The effect of two growth forms of Norway spruce on stand development and radiation interception: a model analysis

Summary Development of tree and canopy structure, and interception of photosynthetically active radiation (PAR) were studied in two model stands of Norway spruce consisting of trees with rapid versus slow site capture. The tree models were derived using Burger's (1953) sample tree material, from which two subpopulations of dominant trees were selected using the rate of horizontal site capture of the tree crowns as the criterion of division. The development of stand structure and interception of PAR were simulated in the two model canopies. The simulation period covered the period from tree age 15–80 years. The average development of the trees in the two subpopulations proved to be very different. The rapidly expanding trees were characterized by low mean within-crown needle area density and a long crown. The slowly expanding trees were smaller but had a higher mean within-crown needle area density. Up to approximately 40 years of age the stand of rapidly expanding trees contained more leaf area and intercepted more radiation than the stand of slowly expanding trees, when canopy cover was held constant. After 40 years of age this relationship was reversed due to the subsequent decline of leaf area in the stand of rapidly expanding trees and the increase in leaf area in the stand of slowly expanding trees. The biological relevancy and silvicultural implications of the simulated patterns of tree and stand development are discussed.     

Spruce budworm growth,development and food utilization on young and old balsam fir trees

Laboratory rearing of spruce budworm, Choristoneura fumiferana, in conjunction with field rearing, gravimetric analyses, a transfer experiment, and foliage chemical analyses at six dates during the period of budworm feeding activity indicated that the age of balsam fir, Abies balsamea, trees (70-year-old mature trees or 30-year-old juvenile trees) affected tree suitability for the spruce budworm via the chemical profile of the foliage. Insects reared on old trees had greater survival and pupal weight, shorter development times, and caused more defoliation than those reared on young trees. Young trees were more suitable for the development of young larvae (instars 2–5), while old trees were more suitable for the development of older, sixth-instar larvae. These results were confirmed by the laboratory transfer experiment. Young larvae fed foliage from young trees had higher relative growth rates (RGR), digestibility (AD), and efficiency of conversion of ingested foliage (ECI) than those fed foliage from old trees. These differences appeared to be related to the high N:tannins ratio, and the high contents of P present in young trees during the development of the young larvae. Old larvae fed foliage from old trees had higher relative growth rates, relative consumption rates (RCR), and digestibility of the foliage than those fed foliage from young trees. The high digestibility of the foliage of old trees was compensated for by a lower efficiency of conversion of digested food (ECD), which in turn resulted in no significant effect of tree age on the efficiency of conversion of ingested foliage by old larvae. The low relative consumption rate of old larvae fed foliage from young trees appeared to be related to the low N:tannins ratio, and the high contents of bornyl acetate, terpinolene, and °-3-carene present in young trees during the budworm sixth instar. Variations in these compounds in relation to tree age may serve as mechanisms of balsam fir resistance to spruce budworm by reducing the feeding rate of sixth instar larvae.     

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.     

Effects of bud phenology and foliage chemistry of balsam fir and white spruce trees on the efficacy of Bacillus thuringiensis against the spruce budworm, Choristoneura fumiferana

Abstract 1 Efficacy of commercial formulations of Bacillus thuringiensis ssp. kurstaki (Btk) against spruce budworm Choristoneura fumiferana was investigated in mixed balsam fir‐white spruce stands. Btk treatments were scheduled to coincide with early flaring of balsam fir shoots, and later with flaring of white spruce shoots. Btk efficacy on the two host trees was compared and examined according to the foliar content of nutrients and allelochemicals and the insect developmental stage at the time of spray. 2 Larvae fed white spruce foliage were less vulnerable to Btk ingestion than larvae fed balsam fir foliage. Higher larval survival on white spruce, observed 10 days after spray, was related to higher foliage content in tannins and a lower N/tannins ratio, which might have induced inactivation of Btk toxins. 3 Larval mortality due to Btk did not depend on spruce budworm larval age. 4 Foliage protection of both host trees was similar in plots treated with Btk: larval mortality due to Btk treatment reduced insect grazing pressure on balsam fir trees; meanwhile, suitability of white spruce foliage seemed to decrease very rapidly, which induced high larval mortality among spruce budworm fed on white spruce trees. Nevertheless, following Btk sprays, 50% more foliage remained on white spruce than on balsam fir trees, because of the higher white spruce foliage production. 5 Both spray timings achieved similar protection of white spruce trees, but Btk treatments had to be applied as early as possible (i.e. during the flaring of balsam fir shoots to optimally protect balsam fir trees in mixed balsam fir‐white spruce stands).     

A Matrix model for Predicting Foliage Weight of Trees by Age Classes

A Leslie-type matrix has been developed to model the annual growth of foliage on individual trees for any species that retains its foliage for more than one year. In order to apply the model to any given species, one must know: (1) age-specific foliage photosynthetic efficiencies; (2) the proportion of foliage of each age class that remains to become one year older; and (3) the diameter of the tree at any two points in time, several years apart. These diameters are used to estimate the maximum eigenvalue of the matrix, which is needed in the determination of the specific numerical matrix which corresponds to a particular individual tree. Although the model is a general one, applicable to any species that has age classes of foliage, it is developed with occasional references to balsam fir, Abies balsamea (L.) Mill. For a data set of 66 dominant and codominant balsam fir trees, the model predicted the weights of the first three age classes of foliage accurately (less than 10% error) over a 6-year period. There was a 23% error for all age classes combined.     

川西亚高山不同年龄紫果云杉径向生长对气候因子的响应

运用树木年轮气候学的基本方法,建立王朗自然保护区紫果云杉在集中分布上限区域的年轮宽度年表,选取差值年表分析不同年龄云杉的径向生长同逐月气候因子的相关及响应关系,结果显示:幼龄组云杉年表的敏感度高于中龄组和老龄组云杉,幼龄组云杉对生长季前及生长季的气温状况显著正相关;中龄组云杉年表仅与当年4月份和7月份的月平均最低气温显著正相关;老龄组云杉的年轮宽度指数同上年生长季(上年8月份)的月平均气温和月平均最低温显著负相关,上年生长季高温的"滞后效应"在老龄组云杉体现的更为突出;幼龄组与中龄组云杉对当年6月份降水持续增加显示出明显的负相关关系,上年12月份的降水会对幼龄组和老龄组云杉径向生长不利。研究表明幼龄组云杉包含的气候信息要优于中龄组和老龄组云杉,在该区域进行相关研究时应根据研究需要选取不同年龄跨度的云杉年表。     

Evidence of a direct chemical plant defense role for maltol against spruce budworm

This study examines the direct chemical defensive role of maltol, a previously identified secondary metabolite found in balsam fir, Abies balsamea (L.) Mill. (Pinaceae), that was detected during herbivory of spruce budworm, Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae). Although used extensively in many industries, in addition to being found in multiple plant species, its functional role in plants remains unknown. The objectives of this study were to quantify the amount of free maltol and its potential conjugated form, maltol glucoside, in various foliage age classes and to evaluate whether constitutive foliage levels of maltol have an impact on spruce budworm fitness in maltol supplementation assays. Gas chromatography–mass spectrometry (GC‐MS) analysis of balsam fir foliage showed that maltol is produced in all foliage age classes tested; however, concentrations were significantly higher in older foliage. Liquid chromatography–mass spectrometry–mass spectrometry (LC‐MS‐MS) analysis showed that maltol also exists in balsam fir in its glucosylated form, a unique discovery in conifers. Similar to maltol, maltol glucoside is also present in current and 1‐year‐old balsam fir foliage and in significantly higher concentration in older foliage. We investigated the impact of maltol‐treated diet on spruce budworm fitness. Maltol additions that reflected constitutive foliage concentrations caused a significant reduction in larval development rate and pupal mass, whereas higher concentrations were required to cause significant mortality. These results suggest that maltol may be an important component of a direct defense strategy in balsam fir against spruce budworm herbivory.     

Spatial patterns of ground vegetation, soil microbial biomass and activity in a mixed spruce-birch stand

Trees directly and indirectly influence the above- and below-ground environment, and can be expected to modify the spatial patterns of organisms associated with the forest floor. This study aimed to examine the effects of a coniferous (Picea abies) and a broad-leaved (Betula pubescens) tree species on the spatial pattern of ground vegetation and soil microbial properties in a mixed stand in central Sweden. I have characterised the species composition of ground vegetation, soil microbial biomass and activity, photosynthetic active radiation (PAR), soil water content and soil pH in the stand, and tested whether the spatial patterns of these variables were related to the positioning of trees. Geostatistics were used to describe the spatial variation in ground vegetation, soil mirobiological properties and the soil surface environment. PAR, soil water content and the cover of the moss Brachytecium reflexum and associated herb species decreased with the influence of spruce trees. Microbial biomass, measured as the amount of phospholipid fatty acids, decreased with spruce influence but increased with the influence of birch trees. Microbial respiration was not affected by spruce but increased with the influence of birch. Ground vegetation and microbial respiration, which were influenced by one tree species only, aggregate on a scale of 4-5 m, corresponding fairly well with patches of a single tree species. Soil microbial biomass, which was affected by both tree species, aggregated on a scale of 7-8 m. roughly corresponding to the distance between patches of spruce and birch trees respectively. I suggest that spruce trees influenced vegetation mainly through shading, and that a difference in the availability of organic matter under birch and spruce trees caused spatial variation in microbial biomass and activity. Thus, spatial patterns in ground vegetation and soil microbial properties may develop in a mixed forest of coniferous-broad leaved trees, as a result of the difference in influence of tree species and nested variation associated with the arrangement of the trees.     

Performance of two Picea abies (L.) Karst. stands at different stages of decline

Summary This is the first in a series of papers on the growth, photosynthetic rate, water and nutrient relations, root distribution and mycorrhizal frequency of two Norway spruce forests at different stages of decline. One of the stands was composed of green trees only while the other included trees ranging in appearance from full green crowns to thin crowns with yellow needles. In this paper we compare the growth and carbohydrate relations of the two stands and examine relationships among growth variables in ten plots. The declining stand produced 65 percent of the wood per ground area compared with the stand in which all trees were green because its foliage produced less wood at any level of leaf area index. The difference in foliage efficiency between the sites could not be explained by differeneces in climate, competition or stand structure. The declining stand appeared to have lower carbon gain as indicated by a smaller increase in reserve carbohydrates before bud break, and weaker sinks for carbohydrates as indicated by less use of the stored carbohydrates than the healthy stand. Thus, growth reduction was probably related to factors which affect both photosynthesis and, even more, the sinks for carbohydrate.     

DOES TREE AGE PREDICT THE OCCURRENCE AND ABUNDANCE OF USNEA LONGISSIMA IN MULTI-AGED SUBMONTANE PICEA ABIES STANDS?

In boreal Scandinavia Usnea longissima is a rare epiphytic lichen associated with old spruce (Picea abies) forests. To test whether tree age was an important determinant of the occurrence and abundance of U. longissima at the scale of a forest stand we estimated thallus numbers on 1008 trees within three submontane spruce stands in south-central Norway. The stands were of all ages and multi-layered and had been selectively cut previous to the 1940s. There was no relationship between number of U. longissima thalli and tree age among trees with dbh of ≥10 cm. The most consistent pattern was a positive relationship between occurrence and number of thalli and stem diameter. After controlling the covariance of stem diameter, there was a tendency for a negative relationship between number of thalli and tree age. The results suggest that in old-growth resembling stands, with already established U. longissima populations, tree size (here measured as stem diameter) is a better predictor of thallus number than tree age. This does not contradict the possibility that other environmental characteristics, associated with stands of older age, may be important for the establishment and growth of U. longissima populations.     

Competition-dependent modelling of foliage biomass in forest stands

Currently, foliage biomass is estimated based on stem diameter or basal area. However, it is questionable whether the relations between foliage and stem observed from plantations of a single tree species can be applied to stands of different structure or species composition. In this paper, a procedure is presented to simulate foliage and branch biomass of tree crowns relative to crown size and light competition. Crowns are divided into layers and segments and each segment is divided into a foliated and an unfoliated fraction. Depending on the competitive status of the segment, leaf area density, specific leaf area and foliated branch fraction are determined. Based on this information, foliage biomass is calculated. The procedure requires a crown shape function and a measure to characterise competition for light and space of each individual segment within the canopy. Simple solutions are suggested for both requirements to enable an application with data that can be measured non-destructively in the field; these were stem position, tree height, crown base height, crown radii and some general crown shape information. The model was parameterised from single trees of Norway spruce and European beech and partly evaluated with independent data close to the investigation plot. Evaluations showed that the model can attribute the ecology of the different crown forms. Modelled foliage distribution for beech and spruce as well as total needle biomass of spruce agreed well with measurements but foliage biomass of beech was underestimated. The results are discussed in the context of a general model application in structured forests.     

Effects of ants on the foraging of birds in spruce trees

I experimentally excluded ants from randomly selected spruce trees Picea abies near colonies of the wood ant Formica aquilonia. Foraging activity of birds in these trees was then compared to the foraging activity of birds in neighboring spruce trees, where ants were allowed to continue foraging. Birds which foraged in the foliage showed the effects of competition with ants: they visited the trees without ants more frequently, and for longer periods. In addition, the insects and spiders that they utilized as food were more abundant in the foliage of trees without ants. Cone-foraging birds, however, which fed on seeds in cones at the tops of the trees, did not show a preference for trees without ants. The differences of tree usage between foliage-gleaning and coneforaging birds can be explained by alteration of the birds' food supply by wood ants: ants did not feed on seeds in cones, and so did not compete with cone-foraging birds. However, foraging wood ants did feed on arthropods living in the foliage, thus reducing the amount of food available to birds there.     

Predicting decay and ground vegetation development in <Emphasis Type="Italic">Picea abies</Emphasis> snag stands

In a Picea abies (L.) Karst. (Norway spruce) mountain forest on the Gandberg site in the northern Swiss Alps, trees were killed by bark beetles in 1992–1997. A combination of field studies and dynamic modelling was used to project snag decay and future ground vegetation succession in these steep, unharvested stands. In permanent plots, ground vegetation cover and natural tree regeneration have been monitored annually since 1994. To obtain additional information on the abundance of snags, logs, boulders and other microsite types in these stands, the relative frequency of the microsite types was quantified along four strip transects on the montane and subalpine elevational levels. A dynamic model of snag decay and ground vegetation development was constructed (modified matrix model approach). Based on field data and literature values, the model was parameterised and initialised separately for the montane and the subalpine level. For model validation, microsite types were quantified in 2001 with the line-intercept method on both elevational levels. Starting from the conditions in the stands before the bark beetle attacks, it was possible to project short-term succession and to accurately simulate the decay and ground vegetation patterns eight years after tree die-back. Long-term simulations suggest that on the montane level, raspberries (Rubus idaeus L.) will be replaced by Picea abies, while on the subalpine level ferns will dominate for a long time.     

Effects of changes in tree species composition on water flow dynamics – Model applications and their limitations

Water-plant relations play a key role in the water cycling in terrestrial ecosystems. Consequently, changes in tree species composition may have distinct effects on the water retention capacity as well as on the pattern of streamflow generation. Such changes may result from modified interception properties and transpiration related to differences in canopy properties and root distribution. In order to evaluate the potential hydrological effects of the current silvicultural conversion from monocultural conifer stands into mixed or pure deciduous stands the hydrological model BROOK90 was applied to two forested upland catchments in Germany. The Rotherdbach catchment (9.4 ha, 93 yr-old Norway spruce) is situated in the Eastern Ore Mountains. The Schluchsee catchment (11 ha, 55-yr-old Norway spruce) is located in the higher altitudes of the Black Forest. The calibrated model is capable to describe rather well the temporal variation of streamflow but also the portions of the individual flow components. Data for a beech scenario were adapted for each site using a standard parameter set for deciduous trees provided by BROOK90. The annual discharge in the fictional beech stand at Rotherdbach is 30 to 50% higher compared to spruce with an increase of soil moisture and especially the slow streamflow components. This mainly results from low interception rates during winter time. In contrast, the spruce stand has a permanently higher interception rate. Effects of tree species conversion are moderate at Schluchsee. The annual discharge of a fictional beech stand at Schluchsee is 7 to 14% higher compared to spruce. There in contrast to Rotherdbach, effects of tree species conversion on soil moisture dynamics are small since vertical percolation in the highly permeable soil dominates and precipitation is abundant. Practical forestry will favorably establish mixed beech–spruce rather than pure beech stands. However, it is critical to simulate mixed stands with BROOK90. Therefore, a simple summation of model results from spruce and beech according to their respective area in a fictional mixed stand can only be a first approximation. Advanced hydrological simulation of mixed stand conditions should regard interactions of tree species and spatial parameter distribution. However, this is not yet feasible due to a distinct lack of information. As a consequence, there is a strong need to collect relevant hydrological and ecophysiological data in mixed stands in the future.     

Does the azimuth orientation of Norway spruce (Picea abies/L./Karst.) branches within sunlit crown part influence the heterogeneity of biochemical,structural and spectral characteristics of needles?

The goal of this study was to determine if selected biochemical, structural and spectral properties of Norway spruce needles are influenced by the azimuth orientation of the branch. Three youngest needle age classes from 20 mature (100 years old or older) Norway spruce trees were sampled from upper branches of the sunlit production crown part from each of the 4 cardinal azimuth orientations. Photosynthetic pigments, soluble phenolic compounds and selected spectral and structural characteristics were determined for each needle age class. The content of photosynthetic pigments and soluble phenolic compounds did not differ among needles from different azimuth-oriented branches, nor did the optical reflectance indices Normalized Difference Vegetation Index (NDVI), Transformed Chlorophyll Absorption in Reflectance Index (TCARI)/Optimized Soil-Adjusted Vegetation Index (OSAVI), Red Edge Inflection Point (REIP) and Landsat Thematic Mapper bands 5 and 4 (TM5/TM4). No variation in volume properties, tissue volume proportions and cross-section shape characteristics of 3rd-year needles rejected our hypothesis that there would be variation in needle structural properties according to the azimuth orientation of branches. Consequently, we concluded that a random sampling of similar-aged needles within the sunlit production crown part may be used to study biochemical or structural and spectral needle properties of a mature Norway spruce growing in forest stands without a significant slope. In addition, the results obtained from a branch of one azimuth orientation should be representative for the whole sunlit portion of the crown. Consequences of these findings for Norway spruce health monitoring using remote sensing techniques are discussed.     

Model simulations of spatial distributions and daily totals of photosynthesis in Eucalyptus grandis canopies

Summary A simulation model for radiation absorption and photosynthesis was used to test the hypothesis that observed nonuniform distributions of nitrogen concentrations in young Eucalyptus grandis trees result in greater amounts of daily assimilation than in hypothetical trees with uniform N distributions. Simulations were performed for trees aged 6, 9, 12 and 16 months which had been grown in plantations under a factorial combination of two levels of fertilization and irrigation. Observed leaf N distribution patterns yielded daily assimilation rates which were only marginally greater (<5%) than for hypothetical trees with uniform distributions. Patterns of assimilation distribution in individual tree crowns closely resembled those for absorbed radiation, rather than for N. These conclusions were unaffected by three choices of alternative leaf area density distributions. The simulation model was also used to calculate hourly and daily rates of canopy assimilation to investigate the relative importance of radiation absorption and total canopy nitrogen on assimilation. Simulated hourly rates of carbon assimilation were often lightsaturated, whereas daily carbon gain was directly proportional to radiation absorbed by the tree crown and to total mass of N in the leaves. Leaf nitrogen concentrations determined photosynthetic capacity, whereas total leaf area determined the amount of radiation absorbed and thus the degree to which capacity was realized. Observed total leaf area and total crown N were closely correlated. The model predicted that nitrogen use efficiences (NUE, mol CO₂ mol^–1 N) were 60% higher for unfertilized than for fertilized trees at low levels of absorbed photosynthetically active radiation (PAR). Nitrogen use efficiency was dependent on fertilizer treatment and on the amount of absorbed PAR; NUE declined with increasing absorbed PAR, but decreased more rapidly for unfertilized than for fertilized trees. Annual primary productivity was linearly related to both radiation absorbed and to mass of N in the canopy.     

Variation of carbon age of fine roots in boreal forests determined from 14C measurements

Background and aims

The main objectives of this study were to determine how the carbon age of fine root cellulose varies between stands, tree species, root diameter and soil depth. In addition, we also compared the carbon age of fine roots from soil cores of this study with reported values from the roots of the same diameter classes of ingrowth cores on the same sites.

Methods

We used natural abundance of ¹⁴C to estimate root carbon age in four boreal Norway spruce and Scots pine stands in Finland and Estonia.

Results

Age of fine root carbon was older in 1.5–2 mm diameter fine roots than in fine roots with <0.5 mm diameter, and tended to be older in mineral soil than in organic soil. Fine root carbon was older in the less fertile Finnish spruce stands (11–12 years) than in the more fertile Estonian stand (3 and 8 years), implying that roots may live longer in less fertile soil. We further observed that on one of our sites carbon in live fine roots with the 1.5–2 mm diameter was of similar C age (7–12 years) than in the ingrowth core roots despite the reported root age in the ingrowth cores – being not older than 2 years.

Conclusions

From this result, we conclude that new live roots may in some cases use old carbon reserves for their cellulose formation. Future research should be oriented towards improving our understanding of possible internal redistribution and uptake of C in trees.     

Wood ants prefer conifers to broadleaved trees in mixed temperate forests

1. Wood ants, species of the subgenus Formica s. str., are one of the most important groups of insects in forest ecosystems due to their high abundance and activity.
2. We assessed the foraging pressure of Formica polyctena and Formica rufa, two dominant wood ant species in the Białowieża Forest. In addition, we compared coniferous and mixed stands in these respects.
3. In mixed stands F. polyctena visited more trees per plot than F. rufa. F. polyctena might forage a larger area due to higher population sizes, which further confirms higher foraging pressure of F. polyctena in comparison with F. rufa.
4. In our study area, host trees of the ant–aphid mutualism were larger in diameter than nonhost trees. The ants also preferred trees situated closer to their nests. In both mixed and coniferous stands, the most visited tree species was Norway spruce. Thus, we can conclude that Norway spruce seems to be preferred by the ant. The decline of living spruce in the Białowieża Forest may endanger their food source and impact the vitality of ant colonies.

     

Relationships between tree dimension and coarse root biomass in mixed stands of European beech (Fagus sylvatica L.) and Norway spruce (Picea abies[L.] Karst.)

Relationships between tree parameters above ground and the biomass of the coarse root system were examined in six mixed spruce-beech stands in the Solling Mountain region in northwest Germany. The selected stands were located on comparable sites and covered an age range of 44 to 114 years. Coarse roots (d?\ge?2 mm) of 42 spruce and 27 beech trees were sampled by excavating the entire root system. A linear model with logarithmic transformation of the variables was developed to describe the relationship between the coarse root biomass (CRB, dry weight) and the corresponding tree diameter at breast height (DBH). The coefficients of determination (R ²) attained values between 0.92 for spruce and 0.94 for beech; the logarithmic standard deviation values were between 0.29 and 0.43. A significantly different effect of tree species on the model estimates could not be detected by an analysis of co-variance (ANCOVA). For spruce, the derived relationships were similar to those reported in previous studies, but not for beech. Biomass partitioning in the tree compartments above and below ground differs significantly between spruce (coarse root/shoot ratio 0.16±0.06) and beech (coarse root/shoot ratio 0.10±0.03) in the mixed stands. These results are similar to those given in other studies involving pure spruce and beech stands on comparable sites in the region, although the ratios of pure stands in other regions growing under different site conditions are somewhat higher. Comparing trees of the same DBH classes, root/shoot ratios of spruce are 1.2 to 3 times higher than those of beech. Dominant spruce trees (DBH>60 cm) attained the highest ratios, suppressed beech trees (DBH<10 cm) the lowest. Site conditions of varying climate and soils and interspecific tree competition are likely to affect root/shoot ratio and DBH-coarse root biomass relationships. The greater variability in beech compared with spruce indicates a high 'plasticity' and adaptability of beech carbon allocation. Thus, the derived equations are useful for biomass estimates of coarse roots involving trees of different ages in mixed stands of spruce and beech in the Solling Mountains. However, application of these relationships to stands in other regions would need further testing.