter layers. One indirect method is based on the penetration of diffuse global radiation measured over the course of a day. The second indirect method uses the LAI-2000 plant canopy analyser (PCA) which measures diffuse light penetration from five different sky sectors simultaneously. The third indirect method uses the Demon portable light sensor to measure the penetration of direct beam sunlight at different zenith angles over the course of half a day. The Poisson model of gap frequency was applied to estimate plant area index (PAI) from observed transmittances using the second and third methods. Litter collection from 11 temperate decidous forests gave values of LAI ranging from 1.7 to 7.5. Estimates based on the needle method showed a significant linear relationship with LAI values obtained from litter collections but were systematically lower (by 6–37%). PAI estimates using all three indirect techniques (fixed light sensor system, LAI-2000 and Demon) showed a strong linear relationship with LAI derived from litter collection. Differences, averaged over all forest stands, between PAI estimates from each of the three indirect methods and LAI from litter collections were below 2%. If we consider that LAI=PAI–WAI (wood area index) then, all three indirect methods underestimated LAI by an additional factor close to the value of WAI. One reason could be a local clumping of architectural canopy components: in particular, the spatial dispositions of branchlets and leaves are not independent, leading to a non-random relationship between the distributions of these two canopy components.     

Seasonality of leaf litter and leaf area index data for various tree species in a cool-temperate deciduous broad-leaved forest,Japan, 2005–2014

This paper reports seasonal data regarding leaf litter for 14 deciduous broad-leaved species and one evergreen coniferous species as well as leaf area index (LAI) data for the 14 deciduous broad-leaved species in a cool-temperate deciduous broad-leaved forest in Japan. The seasonal leaf biomass of various tree species is important for accurately evaluating ecosystem functions such as photosynthesis and evapotranspiration under climate change. However, there is a lack of freely available, long-term data. We collected litterfall every 1 to 4 weeks from September or October to November or December each year from 2005 to 2014 in Takayama, Japan (36°08′46″N, 137°25′23″E, 1420 m a.s.l.). After sorting the litter into leaves (according to species categories), stems + branches, and “other”, we dried and weighed the litter groups. We also collected seasonal leaf data (number of leaves and leaf length and width) for each broad-leaved species, which we recorded every 1 to 4 weeks from April or May to October or November using multiple target shoots. To estimate the LAI in autumn for each deciduous broad-leaved species, we used a semi-empirical model of the vertical integration of leaf dry mass per unit leaf area. To estimate the LAI in spring and summer, we used the relationship between the LAI in autumn and the seasonal leaf data. Our data provide input, calibration, and validation parameters for determining LAI based on satellite remote-sensing observations or radiative transfer models and for use in ecosystem models.     

Responses of sap flux and stomatal conductance of Pinus taeda L. trees to stepwise reductions in leaf area

Herbivory or artificial foliage removal has been shown to affect gas exchange and canopy water relations. In this study, canopy architecture and water relations in response to progressive defoliation were examined in a stand of 8-year-old loblolly pine (Pinus taeda L.) trees, a shade-intolerant, pioneer species common in the south-eastern USA. Sap flux was measured with constant heat sap flow gauges in order to estimate canopy stomatal conductance (Gs) while foliage in the 6 m high stand was harvested in 1 m increments from the bottom up. Leaf-level stomatal conductance and water potential data were also collected. Profiles of silhouette area ratio and specific leaf area showed no trends with crown height, reflecting an open canopy (leaf area index = 1.55). Therefore, short-term changes in Gs with foliage removal were attributed to hydraulic effects rather than influences of changes in mean microclimate conditions on Gs of remaining foliage. A large increase in Gs was observed during the 6 h pruning period which fully compensated for the reductions in foliage area down to 45%. Canopy stomatal conductance and whole plant liquid phase conductance as calculated from sap flux were both influenced by the rate of growth as indicated by the annual basal area increment.     

The chemical composition of pine foliage in relation to the population dynamics of the pine beauty moth,Panolis flammea,in Scotland

Summary This paper reports part of a study to determine why damaging outbreaks of the pine beauty moth, Panolis flammea (D & S) (Lepidoptera: Noctuidae), in Scotland are frequent on lodgepole pine but do not occur on Scots pine, and why outbreaks on lodgepole pine are mainly confined to trees growing in deep unflushed peat. The elongation of shoots and the growth of needles of Scots pine occurred later in the season than did those of lodgepole pine. The foliage of Scots pine generally had a higher level of nitrogen, and consistently had a higher level of phosporus, but had a consistently lower level of tannins than that of lodgepole pine during the period when the larvae were feeding each year. The nitrogen content of the foliage of lodgepole pine growing in an iron pan soil was generally higher than that of lodgepole pine growing in deep peat during the same period but there were no general differences in the phosphorus or tannin contents of lodgepole pine in the 2 soil types. These findings suggest that Scots pine is a more suitable host plant than lodgepole pine and that the foliage of lodgepole pine growing in deep peat is not more suitable than lodgepole pine growing in an iron pan soil. On the basis of the chemical analyses used in this study, it is concluded that the abundance of pine beauty moth in Scotland is not strongly influenced by the nutritional suitability of its host plants.     

Pinus taeda forest growth predictions in the 21st century vary with site mean annual temperature and site quality

Climate projections from 20 downscaled global climate models (GCMs) were used with the 3‐PG model to predict the future productivity and water use of planted loblolly pine (Pinus taeda) growing across the southeastern United States. Predictions were made using Representative Concentration Pathways (RCP) 4.5 and 8.5. These represent scenarios in which total radiative forcing stabilizes before 2100 (RCP 4.5) or continues increasing throughout the century (RCP 8.5). Thirty‐six sites evenly distributed across the native range of the species were used in the analysis. These sites represent a range in current mean annual temperature (14.9–21.6°C) and precipitation (1,120–1,680 mm/year). The site index of each site, which is a measure of growth potential, was varied to represent different levels of management. The 3‐PG model predicted that aboveground biomass growth and net primary productivity will increase by 10%–40% in many parts of the region in the future. At cooler sites, the relative growth increase was greater than at warmer sites. By running the model with the baseline [CO₂] or the anticipated elevated [CO₂], the effect of CO₂ on growth was separated from that of other climate factors. The growth increase at warmer sites was due almost entirely to elevated [CO₂]. The growth increase at cooler sites was due to a combination of elevated [CO₂] and increased air temperature. Low site index stands had a greater relative increase in growth under the climate change scenarios than those with a high site index. Water use increased in proportion to increases in leaf area and productivity but precipitation was still adequate, based on the downscaled GCM climate projections. We conclude that an increase in productivity can be expected for a large majority of the planted loblolly pine stands in the southeastern United States during this century.     

Light regime in relation to plant population geometry

A beam-emission experiment using a Monte Carlo technqiue was attempted in a computer model for the random distribution (RD) of foliage. The RD foliage consisted of 500 leaves of the same size, their individual areas being just 1/100 of the unit land area; i.e., a leaf area index (LAI) of RD foliage was equal to 5. Satisfactory agreement of light transmission under a uniform overcast sky was obtained between the observed values in the RD foliage and theoretical anticipations for any leaf inclination angle and/or leaf area density. This result demonstrated the usefulness of the Monte Carlo technique adopted in this experiment. The Monte Carlo experiment was also applied to a patch-like population. For such a foliage, additional effect of lateral incidence were worth noting in contrast with the microclimates in the corresponding infinite foliage. For every leaf inclination angle the mean light flux density below the middle stratum within a patch-like population was not further attenuate with increasing LAI.     

Growth and crown efficiency of height repressed lodgepole pine; are suppressed trees more efficient?

Leaf area, crown projection area and growth over the last 5 years were measured to assess growth efficiency (GE) and crown efficiency (CE) of dominant (D), codominant (CD) and suppressed (SP) trees growing in height-repressed (P sites) and normally developing (M sites) lodgepole pine stands. Leaf area index (LAI), hydraulic characteristics, and needle nutrient concentrations were also measured. Volume growth of P site trees between 1994 and 1999 was 46% that of M site trees. Volume growth was closely associated with both hydraulic supply capacity (Q*) and leaf area. Height repression was not associated with lower GE, but P site trees had CE that was 24.5% lower than M site trees. Average GE of D and CD trees was 28% lower than that of SP trees, while mean CE for the D trees was 46% greater than that of CD, and 80% greater than for SP trees. Between M and P sites, canopy LAI and Q* per unit leaf area did not differ. Needle nitrogen (N) concentrations of M site trees were 7.6% greater than for P site trees. SP tree needles had the highest concentration of N and phosphorus. The nutrient advantage enjoyed by SP trees presumably allowed them to maintain higher GE for a given Q*/A_l. The fastest growing trees were the large D and CD trees from M sites. As LAI did not differ between sites, height repression on P sites may be a result of total leaf area being distributed among too many small trees.