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.     

Carbon,nitrogen, phosphorus,and sulphur concentration and partitioning in beech ecotypes (Fagus sylvatica L.): phosphorus most affected by drought

Beech seedlings of different drought sensitivity originating from 11 German provenances were grown in pots and cultivated in a greenhouse. The present paper aims to give insights on uptake, transport and use of macronutrients, since the knowledge of drought effects on the nutrition of trees is low compared to water relations. Therefore, the elemental composition, the ratio of inorganic to total content, and the partitioning between roots and shoots of carbon, nitrogen, phosphorus, and sulphur were investigated as affected by provenance and drought treatment. Phosphorus and phosphate concentrations decreased in all tissues after a 3 week drought treatment simulating a summer drought period. In roots carbon increased and nitrate decreased, in stems nitrogen decreased but nitrate increased following drought. The observed effects on phosphorus and phosphate are discussed in terms of lower phosphate mobility in the substrate due to lower water availability. The decrease in the ratio of phosphate to phosphorus in the tissues suggests the use of vacuolar phosphate pools for maintaining organic phosphorus homeostasis. The partitioning of all macronutrients was not affected by drought, although phosphorus and phosphate were significantly lowered in tissues. In most of the parameters studied significant differences between provenances were found. The recently observed drought sensitivity of provenances was not reflected in the strength of concentration changes or partitioning of macronutrients by drought over provenances.     

Modelling variability of wood density in beech as affected by ring age,radial growth and climate

Although it has been recognized as a key parameter of wood quality and a good source of information on growth, annual wood density has been little studied within diffuse-porous trees such as beech ( Fagus sylvatica Liebl.). In this paper we examine the variability encountered in beech ring density series and analyze the influences of ring age, ring width, climate and between-tree variability on density. Thirty ring sequences were sampled from 55-year- old dominant beech trees growing within the same stand; ring density and width were measured using radiography. Ring density proved to be less variable through time than ring width. The relationship between these two variables was less than observed in ring-porous trees and it showed great variation between trees. The sensitivity of ring width and density to climate was also different; width was strongly linked to soil water deficit whereas density was correlated to temperature and August rainfall. Unlike ring width, wood density showed sensitivity towards climatic characteristics of the late growing season. A large part of annual density variability remains unexplained, even using advanced modelled water balance variables. We hypothesize that a significant part of the tree ring is under internal control. We also demonstrated great inter-tree variability (the tree effect) in ring density, which has an influence on density but not on trees response to climate.     

Effect of soil strength on root growth under different water conditions

The objective of this study was to determine the effects of soil water and soil strength on root growth in situations where the individual effects of both of these factors were important. Three grain legumes were grown from pre-germinated seeds for five days on 50-mm compacted columns of two major soils of Sri Lanka. Four or five levels of bulk density (1.1 to 1.8 Mg.m^–3) and five or six levels of matric potential (–0.02 to–2.0 MPa) were used.Soil strength and matric potential effects on root growth were independently significant for most crop and soil combinations. Under high (wet) matric potential (>–0.77 MPa) soil conditions, the effect of soil water on root growth was evident only in its effect on soil strength. Bulk density had a significant effect on root growth independent of soil strength and matric potential in three cases.For all crops and soils, root penetration was 80% of the maximum or greater when the average soil strength (soil water not limiting) was 0.75 MPa or less, and when the average matric potential (soil strength not limiting) was –0.77 MPa or greater (wetter). Root penetration was 20% of the maximum or less when the soil strength was greater than 3.30 MPa (soil water not limiting), and when matric potential (soil strength not limiting) was less than –3.57 MPa. The use of pre-germinated seeds, which contained imbibed water, combined with a lack of water loss from the closed chambers containing the plants is the probable cause for the very low (–3.57 MPa) matric potential that allowed root growth at 20% of the maximum.     

Mapping QTLs for root traits under different nitrate levels at the seedling stage in maize (Zea mays L.)

Nitrogen (N) loss is a worldwide problem in crop production. Apart from reasonable N fertilizer application, breeding N efficient cultivars provides an alternative way. Root architecture is an important factor determining N acquisition. However, little is known about the molecular genetic basis for root growth in relation to N supply. In the present study, an F₈ maize (Zea may L.) recombinant inbred (RI) population consisting of 94 lines was used to identify the QTLs for root traits under different nitrate levels. The lateral root length (LRL), axial root length (ARL), maximal axial root length (MARL), axial root number (ARN) and average axial root length (AARL) were evaluated under low N (LN) and high N (HN) conditions in a hydroponics system. A total of 17 QTLs were detected among which 14 loci are located on the same chromosome region as published QTLs for root traits. A major QTL on chromosome 1 (between bnlg1025 and umc2029) for the AARL under LN could explain 43.7% of the phenotypic variation. This QTL co-localizes with previously reported QTLs that associate with root traits, grain yield, and N uptake. Our results indicate that longer axial roots are important for efficient N acquisition and the major QTL for AARL may be used as a marker in breeding N efficient maize genotypes.     

Report on non-temperature related variations in CO<Subscript>2</Subscript> efflux rates from young tree stems in the dormant season

Respiration rates are reported to increase exponentially with temperature. Respiration rates of woody tissues are commonly measured as CO₂ efflux rates () from that tissue. However, this paper describes clear variations in stem that were not related to temperature for the case of a young beech (Fagus sylvatica L.) and oak (Quercus robur L.) tree during the dormant season. The CO₂ concentration ([CO₂]) in the xylem of the beech tree showed similar temperature-independent variations. The trees were grown in a growth chamber in which radiation patterns and temperature were kept constant. was measured with an IRGA connected to cuvettes surrounding a stem segment. Xylem [CO₂] was measured in situ using a CO₂ microelectrode. Depressions in and [CO₂] occurred during the light period, despite equal temperatures in the light and dark period. Explanations found in literature for discrepancies in the exponential relationship between temperature and are the influence of (1) sap flow or (2) decreased cell water content. However, (1) the variations were observed in the dormant season, when no sap flow was observed yet, and (2) reduced cell water content was not likely to be apparent as differences in stem transpiration rates between the dark and light period were not significant. Hence, previously formulated theories failed to explain our results. This work therefore provides a new ground for discussion on other possible causes of daytime depressions in . One might be the refixation of respired CO₂ by corticular photosynthesis in the stem parts adjacent to the stem segment enclosed by the cuvette.     

Optimized nonradioisotopic amplified fragment length polymorphism method for European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.)

The amplified fragment length polymorphism (AFLP) technique has increasingly been used for the study of forest tree species. A nonradioisotopic AFLP method was optimized for European beech (Fagus sylvatica L.) and found reproducible. However, type ofTaq DNA polymerase and choice of primers must be considered for a consistent AFLP pattern.     

Natural regeneration of forest related to the spatial structure of trees: A study of two forest communities in Western Carpathians,southern Poland

The existence of a relationship between the spatial pattern of trees and the distribution of young individuals beneath the canopy has been tested in the beech (Fagus sylvatica) and spruce (Picea abies) — fir (Abies alba) forests in the mountainous region, using two different methods. The first method was the analysis of spatial pattern of individuals, the second one was based on calculating sums of influences of all trees occurring within analysed plot on a given point on the forest floor. Results of spatial pattern analyses were surprisingly consistent: almost all mature trees and seedlings didplayed a random pattern of spatial arrangement. However, there is a clear, although statistically insignificant tendency towards uniformity of spatial pattern with increasing sizes of analysed trees. Results of comparing sums of influences on regularly distributed points with sums of influences on seedlings or saplings revealed no tendency in forest regeneration to concentrate in places, where the sums were smaller than the average for a plot. This, coupled with the dominance of random spatial pattern of trees, suggests, that viewed on a small spatial scale, influence of competition among forest trees on their spatial arrangement is obscured by other factors, which are not closely related to the distribution of individuals.     

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.     

Depth distribution and composition of seed banks under different tree layers in a managed temperate forest ecosystem

In the present work we examined the composition and distribution across three soil layers of the buried soil seed bank under three different overstory types (Fagus sylvatica, Quercus robur, Pinus sylvestris) and in logging areas in a 4383-ha forest in central Belgium. The objectives were: (1) to investigate whether species composition and species richness of soil seed banks are affected by different forest stands; (2) to examine how abundant are habitat-specific forest species in seed banks under different planted tree layers. The study was carried out in stands which are replicated, managed in the same way (even-aged high forest), and growing on the same soil type with the same land-use history. In the investigated area, the seed bank did show significant differences under oak, beech, pine and in logging areas, respectively in terms of size, composition and depth occurrence. All species and layers taken together, the seed bank size ranked as follows: oakwood > beechwood > logging area > pinewood. The same pattern was found for forest species. Seed numbers of Betula pendula, Calluna vulgaris, Dryopteris dilatata and Rubus fruticosus were significantly higher under the beech canopy. Carex remota, Impatiens parviflora and Lotus sp. showed a significantly denser seed bank in logging areas, while Digitalis purpurea seeds were significantly more abundant in soils under the oak canopy. The fact that the seed bank of an originally homogeneous forest varies under different planted stands highlights that a long period of canopy conversion can affect the composition and depth of buried seeds.     

Root growth inhibition effects of holorganic moder humus layer under spruce (Picea abies KARST.) and beech (Fagus sylvatica L.)

Field observations demonstrated inhibition of root growth by moder humus material of the Of₂ and Oh-layers under beech and spruce. Growth chamber experiments with spruce seedlings showed that root growth on a natural Of₂-substrate is ten times lower than on resin-quartz mixtures. Added phenolic acid (protocatechic a.) inhibited root growth only in sterilized substrate. Leachates of Of₂-material inhibited root growth in a mineral substrate devoid of absorption complex (pure quartz sand), but not in a resin-quartz mixture. It is concluded that inhibiting substances are probably ionized molecules that can withstand biodegradation.     

Mapping QTLs of root morphological traits at different growth stages in rice

Roots are a vital organ for absorbing soil moisture and nutrients and influence drought resistance. The identification of quantitative trait loci (QTLs) with molecular markers may allow the estimation of parameters of genetic architecture and improve root traits by molecular marker-assisted selection (MAS). A mapping population of 120 recombinant inbred lines (RILs) derived from a cross between japonica upland rice 'IRAT109' and paddy rice 'Yuefu' was used for mapping QTLs of developmental root traits. All plant material was grown in PVC-pipe. Basal root thickness (BRT), root number (RN), maximum root length (MRL), root fresh weight (RFW), root dry weight (RDW) and root volume (RV) were phenotyped at the seedling (I), tillering (II), heading (III), grain filling (IV) and mature (V) stages, respectively. Phenotypic correlations showed that BRT was positively correlated to MRL at the majority of stages, but not correlated with RN. MRL was not correlated to RN except at the seedling stage. BRT, MRL and RN were positively correlated to RFW, RDW and RV at all growth stages. QTL analysis was performed using QTLMapper 1.6 to partition the genetic components into additive-effect QTLs, epistatic QTLs and QTL-by-year interactions (Q x E) effect. The results indicated that the additive effects played a major role for BRT, RN and MRL, while for RFW, RDW and RV the epistatic effects showed an important action and Q x E effect also played important roles in controlling root traits. A total of 84 additive-effect QTLs and 86 pairs of epistatic QTLs were detected for the six root traits at five stages. Only 12 additive QTLs were expressed in at least two stages. This indicated that the majority of QTLs were developmental stage specific. Two main effect QTLs, brt9a and brt9b, were detected at the heading stage and explained 19% and 10% of the total phenotypic variation in BRT without any influence from the environment. These QTLs can be used in breeding programs for improving root traits.     

Copper uptake by some forest tree species from an acid sandy soil

Summary The behaviour of some young conifers and broadleaves on a very acid sandy humuspodzol soil was studied in pot trials. Species differed considerably in foliar copper concentration, reaction to N fertilization and development of Cu deficiency symptoms. Foliar copper concentrations were highest for the broadleaves (esp. oak and beech), and was increased by Cu and N fertilization, the last reaction showing an optimum curve. Cu deficiency symptoms could easily be induced in the conifers but not in the broadleaves.     

Manganese dynamics in the rhizosphere and Mn uptake by different crops evaluated by a mechanistic model

Understanding of the mechanisms of Mn supply from the soil and uptake by the plants can be improved by using simulation models that are based on basic principles. For this, a pot culture experiment was conducted with a sandy clay loam soil to measure Mn uptake by summer wheat (Triticum aestivum L. cv. Planet), maize (Zea mays L. cv. Pirat) and sugar beet (Beta vulgaris L. cv. Orbis) and to simulate Mn dynamics in the rhizosphere by means of a mechanistic model. Seeds of three crops were sown in pots containing 2.9 kg soil in a controlled growth chamber. Root and shoot weight, Mn content of plants, root length and root radius were determined 8 (13 days in case of sugar beet) and 20 days after germination. Soil and plant parameters were determined to run nutrient uptake model calculations. Manganese content of the shoot varied from 25 mg kg^-1 for sugar beet to 34 mg kg^-1 for maize. Sugar beet had the lowest root length/shoot weight ratio but the highest relative shoot growth rate, resulting in the highest shoot demand on the root. This is reflected by the Mn influx which was 0.9 × 10^-7, 1.7 × 10^-7 and 2.5 × 10^-7 nmol cm^-1 s^-1 for wheat, maize and sugar beet, respectively. Nutrient uptake model calculations predicted similar influx values. Initial Mn concentration of 0.2 μM in the soil solution decreased to only 0.16 μM for wheat, 0.13 μM for maize and 0.11 μM for sugar beet at the root surface. This shows that manganese transport to the root was not a limiting step. This was confirmed by the fact that an assumed 20 times increase in maximum influx (I_max) increased the calculated Mn influx by 3.7 times. Sensitivity analysis demonstrated that for controlling Mn uptake the initial soil solution concentration (C _Li), the root radius (r₀), I_max and the Michaelis constant (K _m) were the most sensitive factors in the listed order. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effect of temperature on seedling growth under impeding conditions

Studies were carried out under controlled laboratory conditions to evaluate the seedling growth capacities of chickpea (Cicer arietinum L.) varieties Pusa 209 and H208 at constant temperatures of 15, 20, 25 and 28°C (±0.5°C) and of pigeonpea (Cajanus cajan L.) variety Prabhat at 20 and 28°C (±0.5°C). Seedling growth at any given time was found to depend on ‘a’, the growth at no impedance, and ‘b’, the impedance growth factori.e., decrease in growth with increase in impedance, and on temperature. The optimum temperature for chickpea was found to be in the range of 20 to 24°C for better seedling growth characteristics, whereas for pigeonpea, 28°C was found to be more congenial than 20°C. Chickpea varieties differed in their response to temperature. Part of the M. Sc. thesis of first author submitted to the Indian Agricultural Research Institute, New Delhi-110012, India.     

Soil modification by different tree species influences the extent of seedling ectomycorrhizal infection

Established vegetation can facilitate the ectomycorrhizal infection of seedlings, but it is not known whether this interaction is limited by the phylogenetic relatedness of trees and seedlings. We use a series of bioassay experiments to test whether soil modification by different ectomycorrhizal tree species causes different levels of seedling infection, whether the extent of seedling infection is a function of the relatedness of tree and seedling, and whether the effect of trees on seedlings is mediated by biotic or abiotic soil factors. We found that soils from under different tree species do vary in their mycorrhizal infectiveness. However, this variation is not related to the genetic relatedness of trees and seedlings but instead, appears to be an attribute of the overstory species, irrespective of seedling species, mediated through a suite of humus- and base-cation-related abiotic effects on soils. Modification of abiotic soil properties by overstory trees should be considered as an important factor in the effect of different overstory trees on the extent of seedling mycorrhizal infection.     

Evaluation of the mobility and discrimination of Ca, Sr and Ba in forest ecosystems: consequence on the use of alkaline-earth element ratios as tracers of Ca

A comprehensive understanding of Ca cycling in an ecosystem is desirable because of the role of this element in tree mineral nutrition and its status as a major base cation on the soil exchange complex. The determination of the origin of Ca in forests is particularly indicated in regard of important changes linked to acid inputs and intensive logging. Natural strontium isotopes are increasingly used as tracers of Ca in forest ecosystems for qualitative and quantitative assessments. Nevertheless this method is limited to relatively simple systems with two sources of nutrients. Some recent studies coupled Sr/Ca or Sr/Ba ratios to Sr isotopic measurements in order to solve more complex systems. Such method has however associated with it some uncertainties: this approach assumed that Ca, Sr and Ba behave similarly throughout the ecosystem and does not take into account the Ca biopurification processes occurring in some tree’s organs which can alter element ratio. The present work focuses on two deciduous species covering large areas in Europe: European beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.). In order to test the similarity of behaviour between Ca, Sr and Ba, their concentrations were measured extensively in the major compartments of two forest ecosystems. In parallel, the discrimination process inside tree organs was studied in 23 stands for beech and 10 stands for oak. We found that Sr and Ca behave similarly in all soil and tree compartments. By contrast, Ba and Ca appear to have contrasting behaviours, especially in streams, soil solution and soil exchange complex (no correlations between element concentrations). Sr/Ba and Ba/Ca ratios must therefore be used with care as tracer of Ca. The Ca biopurification is absent in roots and slight in bole wood but is large in bark, twigs and leaves. The discrimination factors (DF) between wood and leaves are characteristic of the two species studied and do not change significantly as a function of the soil Ca status (acidic or calcareous soils). Therefore, strontium–calcium DF can be used as a correction factor of the Sr/Ca ratio of leaves when this ratio is used in connection with Sr isotopic ratios. This correction allows to solve systems of tree nutrition with more than two sources of Ca.     

In vitro adventitious bud regeneration from internode segments of beech

Internode explants collected from in vitro grown shoots of two clones of Fagus sylvatica L. (European beech) and five clones of F. orientalis Lipski (Oriental beech) were used to evaluate their bud regeneration capacity. Adventitious shoot-buds formed on callus, which developed from internode segments cultured in a Woody Plant Medium supplemented with different concentrations of either thidiazuron (TDZ) or benzyladenine (BA). After 4 weeks of culture on induction media, the explants were transferred to a proliferation medium supplemented with 2.2 μM BA, 9.1 μM zeatin and 2.9 μM indole-3-acetic acid (IAA) for another 8 weeks. Medium containing TDZ was much more efficient than medium containing BA in inducing adventitious buds, the optimal TDZ concentration being 4.5 μM and the optimal BA concentration 17.8 μM. Genotypic variation in shoot regeneration capacity was observed among the two Fagus species and between clones within each species, with a significant interaction between TDZ concentration and genotype regarding mean bud number. Thidiazuron induction medium supplemented with a range of individual auxins was investigated, and it was found that IAA or indole-3-butyric acid at 2.9 μM enhanced the bud forming capacity of explants. Morphogenic response varied significantly with the position of the internode along the stem. The highest regeneration potential was obtained from apical internodes, while those distal to the apex were the least productive. Elongated shoots of adventitious origin can be readily proliferated by axillary branching. This revised version was published online in June 2006 with corrections to the Cover Date.