Fine Root Area Research Articles

Adapting to climate change: responses of fine root traits and C exudation in five tree species with different light-use strategy.

Trees that are categorised by their light requirements have similarities in their growth strategies and adaptation mechanisms. We aimed to understand the complex responses of elevated air humidity on whole tree fine root carbon (C) exudation (ExC) and respiration rate, morphology, and functional distribution in species with different light requirements. Three light-demanding (LD) species, Populus × wettsteinii, Betula pendula, and Pinus sylvestris, and two shade-tolerant species, Picea abies and Tilia cordata saplings were grown in growth chambers under moderate and elevated air relative humidity (eRH) at two different inorganic nitrogen sources with constant air temperature and light availability. The proportion of assimilated carbon released by ExC, and respiration decreased at eRH; up to about 3 and 27%, respectively. There was an indication of a trade-off between fine root released C and biomass allocation. The elevated air humidity changed the tree biomass allocation and fine root morphology, and the responses were species-specific. The specific fine root area and absorptive root proportion were positively related to canopy net photosynthesis and leaf nitrogen concentration across tree species. The variation in ExC was explained by the trees' light-use strategy (p < 0.05), showing higher exudation rates in LD species. The LD species had a higher proportion of pioneer root tips, which related to the enhanced ExC. Our findings highlight the significant role of fine root functional distribution and morphological adaptation in determining rhizosphere C fluxes in changing environmental conditions such as the predicted increase of air humidity in higher latitudes.

Site conditions more than species identity drive fine root biomass, morphology and spatial distribution in temperate pure and mixed forests

AimThe establishment of broadleaved - conifer mixtures has had positive effects on productivity. However, it is still not clear the degree to which belowground space occupation contributes to these effects and how root traits of the respective species differ between mixed and pure stands. Thus, the aim of this study was to examine how enrichment of beech forests with conifer species impacts fine root biomass and other root traits across variable site conditions. MethodsWe studied 20 mature pure and mixed stands of European beech (Fagus sylvatica), Norway spruce (Picea abies), and Douglas-fir (Pseudotsuga menziesii), and determined differences in biomass, spatial distribution, and morphology of fine roots (≤2 mm, to 60 cm soil depth) among beech – conifer mixtures and pure stands of the respective species in two regions with differing site conditions in Northern Germany. ResultsSite conditions had strong effects on fine root biomass (FRB) and fine root area index (RAI). Both FRB and RAI were significantly higher across all stand types on the sites with lower belowground resource availability. On the less favorable sites, FRB significantly increased with increasing beech proportion. In contrast to specific root area (SRA) and RAI, specific root length (SRL) was not affected by site conditions but differed significantly among species. No overyielding was observed when FRB was plotted against basal area and/or area potentially available (APA), which reflects the aboveground horizontal space configuration. Spatial fine root biomass patterns of all species in both monospecific and mixed stands changed along the vertical soil profile at both sites. FRB decreased with increasing soil depth, with the exception of beech in mixed stands. ConclusionOur data suggest that both amount and distribution of tree fine roots in pure and mixed stands are highly variable and depend primarily on-site conditions and secondarily on tree species identity. By disentangling soil and species identity effects, we conclude that beech has a remarkable potential for adjusting fine root biomass to cope with unfavorable conditions as compared to spruce and Douglas-fir.

Adaptive versus non-adaptive responses to drought in a non-native riparian tree / shrub, Tamarix spp

ABSTRACT Rapidly colonizing species often thrive in a wide-range of conditions due to a high degree of phenotypic plasticity that results in populations of “general purpose” genotypes. Alternatively, species with high genetic variation could rapidly respond to forces of selection such that a local population evolves traits that provide an advantage in its local environment. We tested this generalist versus local adaptation paradigm in a widely distributed, recently introduced riparian tree / shrub, Tamarix spp. Using an eighty-day, mid-summer common garden drought experiment, we tested three inter-related hypotheses: 1) stomatal sensitivity to soil water depletion is lower in genotypes from a low-elevation, ephemerally flowing river than genotypes from a low-elevation, perennially flowing river, indicative of local adaptation to hydrological conditions, 2) stomatal sensitivity to soil water depletion is lower in genotypes from high-elevations with regular freeze-thaw exposure than genotypes sourced from locations with no freeze-thaw exposure, indicative of local adaptation to temperature conditions, and 3) differences among genotypes in drought sensitivity are correlated with differences in fine root area to leaf area ratios (Ar:Al). For the most part, results did not support hypothesis 1, but largely supported hypotheses 2 and 3. Specifically, sap-flux scaled canopy transpiration (El) declined 14 days earlier in the low-elevation populations after drought initiation compared to El of the highest elevation population with the highest freeze-thaw exposure. Root area to leaf area ratios of the two low-elevation populations were also only 42% and 55% of Ar:Al in the highest elevation population. Results indicate that the high degree of reported Tamarix hybridization since its introduction to North America has largely produced a swarm of “generalist” genotypes in terms of drought sensitivity. Nevertheless, rapid changes in ecohydrologic conditions may result in some Tamarix populations becoming maladapted sooner to reductions in available water than others in the western US.

Effects of warming and fertilization interacting with intraspecific competition on fine root traits of Picea asperata

Abstract Aims Competition, temperature and nutrient are the most important determinants of tree growth in the cold climate on the eastern Tibetan Plateau. Although many studies have reported their individual effects on tree growth, little is known about how the interactions of competition with fertilization and temperature affect root growth. We aim to test whether climate warming and fertilization promote competition and to explore the functional strategies of Picea asperata in response to the interactions of these factors. Methods We conducted a paired experiment including competition and non-competition treatments under elevated temperature (ET) and fertilization. We measured root traits, including the root tip number over the root surface (RTRS), the root branching events over the root surface (RBRS), the specific root length (SRL), the specific root area (SRA), the total fine root length and area (RL and RA), the root tips (RTs) and root branching (RB) events. These root traits are considered to be indicators of plant resource uptake capacity and root growth. The root biomass and the nutrient concentrations in the roots were also determined. Important Findings The results indicated that ET, fertilization and competition individually enhanced the nitrogen (N) and potassium (K) concentrations in fine roots, but they did not affect fine root biomass or root traits, including RL, RT, RA and RB. However, both temperature and fertilization, as well as their interaction, interacting with competition increased RL, RA, RT, RB and nutrient uptake. In addition, the SRL, SRA, RTRS and RBRS decreased under fertilization, the interaction between temperature and competition decreased SRL and SRA, while the other parameters were not affected by temperature or competition. These results indicate that P. asperata maintains a conservative nutrient strategy in response to competition, climate warming, fertilization and their interactions. Our results improve our understanding of the physiological and ecological adaptability of trees to global change.

Shifts in ectomycorrhizal exploration types parallel leaf and fine root area with forest age

Abstract The abundance of fine roots and leaves in forests typically peaks during mid‐succession and then declines. If fine root area declines more rapidly than leaf area, this could contribute disproportionately to stand decline. However, trees also partner with symbiotic ectomycorrhizal (EcM) fungi that facilitate nutrient acquisition of fine roots. Ectomycorrhizal fungi can use the carbohydrates provided by their plant partners to produce emanating fungal tissues which increase soil exploration potential but may also prove costly. We suggest two competing hypotheses to frame the response of ectomycorrhizas to stand age: (a) the ‘functional offset’ hypothesis posits that the abundance of ectomycorrhizas with emanating tissues (‘Distance’ mycorrhizas) increases with stand age and (b) the ‘energy‐limited’ hypothesis posits that carbon available for root symbionts decreases with stand age resulting in fewer Distance mycorrhizas. In the first hypothesis, EcM functional traits offset root abundance, while in the second, traits parallel root abundance. To test these competing hypotheses, we sampled fine roots to a depth of 90 cm and used allometric equations to estimate changes in root and leaf area index across a chronosequence of Pinus banksiana stands ranging from 2 to 76 years average tree age. We also examined fine roots microscopically to track changes in the abundance of EcM functional types. We used DNA‐based methods to sequence EcM fungi and confirm roots were of P. banksiana. Both fine root and leaf area of pine increased for the first 30–36 years (until Mid age) and then plateaued, while the ratio of leaf to fine root area was similar across the age gradient. Also, changes to fine root area with stand age depended on soil depth. The abundance of Distance mycorrhizas was lowest at the youngest plots and increased until Mid age, where it peaked, contrary to the functional offset hypothesis. Instead, the abundance of Distance mycorrhizas paralleled changes to leaf area, aligned with the energy‐limited hypothesis. Synthesis. Ectomycorrhizas do not offset the function of roots, rather shifts in exploration type likely reflect adjustments to carbon supply from hosts.

Fine-Root Traits Reveal Contrasting Ecological Strategies in European Beech and Norway Spruce During Extreme Drought.

Trees adjust multiple structural and functional organ-specific characteristics, “traits”, to cope with diverse soil conditions. Studies on traits are widely used to uncover ecological species adaptability to varying environments. However, fine-root traits are rarely studied for methodological reasons. We analyzed the adaptability of the fine-root systems of European beech and Norway spruce to extreme drought within species-specific tree groups at Kranzberger Forst (Germany), focusing on the seasonality of morphological, physiological, and biochemical key traits in view of carbon (C) and nitrogen dynamics. We hypothesized that fine roots of both species adjust to seasonal drought: with beech representing a “fast” (i.e. with fast C turnover), and spruce a “slow” (i.e. with long-term C retention) ecological strategy. We identified three functional fine-root categories, based on root function (absorptive or transport fine roots), and mycorrhizal status of the absorptive fine-roots (mycorrhizal or non-mycorrhizal). Solely the non-mycorrhizal absorptive roots adjusted in a species-specific manner supporting fine-root ecological strategy hypothesis. During drought, beech produced thin ephemeral (absorptive non-mycorrhizal) fine roots with high specific fine-root area and high respiratory activity, representing fast C turnover and enabling effective resource exploitation. These adjustments reflect a “fast” ecological strategy. Conversely, spruce absorptive fine roots did not respond to the soil moisture deficit by growth but instead increased root suberization. Drastically lowered respiratory activity of this functional category facilitated C retention and structural persistence during drought, indicating a “slow” ecological strategy in spruce. Absorptive mycorrhizal fine roots maintained respiration throughout the drought event in both tree species, but in spruce this was the only fine-root category with high respiration. This suggests, that spruce relies heavily on mycorrhizal associations as a method of drought resistance. Accumulation of non-structural carbohydrates and high C concentrations were observed in the transport fine roots of both species, indicating drought-induced osmotic protection of these roots. Thus, functional classification enabled us to determine that fine-root branches of each species are not tied to one sole ecological strategy. The suggested approach helps to better understand the complex interplay between structure and function belowground.

Vertical distribution of fine-root area in relation to stand age and environmental factors in black locust (Robinia pseudoacacia) forests of the Chinese Loess Plateau

To examine the spatiotemporal characteristics of the distribution of fine-root area and its relationship with stand age and environmental factors in black locust (Robinia pseudoacacia L.) on the Chinese Loess Plateau, black locust stands were selected at four sites along a precipitation gradient. Four stands of different ages and a transect along the hillslope were also selected at one of the sites. With increasing stand age, fine-root area at the tree level increased exponentially, and the rooting pattern tended to be deeper for trees up to 15 years old and then shallower thereafter. The temporal changes of fine-root distribution could be quantified using stand age and soil nutrients. At the hillslope scale, fine-root area index (FRAI) was lower while the rooting pattern was deeper in the middle slope than in the upper and lower slopes, and the fine-root distribution could be quantified using elevation and soil properties. At the regional scale, FRAI decreased substantially while exhibiting similar rooting patterns with decreasing soil water and nutrient availability along the precipitation gradient. Humidity index represented the regional environmental variation and could be used to quantify FRAI. These findings will be helpful for improving quantification of fine roots and enhancing the accuracy of ecohydrological models.

Soil type more than precipitation determines fine-root abundance in savannas of Kruger National Park, South Africa

Our aim was to examine how soil type and precipitation affect fine-root abundance in savanna ecosystems across Kruger National Park (KNP), South Africa. Fine-root distributions were measured in four sites that represent the natural factorial combination of soil types (basalt-derived clay or granite-derived sand) and precipitation regimes [wet (~750 mm mean annual precipitation) or dry (~500 mm mean annual precipitation)] that occur in KNP. Root area and biomass (at soil depths of 0–75 cm) were estimated from measurements of root number, length and width in images from minirhizotron tubes at each site. Measurements were made during one mid-season sampling during three subsequent years. Fine-root area was more than twice as large in clay (2.3 ± 0.0 mm2 cm−2) than sand (0.8 ± 0.3 mm2 cm−2) sites but did not differ between wet and dry sites. Root number, length and width, used to derive area, showed similar patterns to fine-root area. Fine-root biomass estimated from these values was 5.5 ± 0.6 Mg ha−1 in clay sites and 2.2 ± 0.9 Mg ha−1 in sand sites. Across the four sites, a change from sand to clay soils had a greater effect on fine-root abundance and distributions than a 50% increase in precipitation from dry to wet sites. Results highlight the importance of soil properties on root dynamics and carbon pools in the region.

A savanna response to precipitation intensity.

As the atmosphere warms, precipitation events are becoming less frequent but more intense. A three-year experiment in Kruger National Park, South Africa, found that fewer, more intense precipitation events encouraged woody plant encroachment. To test whether or not these treatment responses persisted over time, here, we report results from all five years of that experiment. Grass growth, woody plant growth, total fine root number and area and hydrologic tracer uptake by grasses and woody plants were measured in six treated plots (8 m by 8 m) and six control plots. Treatment effects on soil moisture were measured continuously in one treated and one control plot. During the fourth year, increased precipitation intensity treatments continued to decrease water flux in surface soils (0–10 cm), increase water flux in deeper soils (20+ cm), decrease grass growth and increase woody plant growth. Greater root numbers at 20–40 cm and greater woody plant uptake of a hydrological tracer from 45–60 cm suggested that woody plants increased growth by increasing root number and activity (but not root area) in deeper soils. During the fifth year, natural precipitation events were large and intense so treatments had little effect on precipitation intensity or plant available water. Consistent with this effective treatment removal, there was no difference in grass or woody growth rates between control and treated plots, although woody plant biomass remained higher in treated than control plots due to treatment effects in the previous four years. Across the five years of this experiment, we found that 1) small increases in precipitation intensity can result in large increases in deep (20–130 cm) soil water availability, 2) plant growth responses to precipitation intensity are rapid and disappear quickly, and 3) because woody plants accumulate biomass, occasional increases in precipitation intensity can result in long-term increases in woody plant biomass (i.e., shrub encroachment). While results are likely to be site-specific, they provide experimental evidence of large ecohydrological responses to small changes in precipitation intensity.

Viewing forests from below: fine root mass declines relative to leaf area in aging lodgepole pine stands.

In the continued quest to explain the decline in productivity and vigor with aging forest stands, the most poorly studied area relates to root system change in time. This paper measures the wood production, root and leaf area (and mass) in a chronosequence of fire-origin lodgepole pine (Pinus contorta Loudon) stands consisting of four age classes (12, 21, 53, and ≥100 years), each replicated ~ five times. Wood productivity was greatest in the 53-year-old stands and then declined in the ≥100-year-old stands. Growth efficiency, the quantity of wood produced per unit leaf mass, steadily declined with age. Leaf mass and fine root mass plateaued between the 53- and ≥100-year-old stands, but leaf area index actually increased in the older stands. An increase in the leaf area index:fine root area ratio supports the idea that older stand are potentially limited by soil resources. Other factors contributing to slower growth in older stands might be lower soil temperatures and increased self-shading due to the clumped nature of crowns. Collectively, the proportionally greater reduction in fine roots in older stands might be the variable that predisposes these forests to be at a potentially greater risk of stress-induced mortality.Electronic supplementary materialThe online version of this article (doi:10.1007/s00442-016-3621-6) contains supplementary material, which is available to authorized users.

A fast exploration of very deep soil layers by Eucalyptus seedlings and clones in Brazil

Although pioneer studies showed several decades ago that deep rooting is common in tropical forests, direct measurements of fine root distributions over the entire soil profile explored by the roots are still scarce. Our study aimed to compare, 2years after planting, fine root traits of Eucalyptus trees planted from cuttings and from seedlings in order to assess whether the propagation mode has an influence on the capacity of the trees to explore very deep soils. Soils cores were sampled down to a depth of 13.5m at the peak of leaf area index (LAI), 2years after planting, under three Eucalyptus clones (belonging to species E. saligna, E. grandis×E. urophylla, E. grandis×E. camaldulensis) and under E. grandis seedlings in the same Ferralsol soil. LAI was estimated using allometric equations based on destructive sampling of eight trees per genotype.All the genotypes exhibited fine root densities roughly constant between the depths of 0.25 and 6.00m. Changes in fine root traits (diameter, specific root length and specific root area) were low between the topsoil and the root front. The ratios between mean tree height and root front depth ranged from 0.8 to 1.2 for the four genotypes. Although tree vertical extension was roughly symmetric above and belowground for all the genotypes, the depth of the root front ranged from 8.0m for the seedlings and the E. grandis×E. urophylla clone to 11.5m for the E. saligna clone. Soil water content profiles suggested that the four genotypes had the capacity to withdraw water down to a depth of 8–10m over the first 2years after planting. Total fine root length ranged from 3.3 to 6.0km per m2 of soil depending on the genotype. The root area/leaf area ratio ranged from 1.3 to 3.2 and was negatively correlated with LAI across the four genotypes. This pattern suggests that the genotypes more conservative for water use (with a low LAI) invest more in fine root area relative to leaf area than genotypes adapted to wet regions (with a high LAI). The velocity of downward movement of the root front might be a relevant criterion in the last stage of the breeding programs to select clones with a fast exploration of deep soil layers in drought prone regions.

Estimation of carbon stocks of Balganga Reserved Forest, Uttarakhand, India

Forest carbon (C) stock estimation was carried out for tree species of Balganga Reserved Forest (BRF) in district Tehri Garhwal, Uttarakhand, India. The highest total biomass density (TBD) and total carbon density (TCD) were estimated at site III in the altitudinal range of 1800–2600 m as 108.26 and 53.45 Mg⋅ha−1 followed by site II in the range of 1600–1800 m as 83.92 and 41.96 Mg⋅ha−1, and the least at site I in the altitudinal range of 1000–1400 m as 57.22 and 28.61 Mg⋅ha−1 with an average of 83.13 and 41.56 Mg⋅ha−1, respectively. The study found average TBD and TCD of the BRF are 23% more when compared to forests located in the same ecoregion due to difference in diameter at breast height and height of tree species. It also found that the C stock increases with increasing altitude due to fine root area, fine root length growth, and decreasing temperature. The BRF was found to have 16.65 million tons of forest organic C stocks in the year 2014. Accordingly, the C transported to the reservoir from the catchment undergoes aerobic/anaerobic degradation to release greenhouse gases to atmosphere. The study also reveals that oak, devdar, and fir have greater C sequestration capacity compared to other species found in BRF.

Fertilization intensifies drought stress: Water use and stomatal conductance of Pinus taeda in a midrotation fertilization and throughfall reduction experiment

While mid-rotation fertilization increases productivity in many southern pine forests, it remains unclear what impact such management may have on stand water use. We examined the impact of nutrient and water availability on stem volume, leaf area, transpiration per unit ground area (EC) and canopy conductance per unit leaf area (GS) of a pine plantation during its 8th and 9th growing seasons. Treatments consisted of a factorial combination of throughfall reduction (30% reduction in throughfall versus ambient) and fertilization (a complete suite of essential nutrients) beginning in April 2012. Overall, our results indicate that despite unusually high rainfall in the study period and a lack of leaf area index (LAI) response, both EC and GS decreased in response to fertilization and throughfall reduction. Fertilization increased stem volume increment 21% in 2013. Treatment differences were greatest in the growing season of 2013, when EC was on average 19%, 13% and 29% lower in the throughfall reduction (D), fertilization (F) and combined treatment (FD) than the control (C), respectively. The responses of GS to volumetric soil water content (VWC) indicate that lower EC in F was associated with a decrease relative to C in GS at high VWC. Decreases of GS in D relative to C were associated with lower VWC, but little change in the response of GS to VWC. Decreases observed in FD resulted from a combination of these two factors. The pattern of GS responses in the different treatments suggests that structural or physiological changes underlie this fertilization response, possibly in fine root area or hydraulic conductivity. In the short term, this led to large increases in the water use efficiency of stem production, which could suggest greater resiliency to minor water stress. However, impacts on long-term sensitivity to drought remain a concern, as the EC reduction triggered by the fertilization treatment was of comparable magnitude to the 30% throughfall exclusion treatment and the greatest reductions were found in the combined treatment.

Willow Root Development and Morphology Changes Under Different Irrigation and Fertilization Regimes in a Vegetation Filter

Determining the appropriate wastewater dose to supply nutrients while avoiding overirrigation is essential for achieving more balanced above- and belowground development of the plants, thereby ensuring high filtering functionality of a willow vegetation filter system. Aboveground biomass and root development of a 3-year-old willow vegetation filter in response to three increasing doses of secondary-treated urban wastewater under both fertilized and unfertilized conditions were evaluated. Fine, coarse and total root biomass, fine root length density (FRLD), specific fine root length (SFRL) and specific root area (SRA) were assessed in soil core samples collected at four different depths (0–20, 20–40, 40–60 and 60–80 cm) in all treatments. While aboveground biomass increased as wastewater amounts and fertilization increased (from 23.89 Mg ha−1 in non-irrigated condition to 47.07 Mg ha−1 in the highest wastewater dose, from 31.5 Mg ha−1 in unfertilized plots to 40.1 Mg ha−1 in the fertilized), total and fine root biomass decreased when willow was irrigated with the highest wastewater dose (i.e. total root biomass is 2.54 Mg ha−1 in non-irrigated condition—1.36 Mg ha−1 in the highest wastewater dose). The root biomass did not follow the same development pattern, and a significant decrease in fine root biomass was observed when the highest irrigation dose was applied. In these circumstances, we believe that the supply in water and nutrient is sufficient to the development of the willows which react by diminishing the amount of roots proportionally to the aboveground parts. This may lead to a higher transpiration rate and allow willows to avoid a surplus of water in the soil.

The temporal response to drought in a Mediterranean evergreen tree: comparing a regional precipitation gradient and a throughfall exclusion experiment

Like many midlatitude ecosystems, Mediterranean forests will suffer longer and more intense droughts with the ongoing climate change. The responses to drought in long-lived trees differ depending on the time scale considered, and short-term responses are currently better understood than longer term acclimation. We assessed the temporal changes in trees facing a chronic reduction in water availability by comparing leaf-scale physiological traits, branch-scale hydraulic traits, and stand-scale biomass partitioning in the evergreen Quercus ilex across a regional precipitation gradient (long-term changes) and in a partial throughfall exclusion experiment (TEE, medium term changes). At the leaf scale, gas exchange, mass per unit area and nitrogen concentration showed homeostatic responses to drought as they did not change among the sites of the precipitation gradient or in the experimental treatments of the TEE. A similar homeostatic response was observed for the xylem vulnerability to cavitation at the branch scale. In contrast, the ratio of leaf area over sapwood area (LA/SA) in young branches exhibited a transient response to drought because it decreased in response to the TEE the first 4years of treatment, but did not change among the sites of the gradient. At the stand scale, leaf area index (LAI) decreased, and the ratios of stem SA to LAI and of fine root area to LAI both increased in trees subjected to throughfall exclusion and from the wettest to the driest site of the gradient. Taken together, these results suggest that acclimation to chronic drought in long-lived Q. ilex is mediated by changes in hydraulic allometry that shift progressively from low (branch) to high (stand) organizational levels, and act to maintain the leaf water potential within the range of xylem hydraulic function and leaf photosynthetic assimilation.

Fine root dynamics along an elevational gradient in tropical Amazonian and Andean forests

AbstractThe key role of tropical forest belowground carbon stocks and fluxes is well recognised as one of the main components of the terrestrial ecosystem carbon cycle. This study presents the first detailed investigation of spatial and temporal patterns of fine root stocks and fluxes in tropical forests along an elevational gradient, ranging from the Peruvian Andes (3020 m) to lowland Amazonia (194 m), with mean annual temperatures of 11.8°C to 26.4 °C and annual rainfall values of 1900 to 1560 mm yr‐1, respectively. Specifically, we analyse abiotic parameters controlling fine root dynamics, fine root growth characteristics, and seasonality of net primary productivity along the elevation gradient. Root and soil carbon stocks were measured by means of soil cores, and fine root productivity was recorded using rhizotron chambers and ingrowth cores. We find that mean annual fine root below ground net primary productivity in the montane forests (0–30 cm depth) ranged between 4.27±0.56 Mg C ha‐1 yr‐1 (1855 m) and 1.72±0.87 Mg C ha‐1 yr‐1 (3020 m). These values include a correction for finest roots (&lt;0.6 mm diameter), which we suspect are under sampled, resulting in an underestimation of fine roots by up to 31% in current ingrowth core counting methods. We investigate the spatial and seasonal variation of fine root dynamics using soil depth profiles and an analysis of seasonal amplitude along the elevation gradient. We report a stronger seasonality of NPPFineRoot within the cloud immersion zone, most likely synchronised to seasonality of solar radiation. Finally, we provide the first insights into root growth characteristics along a tropical elevation transect: fine root area and fine root length increase significantly in the montane cloud forest. These insights into belowground carbon dynamics of tropical lowland and montane forests have significant implications for our understanding of the global tropical forest carbon cycle.

Differences in fine root traits between early and late-successional tree species in a Chinese subtropical forest

The aim of this study was to compare fine root (≤2 mm diameter) traits (i.e. biomass distribution and architecture) of three tree species (Alniphyllum fortunei, Liquidambar formosana and Cyclobalanopsis glauca) growing in a mixed-species stand in a subtropical forest. Fine root samples were collected using soil cores. The collected samples were scanned with the Win-RHIZO system to analyse architectural parameters and were then ovendried to determine dry mass. Fine roots of the three species were mainly distributed in the top 15 cm of soil and decreased with soil depth across all horizontal distances. C. glauca had the highest fine root biomass at 15‐ 30 cm depth at 1.0 m from the tree trunk. The specific fine root area (SRA) and the length (SRL) were the highest for C. glauca, followed by L. formosana and A. fortunei. These species use different soil exploitation strategies. The early-successional species (A. fortunei and L. formosana) increase their fine root biomass and length throughhighcarboninvestment,whereaslate-successionalspecies(C.glauca)increasenutrientuptakeefficiency viachangesinfinerootmorphologyandhigherSRAandSRLvalues.Insecondarybroadleavedforestmanagement and mixed plantation establishment, root trait differences among tree species and their effects on belowground competition and species coexistence should be considered.

Estimates and relationships between aboveground and belowground resource exchange surface areas in a Sitka spruce managed forest

Our knowledge of the nature of belowground competition for moisture and nutrients is limited. In this study, we used an earth impedance method to determine the root absorbing area of Sitka spruce (Picea sitchensis (Bong.) Carr.) trees, making measurements in stands of differing density (2-, 4- and 6-m inter-tree spacing). We compared absorbing root area index (RAI(absorbing); based on the impedance measure) with fine root area index (RAI(fine); based on estimates of total surface area of fine roots) and related these results to investment in conductive roots. Root absorbing area was a near-linear function of tree stem diameter at 1.3 m height. At the stand level, RAI(absorbing), which is analogous to and scaled with transpiring leaf area index (maximum stomatal pore area per unit ground area; LAI(transpiring)), increased proportionally with basal area across the three stands. In contrast, RAI(fine) was inversely propotional to basal area. The ratio of RAI(absorbing) to LAI(transpiring) ranged from 7.7 to 17.1, giving an estimate of the relative aboveground versus belowground resource exchange areas. RAI(absorbing) provides a way of characterizing ecosystem functioning as a physiologically meaningful index of belowground absorbing area.

Adaptation of fine roots to annual fertilization and irrigation in a 13-year-old Pinus pinaster stand

Effects of fertilization and irrigation on fine roots and fungal hyphae were studied in 13-year-old maritime pine (Pinus pinaster Aït. in Soland), 7 years after the initiation of the treatments. The fertilization trials consisted of a phosphorus treatment, a complete fertilizer treatment (N, P, K, Ca and Mg), and an unfertilized treatment (control). Fertilizers were applied annually and were adjusted according to foliar target values. Two irrigation regimes (no irrigation and irrigation of a set amount each day) were applied from May to October. Root samples to depths of 120 cm were collected in summer of 2005, and the biomass of small roots (diameter 2-20 mm) and fine roots (diameter </= 2 mm) and fine root morphology were assessed. Biomass and length of hyphae were studied by a mesh ingrowth bag technique. Total fine root biomass in the litter and in the 0-120 cm soil profile ranged between 111 and 296 g m(-2). Results derived from the measurements of biomass and root length, or root area, showed that both fertilizer treatments reduced the size of the fine root system, especially in the top soil layers, but did not affect small roots. Compared with control treatments, fine root morphology was affected by both fertilizer treatments with the fine roots having increased specific root length/area, and irrigation tended to reinforce this finer morphology. The amount of hyphae in the mesh ingrowth bags was higher in the fertilization and irrigation treatments than in the controls, suggesting further extension of the root system (ectomycorrhizal infection) and thus of the uptake system. Irrigation had no significant effect on the size of the fine root system, but resulted in a shallower rooting system. Total root to shoot ratios were unaffected by the treatments, but fine root mass:needle mass and fine root area index:leaf area index ratios decreased with increasing nutrient supply. Overall, compared with the control fine roots, increased nutrient supply resulted in a lower fine root biomass but the dynamic fraction of the finest roots was greater. Irrigation had only limited effects on fine root size, distribution and morphology.

A comparison of fine root distribution and water consumption of mature Caragana korshinkii Kom grown in two soils in a semiarid region, China

Water is a key limiting factor for vegetation restoration in the semi-arid areas of China. Caragana korshinkii Kom is a shrub that is widely planted in this region to control soil erosion and land desertification. The objective of this study was to investigate the fine root distribution of mature C. korshinkii and its water consumption, when grown in either silt loam or sandy soils, in order to understand differences between the water cycles of two such soils found in the transition zone between fertile loess hills and desert of the Northern Loess Plateau. Fine root distributions were measured using the trench-profile method. Soil water dynamics were monitored with a neutron probe during two growing seasons. The results showed that fine root area density (FRAD) declined with increasing soil depth in both soils, with 70.7% and 96.6% of the total fine roots being concentrated in the upper 1-m layer of the silt loam and sandy soils, respectively. Water consumption by C. korshinkii in the silt loam was close to that in the sandy soil. Most water consumption in both soil types was from the upper 1-m layer. Little variation in plant available water (PAW) occurred in the 3–6 m soil layer during the whole study period. However, in this layer, the PAW was significantly lower in the silt loam soil than in the sandy soil. Total actual evapotranspiration (ETa) was slightly higher from the sandy soil plots than from those of the silt loam soil during both growing seasons. Our study indicated that mature C. korshinkii effectively uses about the same amount of water from either the silt loam or sandy soils, but that more soil water at depth was extracted from silt loam soil than from sandy soil.