Rooting strategy of naturally regenerated beech in Silver birch and Scots pine woodlands

This study investigated the belowground development and strategy of late-successional European beech (Fagus sylvatica L.) in ageing natural Scots pine (Pinus sylvestris L.) and Silver birch (Betula pendula Roth.) woodlands in a French volcanic mid-elevation area. For this purpose root biomass, root profile and fine-root architecture of competitor trees were examined in 53 mixed pine–beech and 42 birch–beech woodlands along a stand maturation gradient, using the root auger technique (0–75-cm). The total beech fine-root biomass highly correlated with aerial dimensions such as stem height and girth, whereas it moderately correlated with its age, thus indicating the effects of competition. Basic stand biometric data such as stand density and basal area had no significant effect on beech root biomass. Conversely, competition indices taking into account the vertical dimensions of competitor trees were efficient, probably due to redundancy with beech height. At similar age and height, beeches under birch had a greater belowground development than beeches under pine. Each species exhibited specific rooting pattern and plasticity of fine-root architecture along the gradients of stand maturation and competition. Beech had a heart-shaped rooting habit in both mixings, which strongly increased along stand maturation. Its fine-root system adopted a foraging strategy to respond to increasing stand competition. The Scots pine fine-root system was plate-like and showed a low morphological plasticity, thus presumably a conservative strategy. Silver birch exhibited a high biomass and a foraging capacity in the topsoil but a loose root system in the subsoil. The coexistence of pine and beech roots in the upper soil presumably leads to a high belowground competition. Beech root system becomes predominant throughout the soil profile and it adopts an efficient foraging strategy, but at the expense of its belowground development. Conversely, the niche partitioning strategy between beech and birch may explain why beech develops strongly belowground in spite of the fact that birch has a dense rooting and a competitive fine-root architecture. As a consequence, beech mid-term regeneration and development may be facilitated under birch as compared with pine.     

Stem hypoxia and root respiration of flooded maple and birch seedlings

Some researchers have attributed flood tolerance of woody plants to air entering the shoot through stems, leaves, or lenticels and diffusing to the roots to sustain aerobie respiration. The purpose of this study was to determine if internal aeration of roots by lower stems, changes in gross morphology of lower stems, or both, contribute to flood tolerance of certain tree species. Greenhouse-grown seedlings of red maple ( Acer rubrum L.) and river birch ( Betula nigra L.) tolerated at least 30 days of flooding, where as sugar maple ( Acer saccharum Marsh) and European white birch (also called silver birch, Betula pendula Roth) were intolerant. Flood treatment induced lentieel intumescences and adventitious root formation on red maple stems, but only adventitious roots formed on river birch stems. Stem morphology of sugar maple and European birch was unchanged by flooding. Flood stress decreased oxygen consumption capacity of excised roots from both tolerant and intolerant species. Exclusion of oxygen from the lower stems of flooded red maple and river birch prevented lenticel intumescence and adventitious root formation, but flood tolerance and root respiration capacity were unchanged. Neither internal aeration nor changes in stem morphology appear to account for flood tolerance of red maple and river birch.     

Competition for nitrogen sources between European beech (Fagus sylvatica) and sycamore maple (Acer pseudoplatanus) seedlings

To investigate the short‐term consequences of direct competition between beech and sycamore maple on root N uptake and N composition, mycorrhizal seedlings of both tree species were incubated for 4 days (i.e. beech only, sycamore maple only or both together) in an artificial nutrient solution with low N availability. On the fourth day, N uptake experiments were conducted to study the effects of competition on inorganic and organic N uptake. For this purpose, multiple N sources were applied with a single label. Furthermore, fine roots were sampled and analysed for total amino acids, soluble protein, total nitrogen, nitrate and ammonium content. Our results clearly show that both tree species were able to use inorganic and organic N sources. Uptake of inorganic and organic N by beech roots was negatively affected in the presence of the competing tree species. In contrast, the presence of beech stimulated inorganic N uptake by sycamore maple roots. Both the negative effect of sycamore maple on N uptake of beech and the positive effect of beech on N uptake of sycamore maple led to an increase in root soluble protein in beech, despite an overall decrease in total N concentration. Thus, beech compensated for the negative effects of the tree competitor on N uptake by incorporating less N into structural N components, but otherwise exhibited the same strategy as the competitor, namely, enhancing soluble protein levels in roots when grown under competition. It is speculated that enhanced enzyme activities of so far unknown nature are required in beech as a defence response to inter‐specific competition.     

TEMPORAL AND SPATIAL PATTERNS OF SPECIFIC LEAF WEIGHT IN SUCCESSIONAL NORTHERN HARDWOOD TREE SPECIES

Temporal and spatial patterns of specific leaf weight (SLW, g/m²) were determined for deciduous hardwood tree species in natural habitats in northern lower Michigan to evaluate the utility of SLW as an index of leaf photosynthetic capacity. No significant diurnal changes in SLW were found. Specific leaf weight decreased and then increased during leaf expansion in the spring. Most species, especially those located in the understory, then had relatively constant SLW for most of the growing season, followed by a decline in SLW during autumn. Specific leaf weight decreased exponentially down through the canopy with increasing cumulative leaf area index. Red oak (Quercus rubra), paper birch (Betula papyrifera), bigtooth aspen (Populus grandidentata), red maple (Acer rubrum), sugar maple (A. saccharum), and beech (Fagus grandifolia) generally had successively lower SLW, for leaves at any one level in the canopy. On a given site, comparisons between years and comparisons of leaves growing within 35 cm of each other showed that differences in SLW among species were not due solely to microenvironmental effects on SLW. Bigtooth aspen, red oak, and red maple on lower-fertility sites had lower SLW than the same species on higher-fertility sites. Maximum CO₂ exchange rate, measured at light-saturation in ambient CO₂ and leaf temperatures of 20 to 25 C, increased with SLW. Photosynthetic capacities of species ranked by SLW in a shaded habitat suggest that red oak, red maple, sugar maple, and beech are successively better adapted to shady conditions.     

Elevated CO2 differentially alters the responses of coocurring birch and maple seedlings to a moisture gradient

Summary To determine the effects of elevated CO₂ and soil moisture status on growth and niche characteristics of birch and maple seedlings, gray birch (Betula populifolia) and red maple (Acer rubrum) were experimentally raised along a soil moisture gradient ranging from extreme drought to flooded conditions at both ambient and elevated atmospheric CO₂ levels. The magnitude of growth enhancement due to CO₂ was largely contingent on soil moisture conditions, but differently so for maple than for birch seedlings. Red maple showed greatest CO₂ enhancements under moderately moist soil conditions, whereas gray birch showed greatest enhancements under moderately dry soil conditions. Additionally, CO₂ had a relatively greater ameliorating effect in flooded conditions for red maple than for gray birch, whereas the reverse pattern was true for these species under extreme drought conditions. For both species, elevated CO₂ resulted in a reduction in niche breadths on the moisture gradient; 5% for gray birch and 23% for red maple. Species niche overlap (proportional overall) was also lower at elevated CO₂ (0.98 to: 0.88: 11%). This study highlights the utility of of experiments crossing CO₂ levels with gradients of other resources as effective tools for elucidating the potential consequences of elevated CO₂ on species distributions and potential interactions in natural communities.     

Light and tree size influence belowground development in yellow birch and sugar maple

The effects of light and tree size on the root architecture and mycorrhiza of yellow birch (Betula alleghaniensis Britton) and sugar maple (Acer saccharum Marsh) growing in the understory of deciduous forests in southern Québec, Canada were studied. At the study site, small (<50 m²), medium (101–200 m²) and large (201–500 m²) canopy gaps were investigated. From within these gaps, 17 yellow birch and 23 sugar maple saplings from 40 to 600 cm in height were sampled. In both species, root biomass and morphological traits were strongly correlated with tree size, but only weakly with light availability. Increased root biomass was primarily allocated to coarse roots and secondarily to fine roots. Yellow birch roots were longer, had a larger area, more endings and branches and grew more rapidly than sugar maple roots. Mycorrhizal colonization increased with available light and declined with tree age in sugar maple and was positively associated with tree size in yellow birch. The study demonstrates that tree size is a very important determinant of how belowground systems acclimate to understory conditions.     

Patterns of rhizosphere carbon flux in sugar maple (Acer saccharum) and yellow birch (Betula allegheniensis) saplings

Despite its importance in the terrestrial C cycle rhizosphere carbon flux (RCF) has rarely been measured for intact root–soil systems. We measured RCF for 8‐year‐old saplings of sugar maple (Acer saccharum) and yellow birch (Betula allegheniensis) collected from the Hubbard Brook Experimental Forest (HBEF), NH and transplanted into pots with native soil horizons intact. Five saplings of each species were pulse labeled with ¹³CO₂ at ambient CO₂ concentrations for 4–6 h, and the ¹³C label was chased through rhizosphere and bulk soil pools in organic and mineral horizons for 7 days. We hypothesized yellow birch roots would supply more labile C to the rhizosphere than sugar maple roots based on the presumed greater C requirements of ectomycorrhizal roots. We observed appearance of the label in rhizosphere soil of both species within the first 24 h, and a striking difference between species in the timing of ¹³C release to soil. In sugar maple, peak concentration of the label appeared 1 day after labeling and declined over time whereas in birch the label increased in concentration over the 7‐day chase period. The sum of root and rhizomicrobial respiration in the pots was 19% and 26% of total soil respiration in sugar maple and yellow birch, respectively. Our estimate of the total amount of RCF released by roots was 6.9–7.1% of assimilated C in sugar maple and 11.2–13.0% of assimilated C in yellow birch. These fluxes extrapolate to 55–57 and 90–104 g C m^?2 yr^?1 from sugar maple and yellow birch roots, respectively. These results suggest RCF from both arbuscular mycorrhizal and ectomycorrhizal roots represents a substantial flux of C to soil in northern hardwood forests with important implications for soil microbial activity, nutrient availability and C storage.     

The phosphorus status of northern hardwoods differs by species but is unaffected by nitrogen fertilization

Northern hardwood forests in the eastern US exhibit species-specific influences on nitrogen (N) cycling, suggesting that their phosphorus (P) cycling characteristics may also vary by species. These characteristics are increasingly important to understand in light of evidence suggesting that atmospheric N deposition has increased N availability in the region, potentially leading to phosphorus limitation. We examined how P characteristics differ among tree species and whether these characteristics respond to simulated N deposition (fertilization). We added NH₄NO₃ fertilizer (50 kg ha^?1 year^?1) to single-species plots of red oak (Quercus rubra L.), sugar maple (Acer saccharum Marsh.), eastern hemlock (Tsuga canadensis (L.) Carr.), American beech (Fagus grandifolia Ehrh.), and yellow birch (Betula alleghaniensis Britt.), in the Catskill Mountains, New York from 1997 to 2007. Species differences were observed in foliar, litter and root P concentrations, but all were unaffected by a cumulative N fertilization of 550 kg/ha. Similarly, measures of soil P availability and biotic P sufficiency differed by species but were unaffected by N fertilization. Results suggest species exhibit unique relationships to P as well as N cycles. We found little evidence that N fertilization leads to increased P limitation in these northern hardwood forests. However, species such as sugar maple and red oak may be sufficient in P, whereas beech and hemlock may be less sufficient and therefore potentially more sensitive to future N-stimulated P limitation.     

Mycorrhizal infection,phosphorus uptake,and phenology in Ranunculus adoneus: implications for the functioning of mycorrhizae in alpine systems

Phosphorus levels, phenology of roots and shoots, and development of vesicular arbuscular mycorrhizal (VAM) fungi were monitored for two years in natural populations of the perennial alpine herb, Ranunculus adoneus. The purpose of this study was to understand how phosphorus uptake relates to the phenology of R. adoneus and to ascertain whether arbusculus, fungal structures used for nutrient transfer, were present when maximum phosphorus accumulation was occurring. Arbuscules were only present for a few weeks during the growing season of R. adoneus and their presence corresponded with increased phosphorus accumulation in both the roots and shoots of R. adoneus. In addition, phosphorus accumulation and peaks in mycorrhizal development occurred well after plant reproduction and most plant growth had occurred. The late season accumulation of phosphorus by mycorrhizal roots of R. adoneus is stored for use during early season growth and flowering the following spring. In this way R. adoneus can flower before soils thaw and root or mycorrhizal nutrient uptake can occur.     

Phosphorus effect on phosphatase activity in endomycorrhizal maize

Success of a mycorrhizal symbiosis is influenced by the availability of phosphorus (P) in the soil. Maize ( Zea mays L. cv. Great Lakes 586) plants were grown under five different levels of soil P, either in the presence or absence of formononetin or the vesicular‐arbuscular mycorrhizal (VAM) fungus Glomus intraradices Schenck and Smith. We detected physiological differences in mycorrhizal roots very early in the development of symbiosis, before the onset of nutrient‐dependent responses. Under low P levels, VAM roots accumulated a greater shoot dry weight (13%), root P concentration (15%) and protein concentration (30%) than non-VAM roots, although root growth was not statistically significantly different. At higher P levels, mycorrhizal roots weighed less than non-VAM roots (10%) without a concomitant host alteration of growth or root P concentration. Mycorrhizal colonization decreased as soil P increased. Formononetin-treatment enhanced colonization of the root by G. intraradices and partially overcame inhibition of VAM colonization by high soil P concentrations. This is the first report that formononetin improves root colonization under high levels of soil P. Acid phosphatase (ACP) and alkaline phosphatase (ALP) activities were closely related to the level of fungal colonization in corn roots. ACP activity in corn roots responded more to soil P availability than did ALP activity (38% more). These results suggest that ACP was involved in the increased uptake of P from the soil, while ALP may be linked to active phosphate assimilation or transport in mycorrhizal roots. Thus, soil P directly affected a number of enzymes essential in host-endophyte interplay, while formononetin enhanced fungal colonization.     

Lorentzian model of roots for understory yellow birch and sugar maple saplings

Total 66 small (<50m(2)), 24 medium (101-200m(2)) and 36 large (201-500m(2)) canopy gaps at the three sites of yellow birch (Betula alleghaniensis Britton) and sugar maple (Acer saccharum Marsh) forests were established in southern Québec, Canada. Half of the gaps were covered by 8x8m(2) shading cloths to mimic a closed canopy. From these gaps, 46 understory yellow birch and 46 sugar maple saplings with different tree ages and sizes were sampled. Single- and multi-variable linear and nonlinear models of root biomass and traits (root surface area, volume, length and endings) were developed and examined. Lorentzian model as a multi-variable nonlinear model was firstly applied to the simulations using both base diameter and height, and performed the best fit to total root biomass in both species with the highest correlation coefficients (R(2)=0.96 and 0.98) and smallest root mean squared deviations (RMSD=7.85 and 7.02) among all the examined models. The model also accurately simulated small fine root (2.0mm in diameter), coarse fine root (>2.0-5.0mm) and coarse root (>5.0mm) biomass (R(2)=0.87-0.99; RMSD=2.24-6.41), and the root traits (R(2)=0.71-0.99; RMSD=0.19-19.38). The study showed yellow birch roots were longer, larger, had more endings (tips) and grew faster than sugar maple roots. The root traits were largely distributed to small fine roots, sharply decreased from small fine roots to coarse fine roots, the fewest in coarse roots except for root volume. When trees were large, coarse root biomass increased more rapidly than fine root biomass, but vise versa when the trees were small.     

ONTOGENETIC ANALYSIS OF TREE ROOTS IN ACER RUBRUM AND BETULA PAPYRIFERA

The development of long (6-20 m) tree roots was reconstructed from serial transverse sections taken 10-20 cm apart. Measurements of primary xylem diameter showed that the tips of long roots of red maple (Acer rubrum L.) and paper birch (Betula papyrifera Marsh.) are initially small, but they increase in diameter with increasing distance from the stem. A repeating pattern of the width of the first annual ring marks length increments of long roots. The pattern of ring width is paralleled by a pattern of primary xylem diameter. Our interpretation is that primary xylem diameter increases during spring and early summer, stays constant during late summer and decreases in autumn. After injuries to large root tips, new large lateral tips are formed that replace the parent tip. Thus, a single long root is actually formed by a succession of these replacement roots. In a series of sections there may be sudden changes in primary xylem diameter as the successive sections pass from parent root to replacement root at points of injury. These changes in birch are associated with changes in a number of protoxylem poles.     

Biomass and morphology of fine roots in temperate broad-leaved forests differing in tree species diversity: is there evidence of below-ground overyielding?

Biodiversity effects on ecosystem functioning in forests have only recently attracted increasing attention. The vast majority of studies in forests have focused on above-ground responses to differences in tree species diversity, while systematic analyses of the effects of biodiversity on root systems are virtually non-existent. By investigating the fine root systems in 12 temperate deciduous forest stands in Central Europe, we tested the hypotheses that (1) stand fine root biomass increases with tree diversity, and (2) ‘below-ground overyielding’ of species-rich stands in terms of fine root biomass is the consequence of spatial niche segregation of the roots of different species. The selected stands represent a gradient in tree species diversity on similar bedrock from almost pure beech forests to medium-diverse forests built by beech, ash, and lime, and highly-diverse stands dominated by beech, ash, lime, maple, and hornbeam. We investigated fine root biomass and necromass at 24 profiles per stand and analyzed species differences in fine root morphology by microscopic analysis. Fine root biomass ranged from 440 to 480 g m⁻² in the species-poor to species-rich stands, with 63–77% being concentrated in the upper 20 cm of the soil. In contradiction to our two hypotheses, the differences in tree species diversity affected neither stand fine root biomass nor vertical root distribution patterns. Fine root morphology showed marked distinctions between species, but these root morphological differences did not lead to significant differences in fine root surface area or root tip number on a stand area basis. Moreover, differences in species composition of the stands did not alter fine root morphology of the species. We conclude that ‘below-ground overyielding’ in terms of fine root biomass does not occur in the species-rich stands, which is most likely caused by the absence of significant spatial segregation of the root systems of these late-successional species.     

Positive correlation between VAM-induced changes in root exudation and mycorrhizosphere mycoflora

The influence of vesicular-arbuscular mycorrhizal (VAM) fungi on rhizosphere mycoflora was studied together with the possible mechanism involved in this process. Six combinations of VAM fungi and phosphorus fertilizer treatments were applied to Leucaena leucocephala roots and quantitative and qualitative observations were made periodically of the rhizosphere mycoflora and constituents of root exudates. The results obtained indicate that the presence of specific mycoflora in the rhizosphere of mycrorrhizal roots is mediated through root exudates rather than being an outcome of improved P nutrition.     

Fine root biomass in a beech (Fagus sylvatica L.) stand on Paiko Mountain,NW Greece

Abstract

Fine roots represent a small proportion of total plant biomass however they represent the most dynamic component of the root systems of woody plants. There is limited information on the beech fine root production in Mediterranean ecosystems and especially in Greece. We measured live, dead and total fine root biomass (d<2 mm) (LFRB, DFRB and TFRB, respectively) over a growing season in a beech (Fagus sylvatica L.) stand on Paiko mountain, NW Greece, in order to contribute to the generally scarce knowledge of the fine root biomass of beech stands. It was found that TFRB and LFRB increased from May to July and then decreased. LFRB decreased with soil depth while there was no pattern at the change of DFRB with soil depth.     

SEASONAL AND INDIVIDUAL VARIATION IN LEAF QUALITY OF TWO NORTHERN HARDWOODS TREE SPECIES

Seasonal trends in five traits of sugar maple (Acer saccharum Marsh.) and yellow birch (Betula allegheniensis Britt.) leaves thought to influence feeding by herbivores were measured from 17 May through 19 September, 1979. Total nitrogen and water contents declined and toughness increased through the growth season. These seasonal changes were more pronounced in sugar maple than in yellow birch. Total polyphenol contents and tanning coefficients of leaf extracts from both species reached a season high by the end of May and changed very little after that date; these patterns differ from those reported by several other investigators. Sugar maple leaf extracts exhibited much higher tanning coefficients than did those of yellow birch, a finding which is consistent with current plant defense theory. Significant differences in total polyphenol content and tanning coefficients were found between individual trees in yellow birch, but not sugar maple. The relationship between successional status, leaf quality traits, and variability in these traits in forest trees is discussed.     

Effects of environmental variables on vesicular‐arbuscular mycorrhizal abundance in wild populations of Vangueria infausta

Abstract. Vesicular‐arbuscular endomycorrhizal fungi include among its members some of the most widespread root symbiont species. It is not known whether these fungal species show environmental or host preferences. In 13 semi‐arid savanna sites in Botswana, we found positive correlations between individual environmental factors and the abundance of VAM associations in the roots of an important host, the indigenous fruit tree Vangueria infausta (Rubiaceae). The concentration of phosphorus in the leaves of the host was positively correlated with both phosphorus in the soil and the abundance of VAM associations in its roots, indicating direct benefits to the host of this association. Abundance of VAM associations was significantly different between the studied sites and between seasons and was positively correlated with the mean annual rainfall. In addition, there was a negative correlation with phosphorus concentration in the soil. The differences between the summer and winter abundance of VAM associations were positively correlated with the density of bush cover and the amount of grazing. The apparent correlation found between the abundance of VAM in V. infausta and bush and tree community composition is presumably related to correlations between these two parameters and environmental conditions.     

The root sap of birch on a podzol

Summary Root sap analyses were made on birch trees growing in a strongly podzolised heath soil. In the early summer the concentration of calcium and phosphorus appears to vary with the quantity of sap produced, and is independent of root environment. Later in the summer, however, this relationship no longer holds; there now being a connection between rooting zone and nutrient concentration. Roots reaching theC horizon absorb greater quantities of bases, whilst the superficial roots show higher concentrations of phosphorus and ammoniacal nitrogen. Throughout the season potassium uptake remains practically independent of both sap yield and rooting zone. In these soils young pine trees do not penetrate to the subsoil, but a comparison of sap analyses from surface roots of pine and birch shows that birch sap is richer in most nutrients. This, together with the ability of the birch to exploit the subsoil, may be a contributory factor to the soil improving properties of a birch crop.     

Effects of aqueous extracts from leaves and leaf litter on the abundance and diversity of soil gymnamoebae in laboratory microcosm cultures

The distribution and abundance of microbiota in soil and litter may be significantly affected by the quality and quantity of localized patches of leaf organic matter. This study examined the relative effects of aqueous extracts of shed autumn leaves from American beech (Fagus grandifolia), sugar maple (Acer saccharum), red oak (Quercus rubra), and white oak (Quercus alba) on the density and diversity of gymnamoebae in laboratory cultures. Overall, the beech leaf extract produced the most growth of gymnamoebae followed by white oak with leaf extracts from maple and red oak producing least growth. Cultures using natural leaf litter from beneath beech trees had higher densities and diversity of gymnamoebae than leaf-litter cultures from a maple-oak stand. Soil microcosms confirmed that beech leaf extracts produced a higher density of gymnamoeba growth when added to soil cultures compared with maple and oak leaf extracts. Protein content, CHN (carbon and nitrogen content), and pH of the leaf extracts were assayed, but these alone were not sufficiently different to account for the effects. A dilution experiment indicated that some other concentration-dependent factor in the extract may produce the effects.     

Differential drought responses between saplings and adult trees in four co-occurring species of New England

Tree-ring characteristics in four species were examined to address whether co-occurring mature trees of different successional status respond differently to drought, and whether saplings of these species have a greater response to drought than mature trees. We examined saplings and mature trees of paper birch, yellow birch, red maple and sugar maple, which varied in successional status (shade-tolerance) and co-occurred at Harvard Forest, Petersham, Mass., USA. Three drought events in 1964–1966, 1981 and 1995 were identified using climate data. For mature trees, there was no significant interspecific difference in relative changes in ring-width index (RWI) during the 1964–1966 and 1995 drought events. However, the interspecific difference was significant in the 1981 drought event. Response function analysis for mature trees showed that the radial growth of sugar maple was mainly controlled by spring and summer precipitation, red maple by spring and summer precipitation and temperature, yellow birch by winter and summer precipitation, and spring and summer temperature, and paper birch by spring and summer precipitation and spring temperature. Saplings of sugar maple and yellow birch, but not red maple and paper birch, showed significant positive correlations between RWI and annual total precipitation. In the 1995 drought event, saplings and mature trees of red maple and paper birch differed significantly in drought responses, but this was not true in sugar maple and yellow birch. Our results do not support a generally greater response in saplings than in mature trees, nor an early- versus late successional difference in drought responses.