Root biomass and rooting profile of naturally regenerated beech in mid-elevation Scots pine woodlands

The natural establishment of shade-tolerant forest species such as beech (Fagus sylvatica L.) occurs in naturally regenerated Scots pine (Pinus sylvestris L.) woodlands that develop on former pastures and cultivated lands. To examine possible effects of underground competition in beech establishment, we studied the root biomass and the rooting profile of 53 mixed Scots pine-natural beech woodlands in French mid-elevation volcanic areas. Stands were arranged along a maturation gradient. Roots were sampled using the root-auger technique (0 to 75 cm depth every 15 cm). In addition, 23 young beeches were uprooted to study the entire root system. Total beech fine-root biomass was closely correlated with most beech aerial characteristics, (e.g., height, diameter and girth), and correlated moderately with tree age. However, it correlated poorly with basic competition indices such as stand density and basal area. Conversely, competition indices including vertical dimensions of competing trees were correlated with the underground biomass, probably as a result of redundancy with beech height. The rooting profile (fine roots, < 5 mm) of beech and pine were quite similar, and did not change significantly along the stand maturation gradient. Beech has a heart-shaped root system while pine is more plate-like and dimorphic. Beech fine-root biomass progressively surpassed pine biomass throughout the soil layers, thus confirming that it is dynamic and competitive in mature mixed stands. The coexistence of the root systems of beech and pine in the same soil layers presumably results in strong underground competition.     

Variation in fine root biomass of three European tree species: Beech (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.), and Scots pine (Pinus sylvestris L.)

Abstract

Fine roots (<2 mm) are very dynamic and play a key role in forest ecosystem carbon and nutrient cycling and accumulation. We reviewed root biomass data of three main European tree species European beech, (Fagus sylvatica L.), Norway spruce (Picea abies L. Karst.) and Scots pine (Pinus sylvestris L.), in order to identify the differences between species, and within and between vegetation zones, and to show the relationships between root biomass and the climatic, site and stand factors. The collected literature consisted of data from 36 beech, 71 spruce and 43 pine stands. The mean fine root biomass of beech was 389 g m^?2, and that of spruce and pine 297 g m^?2 and 277 g m^?2, respectively. Data from pine stands supported the hypothesis that root biomass is higher in the temperate than in the boreal zone. The results indicated that the root biomass of deciduous trees is higher than that of conifers. The correlations between root biomass and site fertility characteristics seemed to be species specific. There was no correlation between soil acidity and root biomass. Beech fine root biomass decreased with stand age whereas pine root biomass increased with stand age. Fine root biomass at tree level correlated better than stand level root biomass with stand characteristics. The results showed that there exists a strong relationship between the fine root biomass and the above-ground biomass.     

Changes in diversity and storage function of ectomycorrhiza and soil organoprofile dynamics after introduction of beech into Scots pine forests

Diversity and storage function of mycorrhiza as well as soil organoprofile formation were investigated in a chronosequence of a pure Scots pine (Pinus sylvestris L.) stand, of Scots pine stands that were underplanted with beech (Fagus sylvatica L.) and in three pure beech stands of different age. Mycorrhiza diversity was higher in the pure beech stands compared to the pure pine stand. Beech and pine trees in the mixed stands had similar dominant mycorrhiza morphotypes. However, trees in two of the three pure beech stands were mycorrhized with other types. Mycorrhizal abundance and nutrient amounts of mycorrhizae associated with beech trees were higher in the mixed and in the pure beech stands compared to pine mycorrhizae indicating that nutrient uptake was higher in older beech than in older pine trees. Humus quality varied from pine to beech stands. Plant litter storage in the humus layer was highest in the youngest mixed stand and lowest in the oldest beech stand. Humus forms changed from moder grass-type in the pure Scots pine stand to mor-like moder and moder rich in fine humus with increasing age of beeches in the mixed stands. The older beech stands were characterised by oligomull and mull-like moder as the dominating humus forms. The ecologically favourable humus forms, i.e., nutrient rich humus forms in the older beech stands correlate well with the higher mycorrhizal diversity and abundance as well as the higher nutrient storage of their mycorrhizae in these stands. The results are also discussed with regard to the 'base-pump effect' of beech trees.     

Impact of interspecific competition and drought on the allocation of new assimilates in trees

In trees, the interplay between reduced carbon assimilation and the inability to transport carbohydrates to the sites of demand under drought might be one of the mechanisms leading to carbon starvation. However, we largely lack knowledge on how drought effects on new assimilate allocation differ between species with different drought sensitivities and how these effects are modified by interspecific competition. We assessed the fate of ¹³C labelled assimilates in above‐ and belowground plant organs and in root/rhizosphere respired CO₂ in saplings of drought‐tolerant Norway maple (Acer platanoides) and drought‐sensitive European beech (Fagus sylvatica) exposed to moderate drought, either in mono‐ or mixed culture. While drought reduced stomatal conductance and photosynthesis rates in both species, both maintained assimilate transport belowground. Beech even allocated more new assimilate to the roots under moderate drought compared to non‐limited water supply conditions, and this pattern was even more pronounced under interspecific competition. Even though maple was a superior competitor compared to beech under non‐limited soil water conditions, as indicated by the changes in above‐ and belowground biomass of both species in the interspecific competition treatments, we can state that beech was still able to efficiently allocate new assimilate belowground under combined drought and interspecific competition. This might be seen as a strategy to maintain root osmotic potential and to prioritise root functioning. Our results thus show that beech tolerates moderate drought stress plus competition without losing its ability to supply belowground tissues. It remains to be explored in future work if this strategy is also valid during long‐term drought exposure.     

Assessing fine-root biomass and production in a Scots pine stand – comparison of soil core and root ingrowth core methods

Soil core and root ingrowth core methods for assessing fine-root (< 2 mm) biomass and production were compared in a 38-year-old Scots pine (Pinus sylvestris L) stand in eastern Finland. 140 soil cores and 114 ingrowth cores were taken from two mineral soil layers (0–10 cm and 10–30 cm) during 1985–1988. Seasonal changes in root biomass (including both Scots pine and understorey roots) and necromass were used for calculating fine-root production. The Scots pine fine-root biomass averaged annually 143 g/m² and 217 g/m² in the upper mineral soil layer, and 118 g/m² and 66 g/m² in the lower layer of soil cores and ingrowth cores, respectively. The fine-root necromass averaged annually 601 g/m² and 311 g/m² in the upper mineral soil layer, and 196 g/m² and 159 g/m² in the lower layer of soil cores and ingrowth cores, respectively. The annual fine-root production in a Scots pine stand in the 30 cm thick mineral soil layer, varied between 370–1630 g/m² in soil cores and between 210 – 490 g/m² in ingrowth cores during three years. The annual production calculated for Scots pine fine roots, varied between 330–950 g/m² in soil cores and between 110 – 610 g/m² in ingrowth cores. The horizontal and vertical variation in fine-root biomass was smaller in soil cores than in ingrowth cores. Roots in soil cores were in the natural dynamic state, while the roots in the ingrowth cores were still expanding both horizontally and vertically. The annual production of fine-root biomass in the Scots pine stand was less in root ingrowth cores than in soil cores. During the third year, the fine-root biomass production of Scots pine, when calculated by the ingrowth core method, was similar to that calculated by the soil core method. Both techniques have sources of error. In this research the sampling interval in the soil core method was 6–8 weeks, and thus root growth and death between sampling dates could not be accurately estimated. In the ingrowth core method, fine roots were still growing into the mesh bags. In Finnish conditions, after more than three growing seasons, roots in the ingrowth cores can be compared with those in the surrounding soil. The soil core method can be used for studying both the annual and seasonal biomass variations. For estimation of production, sampling should be done at short intervals. The ingrowth core method is more suitable for estimating the potential of annual fine-root production between different site types.     

Contrasting fine-root production,survival and soil CO2 efflux in pine and poplar plantations

Tree root activity, including fine-root production, turnover and metabolic activity are significant components of forest productivity and nutrient cycling. Differences in root activity among forest types are not well known. A 3-year study was undertaken in red pine (Pinus resinosa Ait.) and hybrid poplar (Populus tristis X P. balsamifera cv `Tristis no. 1') plantations to compare belowground root dynamics. We measured fine-root production, mortality and standing crop, as well as soil CO₂ efflux. Pine fine-root production was only 2.9% of that of poplar during three years; 85 pine roots were observed in minirhizotron tubes compared with 4088 poplar roots. Live-root density oscillated seasonally for both species with late winter minimum and autumn maximum. Poplar reached constant maximum live-root length within the first growing season, but pine continued to increase observed fine-root length for three growing seasons. Within the first 100 days following initial appearance, 22% of the pine roots disappeared and 38% of the poplar roots disappeared. Median fine-root longevity of pine was 291 days compared with 149 days for poplar roots. Fine-root longevity increased with depth in the soil, and was greater for roots with initial diameter >0.5 mm. The probability of poplar root death from late February to May was more than three times that in any other season, regardless of root age. Despite the greater poplar root production and live-root length, fine-root biomass and soil CO₂ efflux was greater in pine. Greater metabolic activity in the pine stand may be due to greater fine-root biomass or greater heterotrophic respiration. This revised version was published online in June 2006 with corrections to the Cover Date.     

Fine-root seasonal pattern,production and turnover rate of European beech (Fagus sylvatica L.) stands in Italy Prealps: Possible implications of coppice conversion to high forest

Abstract

The aim of this study was to investigate the possible effects of coppice conversion to high forest on the beech fine-root systems. We compared the seasonal pattern of live and dead fine-root mass (d < 2 mm), production and turnover in three beech stands that differed in management practices. Tree density was higher in the 40-year-old coppice stand than in the stands that were converted from coppice to high forest in 1994 and 2004, respectively. We found that a reduction in tree density reduced the total fine-root biomass (Coppice stand, 353.8 g m^?2; Conversion 1994 stand, 203.6 g m^?2; Conversion 2004 stand, 176.2 g m^?2) which continued to be characterised by a bimodal pattern with two major peaks, one in spring and one in early fall. Conversion to high forest may also affect the fine-root soil depth distribution. Both fine-root production and turnover rate were sensitive to management practices. They were lower in the Coppice stand (production 131.5 g m^?2 year^?1; turnover rate 0.41 year^?1) than in the converted stands (1994 Conversion stand: production 232 g m^?2 year^?1, turnover rate 1.06 year^?1; 2004 Conversion stand: production 164.2 g m^?2 year^?1, turnover rate 0.79 year^?1).     

Drought effects on fine-root and ectomycorrhizal-root biomass in managed Pinus oaxacana Mirov stands in Oaxaca, Mexico

The effects of a severe drought on fine-root and ectomycorrhizal biomass were investigated in a forest ecosystem dominated by Pinus oaxacana located in Oaxaca, Mexico. Root cores were collected during both the wet and dry seasons of 1998 and 1999 from three sites subjected to different forest management treatments in 1990 and assessed for total fine-root biomass and ectomycorrhizal-root biomass. Additionally, a bioassay experiment with P. oaxacana seedlings was conducted to assess the ectomycorrhizal inoculum potential of the soil for each of the three stands. Results indicated that biomasses of both fine roots and ectomycorrhizal roots were reduced by almost 60% in the drought year compared to the nondrought year. There were no significant differences in ectomycorrhizal and fine-root biomass between the wet and dry seasons. Further, the proportion of total root biomass consisting of ectomycorrhizal roots did not vary between years or seasons. These results suggest that both total fine-root biomass and ectomycorrhizal-root biomass are strongly affected by severe drought in these high-elevation tropical pine forests, and that these responses outweigh seasonal effects. Forest management practices in these tropical pine forests should consider the effects of drought on the capacity of P. oaxacana to maintain sufficient levels of ectomycorrhizae especially when there is a potential for synergistic interactions between multiple disturbances that may lead to more severe stress in the host plant and subsequent reductions in ectomycorrhizal colonization.     

Short-term effects of manipulated increase in acid deposition on soil, soil solution chemistry and fine roots in Scots pine (Pinus sylvestris) stand on a podzol

A manipulated increase in acid deposition (15 kg S ha⁻¹), carried out for three months in a mature Scots pine (Pinus sylvestris) stand on a podzol, acidified the soil and raised dissolved Al at concentrations above the critical level of 5 mg l⁻¹ previously determined in a controlled experiment with Scots pine seedlings. The induced soil acidification reduced tree fine root density and biomass significantly in the top 15 cm of soil in the field. The results suggested that the reduction in fine root growth was a response not simply to high Al in solution but to the depletion of exchangeable Ca and Mg in the organic layer, K deficiency, the increase in NH₄:NO₃ ratio in solution and the high proton input to the soil by the acid manipulation. The results from this study could not justify the hypothesis of Al-induced root damage under field conditions, at least not in the short term. However, the study suggests that a short exposure to soil acidity may affect the fine root growth of mature Scots pine.     

An Analysis of Vegetation Restoration on Opencast Oil Shale Mines in Estonia

We compared four types of 30‐year‐old forest stands growing on spoil of opencast oil shale mines in Estonia. The stand types were: (1) natural stands formed by spontaneous succession, and plantations of (2) Pinus sylvestris (Scots pine), (3) Betula pendula (silver birch), and (4) Alnus glutinosa (European black alder). In all stands we measured properties of the tree layer (species richness, stand density, and volume of growing stock), understory (density and species richness of shrubs and tree saplings), and ground vegetation (aboveground biomass, species richness, and species diversity). The tree layer was most diverse though sparse in the natural stands. Understory species richness per 100‐m² plot was highest in the natural stand, but total stand richness was equal in the natural and alder stands, which were higher than the birch and pine stands. The understory sapling density was lower than 50 saplings/100 m² in the plantations, while it varied between 50 and 180 saplings/100 m² in the natural stands. Growing stock volume was the least in natural stands and greatest in birch stands. The aboveground biomass of ground vegetation was highest in alder stands and lowest in the pine stands. We can conclude that spontaneous succession promotes establishment of diverse vegetation. In plantations the establishment of diverse ground vegetation depends on planted tree species.     

Nutritional disturbances of young Scots pines caused by pine bark bugs in a dry heath forest

This study deals with the effect of pine bark bugs (Aradus cinnamomeus Panzer) on the nutrition of young Scots pines (Pinus sylvestris L.). Soil and needle samples for analytical purposes were collected from a young pine stand growing on a dry barren mineral soil afflicted by pine bark bugs.The damage to vascular tissues caused by pine bark bugs disturbed the nutrition of the trees, especially in the top part of the crown. The foliar calcium, magnesium, manganese and sulphur concentrations were highly reduced. Scots pines suffered from a lack of calcium, magnesium, nitrogen and phosphorus. These deficiencies were secondary by nature, because no differences were observed between the nutrient concentrations of the underlying soil of the healthy and affected trees. The symptoms of trees damaged by pine bark bugs resembled most of all calcium deficiency symptoms.     

Survival of black alder (Alnus glutinosa L.), silver birch (Betula pendula Roth.) and Scots pine (Pinus sylvestris L.) seedlings in a reclaimed oil shale mining area

Early survival and growth of black alder, silver birch and Scots pine were investigated on reclaimed extremely stony and heterogeneous calcareous (pH 8) opencast oil shale mining areas (OOSMAs). Biomass allocation, production, leaf and root adaptations, and mineral nutrition in relation to tree species and soil heterogeneity were analysed. The adaptive strategies of tree species in first-year plantations on OOSMA were different. Scots pine allocated 1.5–2 times more biomass into leaves and fine roots than deciduous trees. The lower leaf/fine root biomass ratio was in proportion to the better survival (%) of seedlings, decreasing in the following order: black alder (93%) ≥ Scots pine (83%) > silver birch (64%). Deciduous trees improved mineral nutrition more by fine-root morphological adaptations than Scots pine; e.g. the mean specific root length (SRL, m g^?1) of short roots increased in the following order: Scots pine (62) < black alder (172) < silver birch (314). The effect of soil heterogeneity on growth and adaptations was minor. All studied species suffered from P and N, and deciduous species also from K deficiency. In the first year after planting, black alder was best adapted to the harsh conditions of the post-mining substrate. The approaches of this study can be used for other regions where wastelands require reclamation.     

Morphology,biomass and nutrient status of fine roots of Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis>) as influenced by seasonal fluctuations in soil moisture and soil solution chemistry

A field monitoring study was carried out to follow the changes of fine root morphology, biomass and nutrient status in relation to seasonal changes in soil solution chemistry and moisture regime in a mature Scots pine stand on acid soil. Seasonal and yearly fluctuations in soil moisture and soil solution chemistry have been observed. Changes in soil moisture accounted for some of the changes in the soil solution chemistry. The results showed that when natural acidification in the soil occurs with low pH (3.5–4.2) and high aluminium concentration in the soil solution (>3–10 mg l^–1), fine root longevity and distribution could be affected. However, fine root growth of Scots pine may not be negatively influenced by adverse soil chemical conditions if soil moisture is not a limiting factor for root growth. In contrast, dry soil conditions increase Scots pine susceptibility to soil acidification and this could significantly reduce fine root growth and increase root mortality. It is therefore important to study seasonal fluctuations of the environmental variables when investigating and modelling cause-effect relationships.     

Comparison of Fine Root Dynamics in Scots Pine and Pedunculate Oak in Sandy Soil

In this study, we investigated the relationship between the seasonality of vegetation cover and that of fine root processes in a man-made forest in northern Belgium. Due to their contrasting foliar development, we expected different seasonal patterns of fine root growth and standing biomass between Pedunculate oak (Quercus robur L.), and Scots pine (Pinus sylvestris L.). Biomass and necromass of fine and small roots were estimated by repeated core sampling in February, April, June, August and October 2003. Measurements showed that Pedunculate oaks maintained more live fine roots in winter than Scots pines. However, Scots pines produced more than twice as much fine roots in spring, such that in summer both species had similar root mass. Scots pine root production started before-, but declined during leaf unfolding. Pedunculate oak roots, in contrast, started elongating only after bud break. For both species, fine root production peaked in JuneJuly, but was more than offset by drought-induced mortality at the end of July and early August. Summer drought in 2003 was exceptionally long and intense, significantly reducing leaf area, killing most new roots, and inhibiting root decomposition, such that the obtained results cannot be typical for this forest.     

岷江上游4个栽培树种细根功能性状垂直分布的差异性

细根(直径≤2 mm)功能性状及垂直分布格局是反映植物对土壤资源吸收策略和影响森林地下生态过程的关键。本研究以岷江上游4个人工林树种连香树(Cercidiphyllum japonicum)、白桦(Betula platyphylla)、华山松(Pinus armandii)和油松(P.tabuliformis)为对象,调查不同海拔树木细根功能性状及其在不同土层间的垂直分布格局,并分析细根功能性状分布与构型之间的相关关系。结果表明:阔叶树种比针叶树种有更大的根长密度、生物量、比根长和比表面积,而直径反之;4个树种细根集中在0~20 cm土层,根长密度和生物量在较高海拔地段均显著大于较低海拔,且均随土壤深度增加而减少,但比根长、比表面积和直径无显著的海拔差异,随土层加深也无明显的垂直变化规律;针阔树种间的细根构型差异显著,但不受海拔差异的影响,阔叶树的细根分支强度与一级根数量显著大于针叶树种;一级根数和根尖密度与比根长以及分根比与根长密度和生物量均呈显著正相关,而分叉与几个细根功能参数均呈负相关;随着土层深度增加,细根总生长量明显减少,但细根资源利用效率和策略不变;5个细根功能性状的垂直分布格局差异明显,且受树种特性影响显著,细根分支构型对其垂直分布及资源获取策略具有显著影响。     

Fertilization in a mature Scots pine (Pinus sylvestris L.) stand—effects on fine roots

The effects of liquid and solid fertilizers on fine-root development were studied in a 130-year-old Scots pine (Pinus sylvestris L.) stand. Ingrowth cores,viz. initially root-free mesh bags with sieved mineral soil taken outside the plots and driven to a depth of 30 cm from the soil surface, were subsequently resampled and the amount of fine roots was estimated. The total accretion of both fine-root length and dry weight was comparatively high in the liquid fertilization plot. The most substantial net accretion, however, during the 1984 period was in the control plot. The results of the study is that the ramification pattern of the fine roots was strongly influenced by fertilization. The average number of root tips per unit length was 9.9 cm⁻¹ in the control plot cm⁻¹, compared with both the solid (A and B) and liquid fertilization plots (2.3, 3.2 and 3.3 cm⁻¹, respectively) due to a greater occurrence of mycorrhizal aggregates (‘ball mycorrhiza’). The effects of fertilization on the mineral nutrient concentrations in the fine roots were limited—the strongest effects were in the liquid-fertilized area. The observed increase in the concentration of most mineral nutrients in the latter experimental area, however, may be due to a change in the growth pattern of the fine roots rather than an effect of the fertilizers themselves.     

Testing the enemies hypothesis in forest stands: the important role of tree species composition

Numerous studies conducted in agro-ecosystems support the enemies hypothesis, which states that predators and parasites are more efficient in controlling pest densities in polycultures than in monocultures. Few similar studies, however, have been conducted in forest ecosystems, and we do not yet have evidence as to whether the enemies hypothesis holds true in forests. In a 2-year study, we investigated whether the survival of autumnal moth (Epirrita autumnata) larvae and pupae differs between silver birch monocultures and two-species mixtures of birch with black alder, Norway spruce and Scots pine. We placed young larvae on birch saplings and monitored their survival until the end of the larval period, when we checked whether they had been parasitized. After the larvae had pupated, pupal survival was tested in a field trial. In 2002, the larvae disappeared earlier and their overall survival was lower in birch–pine mixtures than in other stand types. In 2003, survival probability was lowest in birch–pine stands only during the first week and there were no differences between stands in overall survival. Larval parasitism was not affected by tree species composition. Pupal weight and pupal survival were likewise not affected by stand type. Among the predators, wood ants were more abundant on birches growing in birch–pine mixtures than in other stand types probably because colonies of myrmecophilic aphids were common on pines. In contrast, spider numbers did not differ between stand types. Ant exclusion by means of a glue ring around the birch trunk increased larval survival, indicating that ants are important predators of the autumnal moth larvae; differences in larval survival between stands are probably due to differential ant predation. Our results provide only partial support for the enemies hypothesis, and suggest that it is both tree species composition and species diversity which affect herbivore survival and predation.     

Are fine roots of both shrubs and perennial grasses able to occupy the upper soil layer? A case study in the arid Patagonian Monte with non-seasonal precipitation

We tested whether both shrubs and grasses are able to develop similar active fine-root systems in the upper soil layer of the arid Patagonian Monte ecosystem with non-seasonal precipitation. We selected in the field shrub patches consisting of one isolated modal plant of the dominant shrub Larrea divaricata Cav., grass patches formed by one or more bunches of the dominant grass Stipa tenuis Phil. (15 cm diameter), and mixed patches consisting of one individual of L. divaricata with bunches of S. tenuis under its canopy. We assessed the biomass, regrowth, and activity of fine roots (diameter <1.4 mm) of each species in the upper soil (50 cm depth) of each patch type at 3-month intervals. We also measured the N concentration in fine roots to estimate the relative contribution of each species to fine-root biomass of mixed patches. We injected Li⁺ in the soil as a chemical tracer to detect fine-root activity of each species in the upper soil. Fine-root biomass was higher in mixed patches than in grass patches while fine-root biomass in shrub patches did not differ from the two former. We did not find differences in fine-root regrowth among patch types. Li⁺ injection provided evidence of active fine roots of both species in the upper soil when it was wet. N concentration in fine roots suggested the prevalence of fine roots of L. divaricata in the upper soil of mixed patches. Our results support evidence of the ability of fine roots of both the shrub and the grass species to occupy the upper soil. These findings did not support the two-layer model (H Walter, Ecology of tropical and subtropical vegetation, Oliver and Boyd, Edinburgh, 1971) and provide evidence of this model would be less applicable to arid ecosystems with non-seasonal precipitation. Further, our results highlighted some issues deserving more research such as the outcome of belowground competition between neighboring plants of both contrasting life forms, the eventual limited fine-root carrying capacity of the upper soil, and differences in fine-root lifespan between species of both contrasting life form.     

Biomass responses to elevated CO<Subscript>2</Subscript>, soil heterogeneity and diversity: an experimental assessment with grassland assemblages

While it is well-established that the spatial distribution of soil nutrients (soil heterogeneity) influences the competitive ability and survival of individual plants, as well as the productivity of plant communities, there is a paucity of data on how soil heterogeneity and global change drivers interact to affect plant performance and ecosystem functioning. To evaluate the effects of elevated CO₂, soil heterogeneity and diversity (species richness and composition) on productivity, patterns of biomass allocation and root foraging precision, we conducted an experiment with grassland assemblages formed by monocultures, two- and three-species mixtures of Lolium perenne, Plantago lanceolata and Holcus lanatus. The experiment lasted for 90 days, and was conducted on microcosms built out of PVC pipe (length 38 cm, internal diameter 10 cm). When nutrients were heterogeneously supplied (in discrete patches), assemblages exhibited precise root foraging patterns, and had higher total, above- and belowground biomass. Greater aboveground biomass was observed under elevated CO₂. Species composition affected the below:aboveground biomass ratio and interacted with nutrient heterogeneity to determine belowground and total biomass. Species richness had no significant effects, and did not interact with either CO₂ or nutrient heterogeneity. Under elevated CO₂ conditions, the two- and three-species mixtures showed a clear trend towards underyielding. Our results show that differences among composition levels were dependent on soil heterogeneity, highlighting its potential role in modulating diversity–productivity relationships. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible to authorized users.     

Evaluation of different approaches for modelling individual tree seedling height growth

We compared different approaches for modelling height growth of individual beech seedlings in a controlled factorial experiment as well as in field data from naturally regenerated beech seedlings under the canopy of overstorey mature beech trees. Several competition indices, a model of overstorey fine root density, relative photosynthetically active radiation (PAR) values, and soil water values were used in these approaches. In the factorial experiment relative PAR and soil water content were measured and used for the prediction of seedlings height growth. In the field experiment this was done by using relative PAR and estimated fine root biomass as a surrogate for below ground resource availability. The latter approach was compared with a model where we used various competition indices representing the impact of overstorey trees on beech seedlings. Our results suggested that (1) models which combine resource based growth functions are suitable for the prediction of individual height growth of beech seedlings. Resource based models offer the opportunity to investigate on the independent multiplicative effect of irradiance and water supply and their interactions on tree seedlings. It was (2) shown that a combined model could be used not only to predict individual height growth of beech seedlings in a controlled experiment but also in the field. The model parameters of a pure light response function for the controlled factorial experiment are comparable to those obtained in the field study. The results showed (3) that the precision of predicting beech seedlings height growth is comparable between the model types tested within this study. Approximately half of the observed variation in seedlings relative height growth rate could be explained. However, the simple competition index approach provides no information on the environmental factors constraining tree seedlings growth; whereas the multiplicative combined models can be used to get a better understanding of growth dynamics in the field.