The effect of missing rings on stand-age estimation of even-aged forests in northern Hokkaido,Japan

The effectiveness of estimating stand age from increment core samples taken at stump height (20 cm above ground) was tested in an even-aged stand ofBetula ermanii that had regenerated after a forest fire in 1945 in Hokkaido, Japan. Careful cross-dating revealed that annual rings were missing in 11 cores out of a total of 42 cores sampled, and that all these missing rings occurred in the outermost part of the core. These facts indicate that precaution has to be taken in selecting trees from which cores are to be sampled. The present work also revealed that those trees with missing rings have a characteristic appearance, with a thinner stem and less crown foliage than normal trees with complete rings. It was also found that even if this appearance test failed, the possibility of missing rings could be detected from a declining growth pattern, with extremely narrow rings on the increment core which normal trees did not show.     

Spatial variation in sapwood area to leaf area ratio and specific leaf area within a crown of silver birch

Spatial variation in sapwood area to leaf area ratio (Huber value, HV) and specific leaf area (SLA) was examined in branches of closed-canopy trees of silver birch (Betula pendula Roth). HV increased basipetally within a crown and decreased with increasing branch order, but exhibited no significant radial trend along a primary branch. HV was primarily determined by branch position in a crown and branch diameter at the sampling point, being independent of the size of the tree and branch. Greater HV in the lower-crown branches is considered a means to mitigate differences in hydraulic transport capacity between the branches located in different canopy layers. Beside branch position and sampling location on a branch, SLA depended significantly on several other variables characterising tree and branch size. SLA increased basipetally within a crown and along a primary branch, but exhibited no significant trend with branch orders. Because height caused leaf area (A_L) to diminish more rapidly than leaf dry weight, A_L primarily determined the vertical variation in SLA.     

Frequency of partial and missing rings in Acer saccharum in relation to canopy position and growth rate

Locally absent growth rings are known to occur in trees during periods of environmental stress, but little evidence has been available on their frequency in moist temperate forests or on their potential for causing problems in dendroecological studies. In this study, 95 stem disks of Acer saccharum cut from trees representing a wide range of size and age were examined for ring anomalies using three techniques – inspection of disks for partial rings, crossdating of each tree against regional master chronologies, and use of stand history evidence. The number of ring anomalies in Acer saccharum was inversely related to growth rate and vigor. Mean percentage of ring anomalies was 1.3% in dominant canopy trees, 4.6% in trees of the intermediate crown class, and 16.2% in overtopped trees. All of the overtopped trees had partial or missing rings, with a mean of 10 per tree and a range of 2 to 20. Use of crossdating to correct for ring anomalies appeared to be successful in 88% of the canopy trees. However, crossdating results were inconclusive in 32% of the overtopped trees because of low ring width variability, multiple missing rings, and short duration of trouble-free segments. Because overtopped trees are of limited value in reconstructing disturbance history and often cannot be crossdated reliably, we suggest that they normally be excluded from disturbance chronologies.     

Within-crown distribution of male and female cones on clonal Sitka spruce trees

The within-crown distribution of cones (strobili) was mapped on 48 mature grafts of Sitka spruce [Picea schensis (Bong.) Carr.]. The total number of cones per tree was increased by a mainstem injection of 20 mg GA_4/7 and 8 mm wide bark ring. The cones and buds were classified as either lateral or terminal on each individual branch. The distribution of lateral female, terminal female, lateral male and terminal male cones, showed a general progression from the upper distal to lower proximal regions of the tree crown in all 12 clones. Evidence for preferential allocation of cones to particular branch types is presented. The region of the tree with the greates number of cones varied with treatment and reflected differences in the sex ratio of the individual trees. There is evidence for a relationship between branch length, position within the crown and the type of cone produced.     

Overcoming crossdating challenges to generate ring-width chronologies for Sequoia sempervirens across its native range

We utilize a large dendrochronological dataset for Sequoia sempervirens of 1.29 million rings from 5952 series for 278 trees to generate location and regional ring-width chronologies and to investigate constraints on crossdating. Increment cores were collected at regular height intervals along the trunk via free-hanging ropes with some terrestrial deadwood sampled. Inter-annual chronologies span 86–1687 yr for 47 locations in primary and secondary forests across the native Sequoia range. Shared signals based on declining strength of paired correlations with distance, strong correlations within regions, and groupings of principal components guide chronology creation for northern (1032 yr, 108 trees) and southern (865 yr, 52 trees) regions and four sub-regions. The suite of chronologies provides location-specific references as well as more robust regional indices that reflect distinct signals along a climatically-integrative latitudinal gradient. Secondary forests date to near completion, and we use a subset of data to assess crossdating for trees in primary forests. Among tree and landscape attributes, old age is the most important predictor of undated and missing rings, and undated rings align with periods of reduced biomass production distributed across centuries-long lifespans. Crossdating metrics vary by height above the ground with breast height (BH) and 10 m samples having more undated rings than upper samples. At 10 m, samples have the most missing rings. Buttressing affects the lower trunk as BH series have higher ring widths and lower interseries correlations compared to those up to 70 m. Whole-trunk sampling maximizes success for crossdating old, structurally-complex Sequoia and enables dating of crown damage and assessment of biomass dynamics related to environmental change. Reference chronologies, regional assessments, and crossdating insights collectively guide future study of Sequoia for applications including climatology, fire ecology, and seismology.     

Carbon uptake and respiration in above-ground parts of a Larix decidua × leptolepis tree

Summary Shade needles of hybrid larch (Larix decidua × leptolepis) had the same rates of photosynthesis as sun needles per dry weight and nitrogen, and a similar leaf conductance under conditions of light saturation at ambient CO₂ (A_max). However, on an area basis, A_max and specific leaf weight were lower in shade than in sun needles. Stomata of sun needles limited CO₂ uptake at light saturation by about 20%, but under natural conditions of light in the shade crown, shade needles operated in a range of saturating internal CO₂ without stomatal limitation of CO₂ uptake. In both needle types, stomata responded similarly to changes in light, but shade needles were more sensitive to changes in vapor pressure deficit than sun needles. Despite a high photosynthetic capacity, the ambient light conditions reduced the mean daily (in summer) and annual carbon gain of shade needles to less than 50% of that in sun needles. In sun needles, the transpiration per carbon gain was about 220 mol mol^–1 on an annual basis. The carbon budget of branches was determined from the photosynthetic rate, the needle biomass and respiration, the latter of which was (per growth and on a carbon basis) 1.6 mol mol^–1 year^–1 in branch and stem wood. In shade branches carbon gains exceeded carbon costs (growth + respiration) by only a factor of 1.6 compared with 3.5 in sun branches. The carbon balance of sun branches was 5 times higher per needle biomass of a branch or 9 times higher on a branch length basis than shade branches. The shade foliage (including the shaded near-stem sun foliage) only contributed approximately 23% to the total annual carbon gain of the tree.     

Quantitative genetics of yield breeding forPopulus short rotation culture. III. Efficiency of indirect selection on tree geometry

Quantitative analysis of genetic covariances was used to identify the critical morphological components of wood productivity and to evaluate the efficiency of indirect selection for these components at the four levels of biological organization, (1) leaf, (2) branch, (3) main stem, and (4) whole-tree, in 4-yearPopulus deltoides ×P. simonii andP. deltoides ×P. nigra F₁ progeny. A total of 44 morphometric traits measured at the four organizational levels showed varying genetic associations with productivity, variations being dependent on traits, developmental processes (current terminal, sylleptics, and proleptics), and hybridization combinations. Most of the leaf and branch traits on the current terminal and/or sylleptic branches had higher genetic correlations with stem-wood volume than those on proleptics, which resulted in larger indirect selection responses in volume, especially in DxS progeny. Indirect clonal selection on leaf size and area, branching capacity, and branch angle at age 4 years was expected to generate 10–35% more genetic gain per year in 6-year volume than direct selection on 6-year volume in the DxS progeny. The efficiency of indirect selection on the numbers of different order branches and bifurcation ratio was greater than 1.0 relative to that for direct selection for stemwood volume in the D × N progeny. Under the pressure of artificial selection for superior volume production, with the proportion selected=15%, the two F₁ progeny populations exhibited distinct evolutionary divergence in tree geometry. The high-yielding D × S clones displayed a decurrent-like crown with strong apical dominance, whereas the crown ideotype for the high-yielding D × N clones was found to be excurrent-like and surrounded by dense foliage and branches.     

Stem Growth and Epicormic Branch Formation in Understorey Beech Trees (Fagus sylvatica L.)

A study of the formation of epicormic branches on suppressedbeech trees revealed developmental sequences that had not hithertobeen studied in depth. By combining architectural analysis andanatomy, it was possible to trace the history of the plant andthus fully understand the ontogenesis of the individuals studied.The main sequences observed were: (1) the formation of moribundtrees that were gradually covered in epicormic branches fromthe base of the stem to the crown following gradual forest closure;(2) the formation of this new crown of epicormic branches wascombined with a very sharp reduction in stem cambial activity(partial rings, if any) and modifications to the ligneous sections;and (3) resumed girth increment following the establishmentof epicormic branches. After describing in detail the methodsand individual variation, we highlight the spatial and temporalpatterns of the phenomena observed. Lastly, we put forward functionalhypotheses with a view to setting these results against a backgroundof forest tree ontogenesis in general. Copyright 2001 Annalsof Botany Company Fagus sylvatica L., beech, architecture, epicormic shoot, anatomy, growth, annual wood ring, missing ring     

Biology of the bamboo Chusquea culeou (Poaceae: Bambusoideae) in southern Argentina

Over a period of 7 years the biology and phenotypic variability of Chusquea culeou were studied at 5 locations in cool temperate forests of southern Argentina. Excavated rhizomes had an average of 1.1 successful rhizome buds, and an average of 2.1 years elapsed between successive generations of rhizomes. Rhizome buds usually develop within the first four years after a rhizome forms. Height, volume and weight of a culm can be calculated from its diameter 1 m above the ground. Culm size, length of foliage leaf blades, and pattern of secondary branching differed among study sites. Dead culms were numerous and commonly remained erect for more than 7 years after dying. New culm shoots appear in spring and reach full size within a few months. Shoots can grow more than 9 cm/day. Less than half of the shoots survived a year; most were killed by moth larvae. Multiple primary branch buds emerge through the culm leaf sheaths in the second spring. The mean number of branch buds at mid-culm nodes varied between 34.8 and 81.5, and the mean number of primary branches was between 22.8 and 40.8. Number and length of branches, and number and length of foliage leaf blades at each node is related to the position of the node on a culm. Most branches grow about 3 cm and produce 1 to 3 foliage leaves annually. Foliage leaf blades generally live 2 years or more; few survive 6 years. Relative lengths of foliage leaf blades and their spacing along a branch permit recognition of annual cohorts.Both gregarious and sporadic flowering have been reported, and every year a few isolated plants flower and die. Length of the life cycle is unknown. Seedlings require up to 15 years to produce culms of mature size. Foliage branches may live more than 23 years, and culms may survive 33 years. Extensive loss of new shoots to predation suggests that gregarious flowering may be driven by a need to escape parasitism. C. culeou clumps expand slowly. Average annual rate of increase of the number of live culms in a clump was 4.6%. Methods of seed dispersal are undocumented. A dense stand of Chusquea culeou had an estimated phytomass of 179 tons/hectare (dry weight), 28% of which was underground. Net annual production was about 16 t/ha dry weight.     

Development of biomass proportions in Norway spruce (Picea abies [L.] Karst.)

This study tests the hypotheses that (1) the above-ground structure of Norway spruce (Picea abies [L] Karst.) is derivable from the functional balance theory, and that (2) crown ratio is a key source of structural variation in trees of different age and social position. Twenty-nine trees were measured in three stands (young, middle-aged, and mature), with three thinning treatments (unthinned, normal, and intensive) in the two older stands. There was a strong linear relationship between the total cross-sectional area of branches and that of stem at crown base. Foliage mass was linearly related with stem basal area at crown base. Also an allometric relationship was found between foliage mass and crown length. The mean length (weighted by basal area) of branches obeyed an exponential function of crown length. The parameters of most of these relationships were independent of slenderness (tree height/breast height diameter) and tree age However, total branch cross-sectional area per stem cross-sectional area in the young trees was greater than in the older trees. The young trees also had slightly shorter branches than predicted by the mean branch length equation. This was probably caused by branch senescence which had not yet started in the young stand. The older trees had a relatively long lower crown segment which was growing slowly and senescing. It was proposed that a segmented crown structure is characteristic of shade tolerant tree species, and that the structural model could be further developed by making the two segments explicit.     

Structural crown properties of Norway spruce (Picea abies [L.] Karst.) and European beech (Fagus sylvatica [L.]) in mixed versus pure stands revealed by terrestrial laser scanning

How tree morphology develops in mixed-species stands is essential for understanding and modelling mixed-stand dynamics. However, research so far focused on the morphological variation between tree species and neglected the variation within a species depending on intra- and interspecific competition. Our study, in contrast, addresses crown properties of nine mature Norway spruces (Picea abies [L.] Karst.) of a pure stand and compares them with ten spruces growing in mixture with European beech (Fagus sylvatica [L.]). The same was done with 11 pure stand beeches and 12 beeches growing in mixture with spruce. Through application of a terrestrial laser scanner and a new skeletonization approach, we deal with both species’-specific morphological traits such as branch angle, branch length, branch bending, crown volume and space occupation of branches within the crown, some of which were hardly accessible so far. Special attention is paid to distinct differences between trees growing in mixed and pure stands: for spruce, our study reveals significantly longer branches and greater crown volumes in the mixed stand when compared to the pure stand. In case of European beech, individuals growing in mixture show flatter branch angles, more distinct ramification, greater crown volumes and a lower share of a single branch’s space occupation in the total crown volume. The results show that the presented methods yield detailed information on the morphological traits analyzed in this study and that interspecific competition on its own may have a significant impact on crown structures. Implications for production ecology and stand dynamics of mixed-species forests are discussed.     

Crown alterations induced by decline: a study of relationships between growth rate and crown morphology in beech (Fagus sylvatica L.)

One of the first symptoms expressed by declining trees is reduced growth in stem diameter and length increment. The possibility of a relationship between length increment and crown thinning in beech (Fagus sylvatica L.) was investigated by developing a computer model to simulate first order branching patterns of the apical 2 m of monopodially branching beech trees, 70–100 years old, for a range of length increment rates. The model was based on values for branching angle, main axis and branch length increment, number of branches produced per year and branch mortality rates for six healthy and declining trees. Shoot growth rates in the apical 2 m of the sample trees ranged from about 5 cm/year (decline class 3) to 43 cm/ year (healthy). Simulations of branching patterns in the apical 2 m of trees growing at different rates indicated that, when growth rate exceeded about 20 cm/year, total first order branch length and area explored were independent of growth rate. When growth rates fell below this value there was a reduction in total area explored and first order branch length due primarily to the formation of fewer branches. More acute branching angles contributed to a reduction in the area explored. Growth rate-related crown thinning could increase the risk of bark necrosis and secondary pathogen infection during dry and/or hot spells.     

Effects of branch position on water relations and gas exchange of European larch trees in an alpine community

Water relations and gas exchange were studied in the crowns of small European larch (Larix decidua Mill.) trees with respect to branch position. The upper-crown branches showed significantly higher branch sap flux rate (F _la) and branch conductance (g _b) compared to the lower crown (P<0.001). Values of leaf conductance (g _l), transpiration rate (E) and net photosynthesis (A), averaged for different ranges of atmospheric vapour pressure deficit (VPD), were also higher in the upper crown position. We suppose that the up to 2.6-fold smaller soil-to-leaf hydraulic conductance observed in the lower branches (P<0.001, compared to upper branches) could contribute to the decreased values of F _la, g _b, g _l, and E in the lower crown position. Variation in tracheid lumen diameter with respect to crown position (P<0.001) supported the hypothesis that branches growing at the crown base are hydraulically more constrained than branches located at the top of the tree. Leaf area to sapwood area ratio (A _la/A _sa) exhibited 1.4 times smaller values in lower crown (P<0.01), however, this could not compensate the effect of decreased hydraulic conductivity of the lower-crown branches.     

Using dendrochronology to model hemlock woolly adelgid effects on eastern hemlock growth and vulnerability

This study examined the relationship between eastern hemlock (Tsuga canadensis (L.) Carr.) crown condition and changes in radial growth associated with infestation by hemlock woolly adelgid Adelges tsugae (Homoptera: Adelgidae) (HWA). Tree-ring chronologies of eastern hemlock were used to develop a binomial decline index based on three consecutive years of below average growth. Radial growth decline was modeled, using logistic regression, as a function of an extensive array of tree, crown, and site variables that were collected over an 11 year period in Delaware Water Gap National Recreation Area. Some site-related variables such as site-location and aspect were significantly related to decline probabilities when considered individually. However, the total proportion of response variance accounted for was low, and the only site variable included in the final model was mean plot-level HWA infestation level. For every 1% increase in mean percent HWA infestation per plot, there was an 8% increase in the likelihood that a tree would be classified as being in decline. Tree crown variables such as live crown ratio, crown density, and the modified ZB_adj index, a combination of foliage transparency and branch dieback, had the most explanatory power, both individually and in the final model. These crown variables were relatively accurate predictors of the degree of hemlock growth decline during HWA infestation.     

臭冷杉生物量分配格局及异速生长模型

摘 要：臭冷杉是长白山阔叶红松林中重要针叶树种,采用整株收获法分析21株臭冷杉地上地下生物量分配格局。在枝条水平上采用样枝直径（BD）、样枝长度（BL）、样枝所在轮生枝位置（WP）建立活枝、针叶生物量异速生长模型,在植株水平上采用胸径（DBH）、树高（H）、年龄（Age）、树冠长度（CL）、树冠比率（CR）、南北向冠幅（CW1）、东西向冠幅（CW2）等变量建立树干木质、树皮、活枝、针叶、粗根及整株生物量模型。并利用逐步线性回归法获得不同器官生物量最优模型。结果表明：（1）活枝生物量主要集中在树冠中下层,针叶生物量集中在树冠中层。树冠中层和下层枝叶生物量无显著差异（p>0.05）;（2）21株臭冷杉地上生物量和地下生物量变动范围分别为1.026–506.047 kg/株和0.241–112.000 kg/株。粗根、活枝、针叶、树干木质、树皮及枯枝生物量占整株生物量的相对比例分别为18.68%、18.39%、12.02%、39.29%、8.70%和2.92%;（3）地上生物量与地下生物量呈显著线性相关（p<0.001）,拟合线性方程斜率为0.23;（4）枝条水平上,活枝生物量模型解释量超过95%,平均预测误差小于30%。与单变量（BD）活枝生物量模型相比,2变量（BD、BL）和3变量（BD、BL、WP）模型解释量分别提高1.2%和2.0%,平均预测误差分别下降6.26%和9.27%。针叶生物量相对较难预测,模型解释量仅为82.7%,平均预测误差接近50%,模型中增加BL 和WP变量并未提高针叶生物量的预测精度。活枝生物量与BD、BL、WP正相关,针叶生物量与BD正相关,与BL、WP负相关;（5）植株水平上,基于胸径的单变量模型可解释量大于90%,增加树高变量未能显著提高生物量模型的预测精度。年龄决定了臭冷杉的树干生物量,忽视年龄变量将会产生生物量预测误差。树冠特征是影响枝叶生物量预测精度的重要变量。综合考虑模型的可解释量及回归系数显著性可知,胸径是预测臭冷杉不同器官生物量的可靠变量。     

干旱胁迫下新疆杨树冠不同高度叶片水分状况与非结构性碳动态北大核心CSCD

水力结构调整及非结构性碳动态对于认知干旱胁迫下植物生存前景和死亡风险至关重要。该研究以新疆杨(Populus bolleana Lauche.)为对象,通过干旱处理分析树冠不同高度叶片水力性状、光合生理特性、不同功能器官间的可溶性糖、淀粉及非结构性碳水化合物(NSC)含量变化,以甄别树冠不同高度叶片对干旱胁迫的生理响应和适应差异。结果表明:(1)在干旱胁迫下,树冠上部的叶水势、叶含水量、枝条含水量普遍低于下部,饱和膨压渗透势及膨压损失点水势在不同树冠高度间差异不显著;(2)干旱处理组植株净光合速率随树冠高度的增加而降低,叶绿素SPAD值在树冠的上部显著低于下部,而水分利用效率在树冠的上部却高于下部;(3)干旱处理组比叶面积在各树冠高度下显著低于对照组,而胡伯尔值在中部及上部高于对照组但差异不显著;(4)干旱处理组,树冠上部叶片淀粉含量显著高于下部,枝条可溶性糖及NSC含量在树冠上部显著高于下部,韧皮部的可溶性糖、淀粉及NSC含量在不同树冠高度间无显著差异,细根的可溶性糖、淀粉及NSC含量在对照组与干旱处理组间的差异不显著。研究发现,干旱处理下,树冠高度的增加会加剧新疆杨枝叶的干旱胁迫,致使树冠上部枝条木质部发生栓塞的风险大于下部,并导致NSC在不同器官间的分配和组分产生差异,但新疆杨植株可通过水分利用效率和形态上的适应性调整减缓干旱胁迫。     

Branch architecture,light interception and crown development in saplings of a plagiotropically branching tropical tree,Polyalthia jenkinsii (Annonaceae)

To investigate crown development patterns, branch architecture, branch-level light interception, and leaf and branch dynamics were studied in saplings of a plagiotropically branching tree species, Polyalthia jenkinsii Hk. f. & Thoms. (Annonaceae) in a Malaysian rain forest. Lengths of branches and parts of the branches lacking leaves ('bare' branches) were smaller in upper branches than in lower branches within crowns, whereas lengths of 'leafy' parts and the number of leaves per branch were larger in intermediate than in upper and lower branches. Maximum diffuse light absorption (DLA) of individual leaves was not related to sapling height or branch position within crowns, whereas minimum DLA was lower in tall saplings. Accordingly, branch-level light interception was higher in intermediate than in upper and lower branches. The leaf production rate was higher and leaf loss rate was smaller in upper than in intermediate and lower branches. Moreover, the branch production rate of new first-order branches was larger in the upper crowns. Thus, leaf and branch dynamics do not correspond to branch-level light interception in the different canopy zones. As a result of architectural constraints, branches at different vertical positions experience predictable light microenvironments in plagiotropic species. Accordingly, this pattern of carbon allocation among branches might be particularly important for growth and crown development in plagiotropic species.     

辽东栎林内不同小生境下幼树植冠构型分析

以黄土高原黄龙山林区辽东栎林内3个小生境（林下、林隙、林缘）下辽东栎天然更新幼树为研究对象,采用典型抽样法对辽东栎幼树侧枝、叶片和树冠的空间分布状况以及生物量分配状况进行调查分析,探讨微生境与幼树植冠构型特征的关系,明确辽东栎幼树对不同小生境的适应策略,为栎林经营和林分结构优化提供理论依据。结果显示：（1）3种生境下辽东栎幼树构型发生了可塑性变化,林下幼树树冠层次比较单一,林隙与林缘的幼树树冠层次更加丰富。（2）由林下至林缘,幼树的树高、枝下高呈逐渐减小的趋势,而地径变化趋势与之相反;幼树的冠幅、树冠面积、树冠率呈先增加后减小的趋势,并且林下与林隙、林缘的差异显著;幼树的总体分枝率、逐步分枝率、枝径比呈先增加后减小的趋势。（3）3种生境下,幼树的一级枝的枝长、直径与倾角随着树高的增加而呈减小的趋势,但3种生境的差异不显著;林下一级枝主要分布在冠层中上部,而林隙与林缘一级枝主要分布在冠层中下、中上部。（4）由林下至林缘幼树叶长、叶宽、单叶面积和比叶面积逐渐降低,而单株叶数、叶总面积、叶面积指数呈先增大后减小趋势;与其他2种生境相比林下叶片分布趋于冠层上部。（5）幼树地上部分生物量中林下主干生物量占83%,枝和叶生物量只占17%;而林隙与林缘虽然各部位生物量有所差异但比例基本一致,其中主干占66%左右,枝和叶生物量占34%左右。研究表明,林隙生境下幼树的构型优于林缘和林下生境,在今后栎林的经营中,可以通过适当间伐来增加林隙数量,为森林更新和结构的优化提供有利条件。     

Biomass partitioning and water content relationships at the branch and whole-plant levels and as a function of plant size in Elaeagnus mollis populations in Shanxi, North China

Understanding of the biomass (dry weight) allocation and water relations in populations will provide useful information on the growth patterns and resource-allocation dynamics. By destructive sampling, foliage, branch and root biomass were measured in the endangered shrub Elaeagnus mollis populations growing in Shanxi province, North China. Biomass partitioning and water content relationships were compared at the branch and whole-plant levels, and as a function of basal diameter (plant size). The biomass was mainly distributed in the bigger branches at the branch level, and in the branch wood at the whole-plant level, and branch biomass (but not foliage or root biomass) increases significantly with increasing basal diameter. As a result, branch wood became the major biomass pool, even though considerable biomass was also allocated to the roots. However, the relative water content decreased from the periphery of the crown to the interior of the shrub at the branch level, and from the aboveground to the belowground at the whole-plant level though no significant variation among foliage, branches, and roots. Yet it increased significantly for the whole-plant with increasing basal diameter. The ratio of belowground to aboveground biomass was smaller than 1.0, even as a function of basal diameter. These growth responses indicated a strong adaptation to the shrub’s growing conditions. Biomass was primarily allocated above the ground and the aboveground components grew faster than the belowground one.     

Spacial distribution of photosynthetic capacity and performance in a mountain spruce forest of Northern Germany

Summary Biomass distribution and diurnal CO₂ uptake under natural conditions were investigated on Picea abies in a mountainous climate (Solling, Northwest Germany). Spruce has a remarkable variability in leaf characteristics. Even on a single branch in the lower sun crown, needle dry weight and surface area change considerably from the branch base to the tip and accoring to exposure. Only about 18% of the total biomass of the tree was current year's growth, about 40% of the needles were 4 years and older reaching a maximal age of 12 years. The main growing zone was at the border of upper shade and lower sun crown and the main accumulation of dry weight was at a greater tree height than was observed for maximal growth of needle numbers or surface area. The annual, new growth shifted toward the upper sun crown. Maximal daily CO₂ uptake was highest in the lower sun crown on days with variable cloud cover when temperatures were moderate and water vapor pressure deficits were low. Also the annual CO₂ uptake was highest in the lower sun crown, where 4-year-old and older needles contributed about 35% to the annual CO₂ uptake of the tree. Current year growth contributed about 15% of the total CO₂ gain. The upper and lower sun crowns produce about 70% of the total carbon gain. The carbon balance of spruce and the distribution of the production process in relation to needle age and crown level are discussed.