Functional specialization of Eucalyptus fine roots: contrasting potential uptake rates for nitrogen, potassium and calcium tracers at varying soil depths

1. Little is known about the role of deep roots in the nutrition of forest trees and their ability to provide a safety-net service taking up nutrients leached from the topsoil. 2. To address this issue, we studied the potential uptake of N, K and Ca by Eucalyptus grandis trees (6 years of age - 25 m mean height), in Brazil, as a function of soil depth, texture and water content. We injected NO(3)(-)- (15)N, Rb(+) (analogue of K(+)) and Sr(2+) (analogue of Ca(2+)) tracers simultaneously in a solution through plastic tubes at 10, 50, 150 and 300 cm in depth in a sandy and a clayey Ferralsol soil. A complete randomized design was set up with three replicates of paired trees per injection depth and soil type. Recently expanded leaves were sampled at various times after tracer injection in the summer, and the experiment was repeated in the winter. Soil water contents were continuously monitored at the different depths in the two soils. 3. Determination of foliar Rb and Sr concentrations and (15)N atom % made it possible to estimate the relative uptake potential (RUP) of tracer injections from the four soil depths and the specific RUP (SRUP), defined as RUP, per unit of fine root length density in the corresponding soil layer. 4. The highest tracer uptake rates were found in the topsoil, but contrasting RUP distributions were observed for the three tracers. Whilst the RUP was higher for NO(3)(-)- (15)N than for Rb(+) and Sr(2+) in the upper 50 cm of soil, the highest SRUP values for Sr(2+) and Rb(+) were found at a depth of 300 cm in the sandy soil, as well as in the clayey soil when gravitational solutions reached that depth. 5. Our results suggest that the fine roots of E. grandis trees exhibit contrasting potential uptake rates with depth depending on the nutrient. This functional specialization of roots might contribute to the high growth rates of E. grandis trees, efficiently providing the large amounts of nutrients required throughout the development of these fast-growing plantations.

Morphology, biomass and nutrient status of fine roots of Scots pine (Pinus sylvestris) as influenced by seasonal fluctuations in soil moisture and soil solution chemistry

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

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

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

Moderate drought alters biomass and depth distribution of fine roots in Norway spruce

A rain shelter experiment was conducted in a 90-year-old Norway spruce stand, in the Kysucké Beskydy Mts (Slovakia). Three rain shelters were constructed in the stand to prevent the rainfall from reaching the soil and to reduce water availability in the rhizosphere. Fine root biomass and necromass were repeatedly measured throughout a growing season by soil coring. We established the quantities of fine root biomass (live) and necromass (dead) at soil depths of 0-5, 5-15, 15-25, and 25-35 cm. Significant differences in soil moisture contents between control and drought plots were found in the top 15 cm of soil after 20 weeks of rainfall manipulation (lasting from early June to late October). Our observations show that even relatively light drought decreased total fine root biomass from 272.0 to 242.8 g m-2 and increased the amount of necromass from 79.2 to 101.2 g m-2 in the top 35 cm of soil. Very fine roots, i.e. those with diameter up to 1 mm, were more affected than total fine roots defined as 0-2 mm. The effect of reduced water availability was depth-specific, as a result we observed a modification of vertical distribution of fine roots. More roots in drought treatment were produced in the wetter soil horizons at 25-35 cm depth than at the surface. We conclude that fine and very fine root systems of Norway spruce have the capacity to re-allocate resources to roots at different depths in response to environmental signals, resulting in changes in necromass to biomass ratio.

Dynamics of soil exploration by fine roots down to a depth of 10 m throughout the entire rotation in Eucalyptus grandis plantations

Although highly weathered soils cover considerable areas in tropical regions, little is known about exploration by roots in deep soil layers. Intensively managed Eucalyptus plantations are simple forest ecosystems that can provide an insight into the belowground growth strategy of fast-growing tropical trees. Fast exploration of deep soil layers by eucalypt fine roots may contribute to achieving a gross primary production that is among the highest in the world for forests. Soil exploration by fine roots down to a depth of 10 m was studied throughout the complete cycle in Eucalyptus grandis plantations managed in short rotation. Intersects of fine roots, less than 1 mm in diameter, and medium-sized roots, 1-3 mm in diameter, were counted on trench walls in a chronosequence of 1-, 2-, 3.5-, and 6-year-old plantations on a sandy soil, as well as in an adjacent 6-year-old stand growing in a clayey soil. Two soil profiles were studied down to a depth of 10 m in each stand (down to 6 m at ages 1 and 2 years) and 4 soil profiles down to 1.5-3.0 m deep. The root intersects were counted on 224 m(2) of trench walls in 15 pits. Monitoring the soil water content showed that, after clear cutting, almost all the available water stored down to a depth of 7 m was taken up by tree roots within 1.1 year of planting. The soil space was explored intensively by fine roots down to a depth of 3 m from 1 year after planting, with an increase in anisotropy in the upper layers throughout the rotation. About 60% of fine root intersects were found at a depth of more than 1 m, irrespective of stand age. The root distribution was isotropic in deep soil layers and kriged maps showed fine root clumping. A considerable volume of soil was explored by fine roots in eucalypt plantations on deep tropical soils, which might prevent water and nutrient losses by deep drainage after canopy closure and contribute to maximizing resource uses.

Mean age of carbon in fine roots from temperate forests and grasslands with different management

Fine roots are the most dynamic portion of a plant's root system and a major source of soil organic matter. By altering plant species diversity and composition, soil conditions and nutrient availability, and consequently belowground allocation and dynamics of root carbon (C) inputs, land-use and management changes may influence organic C storage in terrestrial ecosystems. In three German regions, we measured fine root radiocarbon (14C) content to estimate the mean time since C in root tissues was fixed from the atmosphere in 54 grassland and forest plots with different management and soil conditions. Although root biomass was on average greater in grasslands 5.1 ± 0.8 g (mean ± SE, n = 27) than in forests 3.1 ± 0.5 g (n = 27) (p < 0.05), the mean age of C in fine roots in forests averaged 11.3 ± 1.8 yr and was older and more variable compared to grasslands 1.7 ± 0.4 yr (p < 0.001). We further found that management affects the mean age of fine root C in temperate grasslands mediated by changes in plant species diversity and composition. Fine root mean C age is positively correlated with plant diversity (r = 0.65) and with the number of perennial species (r = 0.77). Fine root mean C age in grasslands was also affected by study region with averages of 0.7 ± 0.1 yr (n = 9) on mostly organic soils in northern Germany and of 1.8 ± 0.3 yr (n = 9) and 2.6 ± 0.3 (n = 9) in central and southern Germany (p < 0.05). This was probably due to differences in soil nutrient contents and soil moisture conditions between study regions, which affected plant species diversity and the presence of perennial species. Our results indicate more long-lived roots or internal redistribution of C in perennial species and suggest linkages between fine root C age and management in grasslands. These findings improve our ability to predict and model belowground C fluxes across broader spatial scales.

Conventional detection methodology is limiting our ability to understand the roles and functions of fine roots

We lack a thorough conceptual and functional understanding of fine roots. Studies that have focused on estimating the quantity of fine roots provide evidence that they dominate overall plant root length. We need a standard procedure to quantify root length/biomass that takes proper account of fine roots. Here we investigated the extent to which root length/biomass may be underestimated using conventional methodology, and examined the technical reasons that could explain such underestimation. Our discussion is based on original X-ray-based measurements and on a literature review spanning more than six decades. We present evidence that root-length recovery depends strongly on the observation scale/spatial resolution at which measurements are carried out; and that observation scales/resolutions adequate for fine root detection have an adverse impact on the processing times required to obtain precise estimates. We conclude that fine roots are the major component of root systems of most (if not all) annual and perennial plants. Hence plant root systems could be much longer, and probably include more biomass, than is widely accepted.

Left to right: Robert Twilley and Nicole Poret. Washing mangrove fine roots to set up a decomposition experiment, Taylor Slough

This material is based upon work supported by the National Science Foundation through the Florida Coastal Everglades Long-Term Ecological Research program under Cooperative Agreements #DBI-0620409 and #DEB-9910514. This image is made available for non-commercial or educational use only.

Nicole Poret weighing mangrove fine roots to set up a decomposition experiment, Shark River Slough

This material is based upon work supported by the National Science Foundation through the Florida Coastal Everglades Long-Term Ecological Research program under Cooperative Agreements #DBI-0620409 and #DEB-9910514. This image is made available for non-commercial or educational use only.

Left to right: Nicole Poret and Carlos Coronado-Molina deploying bags to determine decomposition rates of mangrove fine roots, Taylor Slough

This material is based upon work supported by the National Science Foundation through the Florida Coastal Everglades Long-Term Ecological Research program under Cooperative Agreements #DBI-0620409 and #DEB-9910514. This image is made available for non-commercial or educational use only.

Glycine metabolism by plant roots and its occurrence in Australian plant communities

Soluble organic nitrogen, including protein and amino acids, was found to be a ubiquitous form of soil N in diverse Australian environments. Fine roots of species representative of these environments were found to be active in the metabolism of glycine. The ability to incorporate [N-15]glycine was widespread among plant species from subantarctic to tropical communities. In species from subantarctic herbfield, subtropical coral cay, subtropical rainforest and wet heathland, [N-15]glycine incorporation ranged from 26 to 45% of (NH4+)-N-15 incorporation and was 2- to 3-fold greater than (NO3-)-N-15 incorporation. Most semiarid mulga and tropical savanna woodland species incorporated [N-15]glycine and (NO3-)-N-15 in similar amounts, 18-26% of (NH4+)-N-15 incorporation. We conclude that the potential to utilise amino acids as N sources is of widespread occurrence in plant communities and is not restricted to those from low temperature regimes or where N mineralisation is limited. Seedlings of Hakea (Proteaceae) were shown to metabolise glycine, with a rapid transfer of N-15 from glycine to serine and other amino compounds. The ability to take up and metabolise glycine was unaffected by the presence of equimolar concentrations of NO3- and NH4+. Isonicotinic acid hydrazide (INH) did not inhibit the transfer of N-15-label from glycine to serine indicating that serine hydroxymethyltransferase was not active in glycine catabolism. In contrast aminooxyacetate (AOA) strongly inhibited transfer of N-15 from glycine to serine and labelling of other amino compounds, suggesting that glycine is metabolised in roots and cluster roots of Hakea via an aminotransferase.

Fine-root production in two secondary forest sites with distinct ages in Eastern Amazon

The objective of this work was to assess the fine-root (≤ 2 mm diameter) production dynamics of two forest regrowths at different ages. Fine-root production was monitored by the ingrowth core method in one 18-year-old site (2 ha) and one 10-year-old site (0.5 ha), both localized in the Apeú region, Northern Pará State, Brazil. The sites were abandoned after successive shifting cultivation, beginning in 1940. Monthly production of live fine-root was similar between sites and was influenced by rainfall seasonality, with higher production during the dry season than the wet season for mass and length. However, mortality in terms of mass was higher in the 10-year-old site than in the 18-year-old site. The seasonality influenced mortality only in the 18-year old site following the pattern observed for live fine-root. The influence seasonal on mortality in terms of length was different between sites, with higher mortality during the wet season in the 10-year-old site and higher mortality during the dry season in the 18-year-old site. Specific root length was higher during the wet season and at the 10-year-old site. Fine-root production was not influenced by the chronosequence of the sites studied, probably fine-root production may have already stabilized in the sites or it depended more on climate and soil conditions. The production of fine-roots mass and length were indicators that generally showed the same pattern.

Potassium supply to cotton roots as affected by potassium fertilization and liming

Cotton (Gossypium hirsutum) is known to have a high requirement for K and to be very sensitive to low soil pH. Most of K reaches plant roots by diffusion in the soil. As K interacts with Ca and Mg, liming can interfere in K movement in the soil, affecting eventually the plant nutrition. The objective of this work was to study the effect of dolomitic lime and 0, 15, 30, 45 and 60 g kg-1 of K on the supply of K to cotton roots. Cotton plants were grown up to 40 days in 5 L pots containing a Dark Red Latosol (Typic Haplusthox) with 68% and 16% of sand and clay, respectively. There was an increase in dry matter yields and in K accumulation due to K fertilization. Root interception of soil K was also increased by K application, but was not affected by lime. Mass flow and diffusion increased linearly with K levels up to 60 mg kg-1, in pots with lime. In pots without lime the amount of K reaching the roots by diffusion increased up to 45 mg kg-1, but decreased at the highest K level. Accordingly, there was more K reaching the roots through mass flow at the highest K level. This happened because there were more fine roots in pots without lime, at the highest K level. As the roots grew closer, there was a stronger root competition leading to a decrease in the amount of K diffused to cotton roots.