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Fine roots (≤2 mm) consume a large proportion of photosynthates and thus play a key role in the global carbon cycle, but our knowledge about fine root biomass, production, and turnover across environmental gradients is insufficient, especially in tropical ecosystems. Root system studies along elevation transects can produce valuable insights into root trait-environment relationships and may help to explore the evidence for a root economics spectrum (RES) that should represent a trait syndrome with a trade-off between resource acquisitive and conservative root traits. We studied fine root biomass, necromass, production, and mean fine root lifespan (the inverse of fine root turnover) of woody plants in six natural tropical ecosystems (savanna, four tropical mountain forest types, tropical alpine heathland) on the southern slope of Mt. Kilimanjaro (Tanzania) between 900 and 4,500 m a.s.l. Fine root biomass and necromass showed a unimodal pattern along the slope with a peak in the moist upper montane forest (~2,800 m), while fine root production varied little between savanna and upper montane forest to decrease toward the alpine zone. Root:shoot ratio (fine root biomass and production related to aboveground biomass) in the tropical montane forest increased exponentially with elevation, while it decreased with precipitation and soil nitrogen availability (decreasing soil C:N ratio). Mean fine root lifespan was lowest in the ecosystems with pronounced resource limitation (savanna at low elevation, alpine heathland at high elevation) and higher in the moist and cool forest belt (~1,800-3,700 m). The variation in root traits across the elevation gradient fits better with the concept of a multi-dimensional RES, as root tissue density and specific root length showed variable relations to each other, which does not agree with a simple trade-off between acquisitive and conservative root traits. In conclusion, despite large variation in fine root biomass, production, and morphology among the different plant species and ecosystems, a general belowground shift in carbohydrate partitioning is evident from 900 to 4,500 m a.s.l., suggesting that plant growth is increasingly limited by nutrient (probably N) shortage toward higher elevations., (Copyright © 2020 Sierra Cornejo, Hertel, Becker, Hemp and Leuschner.)

In high-elevation grasslands, plants can encounter periods with high air temperature while the soil remains cold, which may lead to a temporary mismatch in the physiological activity of leaves and roots. In a climate chamber experiment with graminoid species from three elevations (4400, 2400, and 250 m a.s.l.), we tested the hypothesis that soil temperature can influence photosynthesis and stomatal conductance independently of air temperature. Soil monoliths with swards of Kobresia pygmaea (high alpine), Nardus stricta (lower alpine), and Deschampsia flexuosa (upper lowland) were exposed to soil temperatures of 25, 15, 5, and -2°C and air temperatures of 20 and 10°C for examining the effect of independent soil and air temperature variation on photosynthesis, leaf dark respiration, and stomatal conductance and transpiration. Soil frost (-2°C) had a strong negative effect on gas exchange and stomatal conductance in all three species, independent of the elevation of origin. Leaf dark respiration was stimulated by soil frost in D. flexuosa , but not in K. pygmaea , which also had a lower temperature optimum of photosynthesis. Soil cooling from 15 to 5°C did not significantly reduce stomatal conductance and gas exchange in any of the species. We conclude that all three graminoids are able to maintain a relatively high root water uptake in cold, non-frozen soil, but the high-alpine K. pygmaea seems to be especially well adapted to warm shoot - cold root episodes, as it has a higher photosynthetic activity at 10 than 20°C air temperature and does not up-regulate leaf dark respiration upon soil freezing, as was observed in the grasses from warmer climates.

Saltmarsh plants are exposed to multiple stresses including tidal inundation, salinity, wave action and sediment anoxia, which require specific root system adaptations to secure sufficient resource capture and firm anchorage in a temporary toxic environment. It is well known that many saltmarsh species develop large below-ground biomass (roots and rhizomes) but relations between fine roots, in particular, and the abiotic conditions in salt marshes are widely unknown. We studied fine root mass (<2 mm in diameter), fine root depth distribution and fine root morphology in three typical communities ( Spartina anglica -dominated pioneer zone, Atriplex portulacoides -dominated lower marsh, Elytrigia atherica -dominated upper marsh) across elevational gradients in two tidal salt marshes of the German North Sea coast [a mostly sandy marsh on a barrier island (Spiekeroog), and a silty-clayey marsh on the mainland coast (Westerhever)]. Fine root mass in the 0-40 cm profile ranged between 750 and 2,500 g m -2 in all plots with maxima at both sites in the lower marsh with intermediate inundation frequency and highest plant species richness indicating an effect of biodiversity on fine root mass. Fine root mass and, even more, total fine root surface area (maximum 340 m 2 m -2 ) were high compared to terrestrial grasslands, and were greater in the nutrient-poorer Spiekeroog marsh. Fine root density showed only a slight or no decrease toward 40 cm depth. We conclude that the standing fine root mass and morphology of these salt marshes is mainly under control of species identity and nutrient availability, but species richness is especially influential. The plants of the pioneer zone and lower marsh possess well adapted fine roots and large standing root masses despite the often water-saturated sediment.

Despite their importance for water uptake and transport, the xylem anatomical and hydraulic properties of tree roots have only rarely been studied in the field. We measured mean vessel diameter ( D ), vessel density (VD), relative vessel lumen area (lumen area per xylem area) and derived potential hydraulic conductivity ( K p ) in the xylem of 197 fine- to medium-diameter roots (1-10 mm) in the topsoil and subsoil (0-200 cm) of a mature European beech forest on sandy soil for examining the influence of root diameter and soil depth on xylem anatomical and derived hydraulic traits. All anatomical and functional traits showed strong dependence on root diameter and thus root age but no significant relation to soil depth. Averaged over topsoil and deep soil and variable flow path lengths in the roots, D increased linearly with root diameter from ∼50 μm in the smallest diameter class (1-2 mm) to ∼70 μm in 6-7 mm roots (corresponding to a mean root age of ∼12 years), but remained invariant in roots >7 mm. D never exceeded ∼82 μm in the 1-10 mm roots, probably in order to control the risk of frost- or drought-induced cavitation. This pattern was overlain by a high variability in xylem anatomy among similar-sized roots with K p showing a higher variance component within than between root diameter classes. With 8% of the roots exceeding average K p in their diameter class by 50-700%, we obtained evidence of the existence of 'high-conductivity roots' indicating functional differentiation among similar-sized roots. We conclude that the hydraulic properties of small to medium diameter roots of beech are mainly determined by root age, rendering root diameter a suitable predictor of hydraulic functioning, while soil depth - without referring to path length - had a negligible effect.

Low temperatures are crucial for the formation of the alpine treeline worldwide. Since soil temperature in the shade of tree canopies is lower than in open sites, it was assumed that self-shading may impair the trees' root growth performance. While experiments with tree saplings demonstrate root growth impairment at soil temperatures below 5-7°C, field studies exploring the soil temperature - root growth relationship at the treeline are missing. We recorded soil temperature and fine root abundance and dynamics in shaded and sun-exposed areas under canopies of isolated Pinus cembra trees at the alpine treeline. In contrast to the mentioned assumption, we found more fine root biomass and higher fine root growth in colder than in warmer soil areas. Moreover, colder areas showed higher fine root turnover and thus lower root lifespan than warmer places. We conclude that P. cembra balances enhanced fine root mortality in cold soils with higher fine root activity and by maintaining higher fine root biomass, most likely as a response to shortage in soil resource supply. The results from our study highlight the importance of in situ measurements on mature trees to understand the fine root response and carbon allocation pattern to the thermal growth conditions at the alpine treeline.

Advancing our understanding of tree fine root dynamics is of high importance for tree physiology and forest biogeochemistry. In temperate broad-leaved forests, ectomycorrhizal (EM) and arbuscular mycorrhizal (AM) tree species often are coexisting. It is not known whether EM and AM trees differ systematically in fine root dynamics and belowground resource foraging strategies. We measured fine root productivity (FRP) and fine root turnover (and its inverse, root longevity) of three EM and three AM broad-leaved tree species in a natural cool-temperate mixed forest using ingrowth cores and combined the productivity data with data on root biomass per root orders. FRP and root turnover were related to root morphological traits and aboveground productivity. FRP differed up to twofold among the six coexisting species with larger species differences in lower horizons than in the topsoil. Root turnover varied up to fivefold among the species with lowest values in Acer pseudoplatanus and highest in its congener Acer platanoides. Variation in root turnover was larger within the two groups than between EM and AM species. We conclude that the main determinant of FRP and turnover in this mixed forest is species identity, while the influence of mycorrhiza type seems to be less important.

Species diversity may increase the productivity of tree communities through complementarity (CE) and/or selection effects (SE), but it is not well known how this relationship changes under water limitation. We tested the stress-gradient hypothesis, which predicts that resource use complementarity and facilitation are more important under water-limited conditions. We conducted a growth experiment with saplings of five temperate broad-leaved tree species that were grown in assemblages of variable diversity (1, 3, or 5 species) and species composition under ample and limited water supply to examine effects of species richness and species identity on stand- and tree-level productivity. Special attention was paid to effects of neighbor identity on the growth of target trees in mixture as compared to growth in monoculture. Stand productivity was strongly influenced by species identity while a net biodiversity effect (NE) was significant in the moist treatment (mostly assignable to CE) but of minor importance. The growth performance of some of the species in the mixtures was affected by tree neighborhood characteristics with neighbor size likely being more important than neighbor species identity. Diversity and neighbor identity effects visible in the moist treatment mostly disappeared in the dry treatment, disproving the stress-gradient hypothesis. The mixtures were similarly sensitive to drought-induced growth reduction as the monocultures, which may relate to the decreased CE on growth upon drought in the mixtures.

While most temperate broad-leaved tree species form ectomycorrhizal (EM) symbioses, a few species have arbuscular mycorrhizas (AM). It is not known whether EM and AM tree species differ systematically with respect to fine root morphology, fine root system size and root functioning. In a species-rich temperate mixed forest, we studied the fine root morphology and biomass of three EM and three AM tree species from the genera Acer, Carpinus, Fagus, Fraxinus, and Tilia searching for principal differences between EM and AM trees. We further assessed the evidence of convergence or divergence in root traits among the six co-occurring species. Eight fine root morphological and chemical traits were investigated in root segments of the first to fourth root order in three different soil depths and the relative importance of the factors root order, tree species and soil depth for root morphology was determined. Root order was more influential than tree species while soil depth had only a small effect on root morphology All six species showed similar decreases in specific root length and specific root area from the 1st to the 4th root order, while the species patterns differed considerably in root tissue density, root N concentration, and particularly with respect to root tip abundance. Most root morphological traits were not significantly different between EM and AM species (except for specific root area that was larger in AM species), indicating that mycorrhiza type is not a key factor influencing fine root morphology in these species. The order-based root analysis detected species differences more clearly than the simple analysis of bulked fine root mass. Despite convergence in important root traits among AM and EM species, even congeneric species may differ in certain fine root morphological traits. This suggests that, in general, species identity has a larger influence on fine root morphology than mycorrhiza type.

Leaf and fine root morphology and physiology have been found to vary considerably among tree species, but not much is known about intraspecific variation in root traits and their relatedness to leaf traits. Various aspen progenies (Populus tremula and P. tremuloides) with different growth performance are used in short-rotation forestry. Hence, a better understanding of the link between root trait syndromes and the adaptation of a deme to a particular environment is essential in order to improve the match between planted varieties and their growth conditions. We examined the between-deme (genetic) and within-deme (mostly environmental) variation in important fine root traits [mean root diameter, specific root area (SRA) and specific root length (SRL), root tissue density (RTD), root tip abundance, root N concentration] and their co-variation with leaf traits [specific leaf area (SLA), leaf size, leaf N concentration] in eight genetically distinct P. tremula and P. tremuloides demes. Five of the six root traits varied significantly between the demes with largest genotypic variation in root tip abundance and lowest in mean root diameter and RTD (no significant difference). Within-deme variation in root morphology was as large as between-deme variation suggesting a relatively low genetic control. Significant relationships existed neither between SLA and SRA nor between leaf N and root N concentration in a plant. Contrary to expectation, high aboveground relative growth rates (RGR) were associated with large, and not small, fine root diameters with low SRA and SRL. Compared to leaf traits, the influence of root traits on RGR was generally low. We conclude that aspen exhibits large intraspecific variation in leaf and also in root morphological traits which is only partly explained by genetic distances. A root order-related analysis might give deeper insights into intraspecific root trait variation.